SYNTHESIS AND PHYLOGENETIC COMPARATIVE ANALYSES
OF THE CAUSES AND CONSEQUENCES OF KARYOTYPE
EVOLUTION IN ARTHROPODS
by
HEATH BLACKMON
Presented to the Faculty of the Graduate School of
The University of Texas at Arlington in Partial Fulfillment
of the Requirements
for the Degree of
DOCTOR OF PHILOSOPHY
THE UNIVERSITY OF TEXAS AT ARLINGTON
May 2015
Copyright © by Heath Blackmon 2015
All Rights Reserved
ii
Acknowledgements
I owe a great debt of gratitude to my advisor professor Jeffery Demuth. The
example that he has set has shaped the type of scientist that I strive to be. Jeff has given
me tremendous intelectual freedom to develop my own research interests and has been
a source of sage advice both scientific and personal. I also appreciate the guidance,
insight, and encouragement of professors Esther Betrán, Paul Chippindale, John Fondon,
and Matthew Fujita. I have been fortunate to have an extended group of collaborators
including professors Doris Bachtrog, Nate Hardy, Mark Kirkpatrick, Laura Ross, and
members of the Tree of Sex Consortium who have provided opportunities and
encouragement over the last five years. Three chapters of this dissertation were the
result of collaborative work. My collaborators on Chapter 1 were Laura Ross and Doris
Bachtrog; both were involved in data collection and writing. My collaborators for
Chapters 4 and 5 were Laura Ross (data collection, analysis, and writing) and Nate
Hardy (tree inference and writing).
I am also grateful for the group of graduate students that have helped me in this
phase of my education. I was fortunate to share an office for four years with Eric Watson.
He helped build my appreciation for the work of early evolutionary biologists and was
always willing to listen to me talk about comparative methods or beetle natural history. I
also benefited greatly from my interactions with other graduate students. Richard
Adams, Daren Card, Nicholas Long, James McQuillan, Matthew Mosley, Jacobo ReyesVelasco, David Sanchez, and Drew Schield have become good friends and scientists that
I respect.
My interest in biology began early and was nurtured by my parents and William
Voss the former curator of the Fort Worth Museum of Science and History. While my
parents were not particularly fond of the animals I brought home they always encouraged
iii
my pursuits without hesitation. I also owe thanks to Matthew Flechter who was my
constant companion in collecting and studying every creature we could find. Most
importantly, I owe a most heartfelt thank you to my children Abbigail and Leslie. They
both have spent many hours in coffee shops, labs, offices, and the field as I worked on
the research contained in this dissertation. They have been understanding,
encouaraging and a source of my desire to excel.
February 20, 2015
iv
Abstract
SYNTHESIS AND PHYLOGENETIC COMPARATIVE ANALYSES
OF THE CAUSES AND CONSEQUENCES OF KARYOTYPE
EVOLUTION IN ARTHROPODS
Heath Blackmon, PhD
The University of Texas at Arlington, 2015
Supervising Professor: Jeffery P. Demuth
Cytogenetic data in the form of karyotypes are complex, highly-variable traits that
offer opportunities to detect changes in genome organization, uncover phylogenetic
history, and distinguish cryptic species. However, the synthesis of cytogenetic data
across large taxonomic scales has been rare. Here I report the insights gained through a
synthesis of all available karyotype data from Arthropoda.
The first chapter focuses on two databases that were built to make these data
openly available, and broad insights into the evolution of sex determination and
chromosome number in Arthropoda that the collected data have made possible. This is
followed by four chapters that use these data to address fundamental questions in
evolutionary biology. In chapter two I address the question, why do some clades
frequently lose Y chromosomes while they are rarely lost in others? I propose the fragile
Y hypothesis that suggests meiotic mechanisms are of central importance in explaining
the phylogenetic distribution of Y chromosome loss. In chapter three I address the
question, why do some clades exhibit near stasis in chromosome number while closely
related clades show great variation? Using data from beetles in a comparative
framework I show that when species evolve traits that reduce effective population size
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rates of chromosome number evolution increase dramatically suggesting that observed
differences in chromosome number are underdominant while segregating and are fixed
through random drift in small populations. In chapter four I address the question, does
low chromosome number increase the probability of evolving haplodiploidy? I develop a
novel comparative method to test this long-standing hypothesis first proposed by Bull
(1983). The results indicate that low chromosome number increases the probability of
transitions to haplodiploidy. In chapter five I address the question, do eusocial
Hymenoptera have higher chromosome number than solitary Hymenoptera? Using a
larger dataset than previous studies, I show that there is no support for an absolute
difference in chromosome number of solitary and eusocial Hymenoptera. Instead, I find
that eusocial Hymenoptera have much higher rates of chromosome evolution when
compared to their solitary relatives suggesting variable selection pressure or reduced
effective population size in eusocial hymenoptera. As a body of work these analyses
illustrate that chromosome number and meiotic mechanisms can impact the evolution of
sex determination systems and that the evolution of chromosome number is often
strongly influenced by the traits an organism exhibits.
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Table of Contents
Acknowledgements .............................................................................................................iii
Abstract ............................................................................................................................... v
List of Illustrations .............................................................................................................. xi
List of Tables ......................................................................................................................xii
Chapter 1 Synthesis of cytogenetic data in Insecta. ........................................................... 1
Abstract ........................................................................................................................... 1
Introduction ..................................................................................................................... 1
Methods .......................................................................................................................... 3
Results ............................................................................................................................ 5
Order summaries ........................................................................................................ 5
Chromosome number ............................................................................................... 17
Sex chromosome systems in insects ....................................................................... 19
Male and female heterogamety ................................................................................ 20
Sex-limited chromosome evolution .......................................................................... 21
Complex sex chromosomes ..................................................................................... 24
Haplodiploidy ............................................................................................................ 24
Parthenogenesis ....................................................................................................... 25
Conclusions .............................................................................................................. 26
Chapter 2 Estimating tempo and mode of Y chromosome turnover:
explaining Y chromosome loss with the fragile Y hypothesis ........................................... 27
Abstract ......................................................................................................................... 27
Introduction ................................................................................................................... 28
Forces promoting Y(W)-chromosome degeneration ................................................ 30
Forces promoting Y(W)-chromosome retention ....................................................... 30
vii
Methods ........................................................................................................................ 32
Data collection .......................................................................................................... 32
Comparative analysis ............................................................................................... 37
Results .......................................................................................................................... 40
Discussion .................................................................................................................... 48
Conclusion .................................................................................................................... 53
Chapter 3 Assessing the impact of key ecological and phenotypic transitions
on the rate of karyotype evolution: drift drives the evolution of chromosome
number. ............................................................................................................................. 55
Abstract ......................................................................................................................... 55
Introduction ................................................................................................................... 55
Methods ........................................................................................................................ 59
Data collection .......................................................................................................... 59
Phylogenetic model based analyses ........................................................................ 60
Scaled variance estimates ....................................................................................... 61
Results .......................................................................................................................... 62
Data collection .......................................................................................................... 62
Selection on chromosome number........................................................................... 63
Phylogenetic model based rate estimates................................................................ 64
Estimates based on scaled variance ........................................................................ 67
Discussion .................................................................................................................... 68
The role of mutation.................................................................................................. 70
Comparison with previous work ............................................................................... 70
Conclusion .................................................................................................................... 71
Chapter 4 Genome structure and the origin of haplodiploidy ........................................... 73
viii
Abstract ......................................................................................................................... 73
Introduction ................................................................................................................... 73
Methods ........................................................................................................................ 75
Data collection .......................................................................................................... 75
Phylogenetic reconstruction ..................................................................................... 75
Phylogenetic ANOVA ............................................................................................... 77
Originating condition test .......................................................................................... 77
Validation testing ...................................................................................................... 80
Results .......................................................................................................................... 81
Phylogenetic ANOVA ............................................................................................... 81
Validation testing for originating condition test ......................................................... 81
Chromosome number and haplodiploidy.................................................................. 82
Discussion .................................................................................................................... 83
Chapter 5 Recombination, chromosome number and eusociality in the
Hymenoptera. .................................................................................................................... 85
Abstract ......................................................................................................................... 85
Introduction ................................................................................................................... 85
Methods ........................................................................................................................ 89
Data collection .......................................................................................................... 89
Phylogeny ................................................................................................................. 89
Comparative analysis ............................................................................................... 90
Results .......................................................................................................................... 91
Eusociality-depended differences in chromosome number ..................................... 91
Rates of chromosome number evolution.................................................................. 92
Discussion .................................................................................................................... 93
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Appendix A Cytogenetic data for invertebrates ................................................................ 97
Appendix B Genbank accesion numbers ........................................................................ 352
Appendix C Carabidae trait data ..................................................................................... 367
Appendix D Acari trait data ............................................................................................. 372
Appendix E Hymenoptera trait data ................................................................................ 388
Appendix F R code for ancestral condition test .............................................................. 422
References ...................................................................................................................... 428
Biographical Information ................................................................................................. 447
x
List of Illustrations
Figure 1.1 Distribution of chromosome number across hexapods. .................................. 19
Figure 1.2 Genus level distribution of sex determination characteristics in insects.......... 20
Figure 2.1 Taxonomic instability indices based on 500 maximum likelihood tree
inferences. ......................................................................................................................... 36
Figure 2.2 Models of sex chromosome system transitions. .............................................. 38
Figure 2.3 Cladogram illustrating the available cytogenetic data and distribution of sex
chromosome systems in Coleoptera................................................................................. 42
Figure 2.4 Sex chromosome system transition rate estimates. ........................................ 44
Figure 2.5 Distribution of PPS datasets in the suborder Adephaga. ................................ 46
Figure 2.6 Distribution of PPS datasets in the suborder Polyphaga. ................................ 48
Figure 3.1 Haploid number of autosomes in the two major suborders of Coleoptera. ..... 63
Figure 3.2 Phylogenetic based estimates of the rate of chromosome number
evolution. ........................................................................................................................... 65
Figure 4.1 Four steps in the originating condition test. ..................................................... 79
Figure 4.2 Empirical p-values calculated for simulated datasets. ..................................... 82
Figure 4.3 Originating condition test results...................................................................... 83
Figure 5.1 Analysis of the rate of chromosome number evolution. .................................. 93
xi
List of Tables
Table 1.1 Tree of sex database ontology............................................................................ 4
Table 1.2 Coleoptera karyotype database ontology ........................................................... 5
Table 1.3 Meiotic mechanisms in Coleoptera ..................................................................... 5
Table 1.4 Mean number of autosomes in genera with variation in either the number of sex
chromosomes.................................................................................................................... 23
Table 2.1 Marginal likelihoods and model comparisons for models of sex chromosome
evolution. ........................................................................................................................... 45
Table 2.2 Summary of mammals with achiasmatic X-Y segregation or Y chromosome
losses. ............................................................................................................................... 53
Table 3.1 Distribution of traits likely to effect population size. .......................................... 66
Table 3.2 Genus level phylogenetic and scaled variance based estimates of rate of
chromosome evolution. ..................................................................................................... 68
xii
Chapter 1
Synthesis of cytogenetic data in Insecta.
Abstract
Perhaps the most basic description of a species nuclear genome is the number of
discrete chromosomes it is divided among and any differences between the genomes of males
and females. However, the difficulty of collecting this scattered data means that most karyotype
data have never been analyzed using phylogenetically corrected statistical approaches.
Karyotype records are often found in taxon specific journals or in studies with limited geographic
focus; to remove this barrier to anlaysis we have constructed two databases. The Tree of Sex
database brings together all available chromosome number and sex determination data available
in plants, vertebrates, and invertebrates. The Coleoptera karyotype database contains all
karyotype data available for Coleoptera including meiotic mechanisms. These two databases
provide open access to records for over 20,000 species. In this chapter we focus our discussion
on data available for Hexapoda and the insights it reveals. The distribution of sex determination
data indicate that male heterogamety (XY, XO) is likely the ancestral sex determination method in
insects. Female heterogamety appears to be a derived condition which has arrised 2 to 4 times
in Insecta. Haplodiploidy has one ancient origin in the ancestor of all Hymenoptera and likely
another in the ancestor of all Thysanoptera but many more recent origins in other orders.
Pathenogenesis is present in almost all orders of insects but likely represent multiple recent
origins. Finally, chromosome number varies across two orders of magnitude within insects but
we find little evidence for traditional explanations for this variation.
Introduction
At the highest level genome architecture can be described by the number of
chromosomes an organism has and the type of sex chromosome system present; this information
is described by the α karyotype. This level of genome architecture is important to our
understanding of basic processes in biology. For instance, changes in genome structure have
been shown to play an important role in speciation (Presgraves 2008; Yoshida et al. 2014), gene
1
movement (Betran et al. 2002; Meisel et al. 2009), adaptation (Nguyen et al. 2013; Guerrero and
Kirkpatrick 2014), and sex chromosome evolution (Vicoso and Bachtrog 2013; Blackmon and
Demuth 2014a). Particularly during the 1970’s karyotype data was the focus of intense research
and debate. White’s stasipatric model of speciation invoked changes in the number of
chromosomes fixed through drift in isolated populations that then acted as isolating barriers
between incipient species (White 1978). Later though, theoretical work showed that the
probability of fixing a change that could act as a strong isolating barrier upon secondary contact
was unlikely with effective population sizes that at the time seemed reasonable (Lande 1985).
The development of modern sequencing technology also led to a decline in the number of
researchers actively analyzing karyotype data outside of cytotaxonomic settings.
The development of sequencing technologies and statistical phylogenetics now allow us
to estimate large time calibrated phylogenies, and the development of comparative methods
allows for statistical models of trait evolution. Together these developments provide a
quantitative framework for the analysis of karyotype data and the ability to test many long
standing theories in evolutionary biology. For instance: What forces control the number of
chromosomes present in different clades, and why do some clades exhibit stasis while others are
highly variable? How often do sex chromosomes experience turnover? Is the decay of sexlimited chromosomes (Y or W) inevitable? What governs the extinction of sex-limited
chromosomes?
Insects are a tremendously successful group accounting for a great majority of animal
species on Earth, and can be found in almost all terrestrial and freshwater habitats (Gullan and
Cranston 2010; Mora et al. 2011). This diversity at the taxonomic level in insects is matched by
variation in chromosome number and sex chromosome systems making insects vital in building
our understanding of high-level genome evolution. Additionally there is a long history of
cytogenetic research in entomology providing a rich source of data (Stevens 1906; Nichols 1910;
Metz 1916; Goldsmith 1919; Hughes-Schrader 1924; Bridges 1935). The last broad synthesis of
insect cytogenetics was completed by White approximately four decades ago (1977). At the
2
ordinal level syntheses and compilations have been more common (Blattodea (White 1976),
Coleoptera (Smith and Virkki 1978), Lepidoptera (Robinson 1971), Odonota (Kiauta 1972)).
Cytogenetic studies in the intervening years have produced thousands of additional records that
are scattered among hundreds of journal articles and dissertations often with narrow taxonomic or
geographic focus. This has made it difficult to analyze large-scale patterns of karyotype evolution
or even determine what data are available for a clade. To eliminate this barrier we created the
Coleoptera karyotype database (www.uta.edu/karyodb) that contains 4,797 records, and worked
with the Tree of Sex Consortium (www.treeofsex.org) to build the tree of sex database that
contains 10,985 records for insects. We envision these resources as long-term repositories that
will be regularly updated; and allow open access to data that were previously scattered and often
available only through subscription-based publications. Here we systematically characterize and
catalogue sex determination mechanisms and chromosome number in all orders of insects as
well as describe insights from the broad-scale phylogenetic patterns of the distribution of sex
chromosomes systems and chromosome numbers across insects.
Methods
To build our databases we began by incorporating records from important compilations
(Makino 1951; Robinson 1971; Kiauta 1972; White 1976; Smith and Virkki 1978). Next we
examined all articles citing these compilations and then performed an extensive literature search
using Google Scholar and Web of Science using order and family names in conjunction with the
terms: karyotype, cytotaxonomy, cytogenetic, parthenogenesis, haplodiploidy, polyploidy, sex
chromosomes and chromosome number. Literature searches were performed in English,
Spanish, Portuguese, French, and German. To the extent possible, we reconciled historical
karyotype data with currently accepted taxonomy. The records in the database are drawn from
approximately 453 published records including primary research papers, review papers and
previous compilations in books.
The Tree of Sex database was developed in conjunction with the NESCent traitDB
project (Vision and Cranston 2014). This database uses a standardized ontology (Table 1.1) for
3
all animals. The web interface for this database requires users to select a taxonomic group of
interest and then records can be selected based on any combination of the traits present in the
focal taxonomic group.
Table 1.1 Tree of sex database ontology
Trait
Sexual system1
Genotypic (sex determination)
States
Gonochorous, hermaphrodite, parthenogenetic
Male heterogametic, female heterogametic, genetic sex
determination, polygenic
Karyotype
ZO, ZW, XY, XO, WO, homomorphic, complex XY, complex
ZW
Molecular basis
Dosage, Y dominant, W dominant
Chromosome number (female)2
integer number
Chromosome number (male)2
integer number
Predicted ploidy
1,2,3,4
Haplodiploidy (sex determination)
Arrhenotoky, paternal genome elimination, other
Environmental (sex determination)
TSD, TSD Ia, TSD Ib, TSD II, size, density, pH, ESD, other
Polyfactorial (sex determination)
Yes, no
1 Gonochorous is used to indicate that sexes are separate akin to dioecy in plants.
2 The diploid number is reported for chromosome number.
The Coleoptera Karyotype Database was developed locally and is hosted by the
University of Texas at Arlington. We store the karyotype data in an SQL database that can be
queried using a dynamically updated webpage. Users generate database queries by making
selections at up to three taxonomic levels (suborder, family, or genus) and or selections of up to
three karyotype characters (sex chromosome system, b-chromosomes, and reproductive mode).
Once a user has defined a query, it is used to produce an html table. The website also allows
users to export a comma separated text file of their results for offline analysis. The ontology used
in this database (Table 1.2) maintains conventions that have a long history in Coleoptera
cytogenetics where karyotypes are normally reported as the meioformula that describe not only
the number of autosomes and the sex chromosome system but also the behavior of the sex
chromosomes during meiosis.
4
Table 1.2 Coleoptera karyotype database ontology
Trait
Reproductive mode
Autosomes
Sex chromosome system
B chromosomes
Predicted ploidy
States
Diplodiploid, Haplodiploid,
Parthenogenetic
Integer number
XY, XO, Xyp, Xyr, Xy+, XXY, NeoXY
Integer number
Integer number
Testes squashes are the most common method employed in beetle karyotyping, and the
karyotypes reported are meioformulas. For example, a commonly reported karyotype is “9+Xyp”
which indicates 9 autosomes, an XY sex determination system, and that a small Y chromosome
remains at a distance from the X during meiosis. Most organisms, require homologous
chromosomes to come together and form chiasmata during the first meiotic division to faithfully
segregate the chromosomes into the gametes. However, many beetles particularly in the
suborder Polyphaga exhibit various forms of non-chiasmatic segregation (Table 1.3).
Table 1.3 Meiotic mechanisms in Coleoptera
Type of meiosis
Chiasmatic
Sex chromosome
systems
XY, NeoXY, XXY
Achiasmatic
XY, NeoXY, XXY
Asynaptic
Xyp, Xyr, Xy+
Description
Canonical form of meiosis where homologous
chromosomes pair and crossover occurs allowing for
recombination between maternal and paternal copies of
each chromosome.
Meiosis in females is chiasmatic but in males all
chromosomes tightly pair (synapse) but crossover and
recombination are absent.
Meiosis of autosomes and sex chromosomes is
chiasmatic in females. However in males autosomes
have chiasmatic meiosis but the sex chromosomes
though paired remain visibly separated with no
opportunity for chiasmata or recombination.
Results
Order summaries
Below we briefly discuss the data available and the distribution of chromosome number
and sex chromosome systems in each insect order. For completeness we also include the
Entognatha (Collembola, Diplura and Protura) wingless arthropods, which, together with insects,
5
make up the subphylum Hexapoda. All records included in the tree of sex database and the
coleoptera karyotype database are included in Appendix A.
Collembola: There are 8000 described species of springtail (Cicconardi et al. 2013)
across four separate suborders (considered as distinct orders by some authors). Reproduction is
sexual in most species although parthenogenesis is described from 21 species. Species from
three out of the four suborders of Collembola are male heterogametic with either XY or XO sex
chromosome karyotypes. Members of the order Symphypleona, however, are characterized by
paternal genome elimination: Both sexes develop from fertilized eggs with a X1X2/X1X2 sex
chromosome karyotype, but during early development two X-chromosomes are eliminated in
males rendering them X1X2OO, which is followed by the elimination of the rest of the paternal
genome from the male germline later in development (Dallai et al. 1999; Dallai et al. 2000).
Diploid chromosome numbers across the four suborders ranges from 8 to 22.
Diplura: Two-pronged bristletails contain approximately 1,000 described species, and
ecological studies have revealed that many species reproduce sexually, and in some groups
females even guard their eggs. Unfortunately, we have been unable to find any cytological
investigations revealing the presence or absence of sex chromosomes or chromosome numbers
in this lineage.
Protura: Approximately 700 species of these small primarily soil dwelling hexapods,
known as coneheads, have been described. Chromosomal sex determination of the XY type has
been identified in three Italian taxa from the families Acerentomidae and Eosentomidae (Fratello
and Sabatini 1989). Records for the species Eosentomon transitorium indicate chromosome
numbers ranging from 12-20 and both homomorphic and heteromorphic XY sex chromosomes,
this variation is likely due to multiple cryptic species within E. transitorium.
Microcoryphia (Archaeognatha): Since their divergence ~250 million years ago Jumping
bristletails are among the least morphologically changed insects, and approximately 500 species
have been described. The method of sex determination has not been identified and cytological
6
data are limited to the diploid chromosome number of 32 in Machilis noctis and 30 in Dilta littoralis
(Bach and Petitpierre 1978). Reproduction by parthenogenesis has been identified in 5 taxa.
Zygentoma: The silverfish are the sister group of all other insects and have approximately
300 extant species. Cytogenetic data are available for four species and reveal an XO sex
chromosome karyotype with diploid chromosome number of 34 in three taxa and 58 in one
(Makino 1951). Two additional species have been reported to reproduce parthenogenetically
which belong to two separate families (Ateluridae and Nicoletiidae) (Molero-Baltanás et al. 1998).
Ephemeroptera: Approximately 3,000 species of mayflies have been described, and
cytogenetic data are available for 19 species. XY sex chromosome karyotypes have been found
in six species from five genera, and two species have XO sex chromosome karyotypes (Kiauta
and Mol 1977). Eleven species belonging to eight genera reproduce parthenogenetically (Gibbs
1977). Diploid chromosome number in this group ranges from 10 (which is found in three species
of the family Baetidae) to 20 in Ecdyonurus dispar.
Odonata: The dragonflies and damselflies contain approximately 5,500 described
species, and have a long history of cytogenetic studies. Data are available for over 400 species
representing 149 genera. 400 taxa have XO sex chromosome karyotypes and this is widely
accepted as the ancestral state for the order (Kiauta and Mol 1977). XY sex chromosome
karyotypes have been observed in 20 species, which are thought to originate from fusions
between autosomes and the ancestral X chromosome. Such fusions should result in a reduction
in the number of autosomes of XY species in comparison to closely related XO species. Of the 7
genera having both XO and XY taxa, 4 genera show this expected pattern of lower chromosome
number in XY species, while the other three genera have equal numbers of chromosomes in both
XY and XO species. Ishnura hastate, a North American damselfly that has colonized the Azores
is the only documented instance of parthenogenetic reproduction in the order (Rivera et al. 2005).
Chromosome number ranges from 6 in the dragonfly Macrothemis hemichlora to 30 in an
unidentified damselfly in the genus Mecistogaster.
7
Psocoptera: The Psocoptera or book lice are sister to the sucking lice (the Phthiraptera)
and contain approximately 3,000 species. Sex determination karyotypes are known for species in
23 families, all of which display XO sex chromosome karyotype, with the exceptions of neo-Y
systems in Amphipsocus japonicas and Kolbia quisquiliarum (Golub and Nokkala 2009).
Parthenogenesis has been reported in about 30 species. Like Phthirapteran and Hemipteran
insects, all Psocopteran have holocentric chromosomes. Diploid chromosome number ranges
from 14 in genera Elipsocus and Loensia to 30 in the family Psyllipsocidae.
Phthiraptera: All approximately 5,000 species of lice are ectoparasites on bird and
mammalian hosts. There are published karyotypes for 18 species, which show that both sexes
are diploid, but that with the exception of a single species, no sex chromosomes can be
distinguished (Tombesi and Papeschi 1993). This suggests that in most species sex
chromosomes are either homomorphic or absent, and only Bovicola limbata was found to have
differentiated X and Y chromosomes (Golub and Nokkala 2004). A recent molecular analysis in
the human body louse suggested that this species may have paternal genome elimination (PGE)
where males, although diploid, only transmit their maternal chromosomes to their offspring.
Parthenogenesis has been described in four species. Diploid chromosome number in lice range
from 10 in several species to 16 in the genera Hoplopleura and Polyplax.
Thysanoptera: There are approximately 5,000 described species of thrips. It is generally
assumed that all thrips species are haplodiploid, which would make them the only other
haplodiploid insect order besides Hymenoptera. However, only a small percentage of thrips (24
species) have been studied by cytogenetic methods and as a result, haplodiploidy has been
confirmed only in 2 out of the 8 families (Brito et al. 2010). Like other groups of insects with
haplodiploidy, some thrips display mating systems with high levels of sib-mating and females
appear to have control over the sex ratios they produce (Choe and Crespi 1997).
Parthenogenesis occurs frequently and is described in 59 species. Diploid chromosome number
in thrips ranges from 20 (reported in two families) to 106 in Aptinothrips rutua.
8
Hemiptera: There are approximately 90,000 described species of hemipterans, divided
into four suborders. In three of these the sex determining systems are relatively homogeneous,
mostly of the XO type with occasional occurrence of complex male heterogametic karyotypes and
origins of neo-Y chromosomes. The suborder Sternorrhyncha, which includes white flies, aphids
and scale insects is much more diverse though. Each of these three clades displays a different
and unique set of sex determining systems: White flies are haplodiploid, with males developing
from unfertilized eggs. Most aphids reproduce through cyclic parthenogenesis, where a species
goes through several rounds of parthenogenesis followed by a single generation of sexual
reproduction. All parthenogenetic and sexually produced offspring have a XX sex chromosome
karyotype, and males are produced by random elimination of one of the two X-chromosomes
during early development, resulting in XO males (Wilson et al. 1997). Some aphid species have
lost their sexual life cycle, and reproduce exclusively through parthenogenesis. Scale insects
display one of the most diverse array of sex determination systems. Sex determination in a
number of basal clades is of the XO type, but there have been at least two independent
transitions to systems with haploid males within scale insects. Haplodiploidy evolved in the tribe
Iceryiini, while paternal genome elimination (PGE) evolved in the neococcids and is the most
common mode of reproduction among scale insects (found in approximately 6000 species;
(Gavrilov 2007)). In some species the paternal genome is silenced (heterochromatinized) in
somatic cells and eliminated from the germline, while in others the paternal genome is lost
entirely from all cells during early development (Ross et al. 2010). Finally, a number of species in
the tribe Iceryiini are unique among insects having evolved true hermaphroditism, where
individuals produce both male and female gametes and reproduce through self fertilization (Ross
et al. 2010). Diploid chromosome number ranges widely among hemipterans, ranging from 4 in
some scale insect species of the family Monophlebidae, while the highest is 192 and was
reported in the scale insect Apiomorpha macqueeni.
Blattodea: About 4,500 species of roaches have been described. Cytogenetic data for
over 100 species are available, and all of the sexual species have XO sex chromosome
9
karyotypes. The overwhelming majority of roaches possess a metacentric X chromosome which
may make centric fusions with autosomes unlikely, and might explain the rarity of complex sex
chromosomes in this order (White 1976). Two species that reproduce parthenogenetically are
reported. Chromosome number ranges from 16 in Lophoblatta fissa to 80 in Macropanesthia
rhinoceros.
Isoptera: Long considered an independent order, recent molecular studies indicate that
the termites are actually a highly derived clade that nests within Blattodea as sister to the genus
Cryptocercus (Ware et al. 2008). Approximately 2,600 species of termites have been described.
Termites are eusocial with overlapping generations and multiple castes including soldiers and
sterile workers that care for the young. Unlike the eusocial Hymenoptera, termites are
diplodiploid. Cytogenetic data is available for 83 species representing 4 families and 41 genera.
Sex chromosomes have been identified in 63 taxa, and the most frequently observed sex
chromosome karyotype found in 51 species is X1X2Y1Y2 (Bergamaschi et al. 2007). One species
(Stolotermes victoriensis) has XO sex chromosome karyotype (Luykx 1990). Chromosome
number in Isoptera ranges from a high of 98 in Mastotermes darwiniensis to a low of 30 in
Cryptotermes domesticus.
Mantodea: With about 2,300 described species the mantids are the sister group of
Blattodea and Isoptera, and are the earliest branching order of dictyopterans. With the exception
of one parthenogen all species exhibit male heterogamety. XO sex chromosome karyotypes are
found in approximately 60 percent of the studied species, but complex sex chromosome
complements are also common. Specifically, X1X2Y sex chromosome karyotype has been
documented in approximately 40 species, and a single XY species has also been reported. The
X1X2Y species were suggested to form a monophyletic group and their sex chromosome derived
from a reciprocal translocation between a metacentric autosome and a metacentric X
chromosome, resulting in two X chromosomes both of which have a single arm that
chiasmatically pairs with one of the arms of the ancestral autosomes that became the Y
chromosome. Achiasmatic male meiosis has evolved multiple times within mantids (White 1976).
10
Chromosome numbers in mantids range from a low of 16 found in several groups to a high of 40
in Leptomantis parva and an unidentified species in the genus Humbertiella.
Zoraptera: Approximately 30 species of zorapterans have been described. These
primarily tropical insects live in small colonies of less than 200 individuals. The colonies exhibit a
polygynous mating system with the dominant males responsible for the majority of successful
mating attempts. Zorotypus hubbardi is the only species that has been studied cytogenetically
and it exhibits XY sex chromosome karyotype with a diploid chromosome number of 38
(Kuznetsova et al. 2002).
Orthoptera: The order Orthoptera contains over 20,000 species. The large size of
chromosomes and low diploid number have made Orthoptera an important group for our general
understanding of chromosomes and cytogenetics. XO sex chromosome karyotypes are found in
about 80% of the species, and is considered the ancestral mode of sex determination in this
clade. However, many species within Saltatoria have XY and X1X2Y sex chromosome karyotypes
(Castillo et al. 2010a). Parthenogenesis has been reported in 10 species (Lehmann et al. 2011).
Extensive cytogenetic work on natural populations has revealed many examples of chromosomal
variation within and between species, including inversions, translocations, fusions, fissions, sex
chromosome rearrangements, and supernumerary B-chromosomes (Karamysheva et al. 2011).
Diploid chromosome number in Orthoptera ranges from a low of 8 in Dichroplus silveiraguidoi to a
high of 26 in Conometopus sulcaticollis.
Phasmatodea: There are approximately 3,000 species of stick insects. Data are available
for 144 taxa, 37 of which reproduce parthenogenetically, and 83 of which have identified sex
chromosomes. The majority of stick insects have XO sex chromosome karyotypes which likely is
the ancestral system (68 species, present in 36 of 46 studied genera), and a minority exhibits XY
sex chromosome karyotypes (13 species, 8 genera) (White 1976). One species, Didymuria
violescens, has males with both XO and XY sex chromosome karyotypes. Chromosome number
in this group is highly variable and polyploidy is well documented in parthenogenetic taxa. Mean
chromosome number for all species that have identified sex chromosomes is 36.5, while
11
parthenogenetic species have a mean chromosome number of 49.1. Diploid chromosome
number in stick insects ranges from a low of 22 found in several species to 80 in Sipyloidea
sipylus.
Embiidina (Embioptera): The order of webspinners contains approximately 300 described
species. Data on sex determination are available for 4 taxa from the family Embiidae and 5 from
the family Oligotomidae (White 1976). All studied sexually reproducing Embiidina have XO sex
chromosome karyotypes, and female diploid chromosome numbers range from 20-24. Two
parthenogenetic species have been identified, one of which, Haploembia solieri, occurs as both a
diploid and triploid race.
Notoptera: The order Notoptera unites two small insect groups, Grylloblattodea and
Mantophasmatodea, which have at times been considered independent orders, and together
have approximately 40 extant species. Cytogenetic data are available only for two species of the
family Gryllobattodea, Grylloblatta campodeioformis and Galloisiana nipponensis and both
species have XY sex chromosome karyotypes (White 1976). The diploid chromosome number of
G. nipponensis is 30.
Plecoptera: There are approximately 2,000 species of stoneflies. Cytogenetic data are
available for 16 species from the families Perlidae and Perlodidae, both members of the suborder
Systellognatha. The sex chromosome system has been identified in 11 taxa, and 7 species have
X1X2O and 3 have XO sex chromosome karyotypes, while only one has XY. The available
cytogenetic evidence indicates that the multiple sex chromosome systems in Plecoptera are the
result of fission rather than fusion. For instance, in the genus Perla species with multiple sex
chromosomes have more chromosomes than Perla species with either XY or XO sex
chromosomes (2n=22 and 26 versus 10, 19 and 21) (Matthey and Aubert 1947). This along with
the absence of Y chromosomes in these species suggests that the multiple sex chromosome
systems originated from fissions of the ancestral X chromosome. Diploid number in this group
ranges from 10 to 33.
12
Dermaptera. Approximately 2,000 species of earwigs have been described. Cytogenetic
data from over 50 species are available (White 1976), and all are male heterogametic, with about
half having XY sex chromosomes, and the other half having complex sex chromosomes (X1X2Y
and X1X2X3Y). Two earwig species are XO. The chromosomes of Dermaptera seem to be
holocentric (White 1972). Sex chromosome polymorphism has been documented in Forficula
auricularia, where XY and X1X2Y males coexist within populations, and X1X2Y males have an
additional chromosome, whose origin is unclear (Henderson 1970). Chromosome number varies
from a low of 4 in Hemimerus bouvieri to a high of 30 in Arixenia esau.
Hymenoptera: There are approximately 100,000 species of Hymenoptera and it is
assumed that all of them have a haplodiploid sex determining system, which has been confirmed
in all of the 1,300 species for which karyotype data are available (Consortium 2014).
Chromosome number varies from 1 in Myrmecia croslandi to 57 in Dinoponera lucida.
Coleoptera: Beetles, with 350,000 described species, have been the focus of intense
cytogenetic investigation, and karyotypes have often been used to identify cryptic species and to
resolve phylogenetic relationships (Smith and Virkki 1978). Cytogenetic data for over 4,900 taxa
exist, and all sexually reproducing beetles are male heterogametic with over 3,000 species
possessing heteromorphic XY sex chromosome karyotypes. Of the remaining species, over 700
are XO and more than 100 are asexual (Blackmon and Demuth 2014a). There are at least two
origins of haplodiploidy and one of PGE in beetles (Jordal et al. 2000). Coleoptera consists of 4
extant suborders. The suborder Archostemata has only 42 extant species, two species in this
group have been studied. Distocupes varians has 9 autosomes and XO sex determination, while
Micromalthus debilis, has a diploid chromosome number of 20 and cyclic parthenogenesis,
paedogenesis (reproduction by sexually mature larvae), and haplodiploidy (Normark 2013). The
suborder Myxophaga has approximately 65 species but only Ytu zeus has been investigated.
Diploid number in this species was 20 and the sex chromosomes where distant pairing XY.
Adephaga, the second largest order of beetles, contains approximately 40,000 described species,
and data are available for 1,221 species from 7 families. Chromosome number in Adephaga is
13
lowest in Graphipterus serrator, which has a diploid chromosome number of 8, and highest in
Dixus capito obscuroides, which has 70 chromosomes. Polyphaga is the largest suborder of
beetles and contains over 300,000 described species with XY sex chromosomes by far the most
common (over 1884 species from 43 families). XO sex chromosomes have been recorded in 18
families, and complex sex chromosome systems have been found in 12 families. Diploid
chromosome number ranges from 4 in Chalcolepidius zonatus to 66 in Disonycha bicarinata.
Polyploidy is frequent in parthenogenetic species, and parthenogenesis has been identified in 16
families. True haplodiploidy has evolved at least once in the subfamily Scolytinae, and it is
thought that all Xylobrini (> 1200 taxa) are haplodiploid but this has been investigated only in a
handful of species. Another scolitine not closely related to the tribe Xylobrini, Hypothenemus
hamperi, exhibits functional haplodiploidy in the form of paternal genome elimination (Brun et al.
1995).
Strepsiptera: Twisted-wing insects are a highly derived and enigmatic group of
endoparasitic insects and contain over 500 species. Their phylogenetic placement was debated
for some time, but most complete studies to date seem to indicate a close relationship with
Coleoptera (Misof et al. 2014). Cytogenetic evidence for this group exists for just two species.
The diploid number of Xenos peckii was identified as 16 and in an unidentified species of Xenos
from Brazil, 3 autosomes and an XY sex chromosome karyotype was observed (Ferreira et al.
1984). There are scattered reports of parthenogenesis in this family based on collecting only
females, but the only convincing case of parthenogenesis is in Stichotrema dallatorreanum, a
species that is facultative parthenogenetic with isolated females reproducing for multiple
generations (Kathirithamby et al. 2001).
Neuroptera: With over 5,000 described species, this group has cytogenetic information
for 72 taxa belonging to five families. XY sex determination is dominant in the group and is found
in 70 taxa and in all studied families. The Y chromosome has been lost at least twice, once in the
Sisyrid Climacia areolaris and again in the Mantispid Plega dactyloya (Hughes-Schrader 1975b).
The family Mantispidae also contains a species, Entanoneura phithisica that has a X1X2X3Y1Y2Y3
14
sex chromosome system (Hughes-Schrader 1969). E. phithisica has 7 autosomes, a reduction of
2 when compared to E. limbata, which indicates that it was likely a conversion of two of the dot
like autosomes to produce this multiple sex chromosome system. Diploid chromosome number
in the group ranges from 10 to 26.
Megaloptera: This group has approximately 300 described species, and along with
Raphidioptera was formerly considered part of Neuroptera. Four of these species have been
examined cytogenetically and all have XY sex chromosome karyotypes, and diploid chromosome
numbers range from 9 to 11 (Takeuchi et al. 2002).
Raphidioptera: There are approximately 300 species of extant described snakeflies.
Cytogenetic data are limited to 6 species from the genera Agulla and Rhaphidia, all with XY sex
chromosomes (Hughes-Schrader 1975a). All studied species have had 12 autosomes.
Trichoptera: The caddisflies contain approximately 11,000 extant species, and are the
sister group of Lepidoptera. Sex determination data are limited to 15 taxa, all of which have a ZO
system (Lukhtanov 2000). An additional 6 taxa are likely parthenogenetic (Corbet 1966). Diploid
chromosome number has been identified for 44 species in this group and varies from a low of 12
in Limnephilus affinis to a high of 100 in Agrypnetes crassicornis.
Lepidoptera: There are over 160,000 described butterfly species. Like Trichoptera all
investigated Lepidoptera are female heterogametic suggesting that the ancestor of these two
orders evolved female heterogamety. While there have been many cytogenetic studies in
butterflies, testes squashes cannot reveal the karyotype of the heterogametic sex as commonly
reported in XY groups and most cytogenetic studies have only reported the chromosome number.
The sex chromosome system has been identified in only 40 of the 1,219 studied species, and
Lepidoptera have both ZW and ZO systems. Based on the distribution of ZO within Lepidoptera
as well as its sister group Trichoptera, it is believed that butterflies were ancestrally ZO
(Lukhtanov 2000). ZO sex chromosomes have been identified in 6 families of Lepidoptera, three
of which (Arctiidae, Gelechiidae, and Saturniidae) also have species with W chromosomes.
These ZW sex chromosome systems are usually found with matching reductions in the number of
15
autosomes, lending support to the idea that fusions between the autosomes and the ancestral Z
chromosome are the source of new W chromosomes or even multiple ZW chromosome system.
The genus Samia within Saturniidae offers a particularly striking example with species that have
13, 12, and 11 autosomes having ZO, ZW, and Z1Z2W sex chromosome systems respectively
(Yoshido et al. 2005). Some of these transitions may have also coincided with a transition of the
sex determining mechanism, from an ancestral Z counting state to a dominant feminizing allele
on the W, as is found in Bombyx mori (Kiuchi et al. 2014). Chromosome number in Lepidoptera is
also highly variable with diploid numbers ranging from 14 to 382. Parthenogenesis has been
identified in 16 taxa, 10 of which belong to the family Psychidae.
Diptera: Approximately 125,000 species of Diptera have been described. The ease with
which polytene chromosomes can be visualized has led to a cytogenetic data for over 1,400 taxa
(White 1949). The majority of species (1188) have XY sex chromosome systems, but the family
Tephritidae has evolved ZW sex chromosome systems and several lineages lack heteromorphic
sex chromosomes all together. White (White 1949) proposed a classification of Diptera based on
cytological grounds. Lower Diptera (Nematocera) were divided into four groups. The most
primitive Diptera (superfamily Tipuloidea) are characterized by the presence of chiasmata in both
sexes, and cytologically distinguishable XY sex chromosomes. A second assemblage of families
(including Culicidae, Chironomidae, and Simuliidae) generally lack cytologically distinguishable
XY sex chromosomes, but have retained chiasmata in males. The third group, which includes
Bibionidae and Thaumaleidae, is characterized by heteromorphic XY sex chromosomes but a
lack of chiasmata in males. A fourth cytological group of Nematocera is characterized by a highly
specialized chromosome cycle, the loss of the Y chromosome, and sex is determined by
elimination of the paternal X chromosome, and includes the families Sciaridae and
Cecidomyidae. Most families of Diptera, including Drosophila, fall within the suborder Brachycera
(higher Diptera). All Brachycera appear to lack chiasmata in males, and most species have
heteromorphic XY chromosomes. At least 9 families have evolved parthenogenetic species, with
the greatest concentration of parthenogens in the family Chironomidae (where 25 of the 52
16
studied taxa are parthenogenetic). In contrast to the lability of their sex determination systems,
chromosome number varies less in this group. A number of families have genomes with just 3
autosomes, while Tabanidae has the highest with 12 autosomes. A recent study has
demonstrated that while many Diptera species have male heterogamety, the chromosome that is
sex-linked can differ among families. In particular, the sex chromosome of Drosophila was found
to be autosomal in several outgroup species from different families (Vicoso and Bachtrog 2013;
Blackmon and Demuth 2014b).
Mecoptera: Scorpion or hang flies, with approximately 550 extant species, are the sister
group of Diptera, but have not been widely studied cytogenetically. 13 of the 14 taxa in which sex
chromosomes have been identified exhibited XO sex determination, and one species, Boreus
brumalis, has a X1X2Y sex chromosome complement (Xu et al. 2013). Achiasmatic male meiosis
appears to have evolved independently in different species (Ullerich 1961). The number of
autosomes in Mecoptera ranges from 11 to 28.
Siphonaptera: Fleas have an estimated 2,500 species, and recent molecular evidence
indicates that fleas are actually highly derived Mecopterans most closely related to the family
Borediae (Whiting 2002). Despite their abundance and medical importance, cytogenetic data are
available for only 7 taxa (Thomas 1990). The sex chromosome system has been identified in six
taxa, two of which have XY sex chromosomes and four that have multiple sex chromosomes, a
characteristic that would lend support to the close association of Siphonaptera with Borediae that
also exhibits multiple sex chromosomes. The number of autosomes in this group ranges from a
low of 3 in Xenopsylla prasadi to a high of 10 in Leptopsylla musculi.
Chromosome number
Chromosome numbers differ dramatically among and within insect groups, with some
groups having stable karyotypes, while others are highly variable. Figure 1.1 shows variation in
diploid chromosome numbers across orders of insects, and both the mean as well as the
variance in chromosomes numbers differ dramatically among insect orders. Lepidoptera, for
example, have an average of 60 chromosomes, ranging from 14 to 380. Diptera, on the other
17
hand, only have 22 chromosomes on average, and reported chromosome numbers vary from 12
to 52. Differences in chromosomal mutation rates could contribute to this diversity in chromosome
numbers across taxa, for instance, genome rearrangement rates are an order of magnitude
higher in Lepidoptera than in Diptera (d'Alencon et al., 2010). However, a popular explanation for
this variance is the presence or absence of localized centromeres. While the majority of insects
have monocentric (i.e. acrocentric or metacentric) chromosomes, a significant number of insects,
including for example all Lepidoptera and Hemiptera have holocentric chromosomes. Here
localized centromeres are absent and even highly fragmented chromosome can segregate
successfully during meiosis and might enable more flexible karyotypes and a higher number of
chromosomes (Melters et al., 2012). To evalaute this hypothesis we conducted a phylogenetic
ANOVA. We used an existing phyogeny and the mean chromosome number for each order from
the Tree of Sex database (appendix A) (Trautwein et al. 2012; Ashman et al. 2014). We found no
significant difference with this approach (F=0.26, P-value=0.81).
18
Figure 1.1 Distribution of chromosome number across hexapods.
Diploid chromosome numbers reported in hexapods. Boxes represent the range of 25th to 75th
percentile. Outliers are plotted as individual points. The color of the order names indicates
whether chromosomes are holocentric (red), monocentric (blue), or unknown (gray). The number
of species for which data are available and the size of each order are indicated in parentheses.
Sex chromosome systems in insects
Most insects reproduce sexually (Normark 2003) and almost all insects are gonochoristic,
i.e. individuals are either male or female throughout their life. Hermaphroditism is largely absent
in insects (Jarne and Auld 2006). Throughout the tree of life, sex can be determined by many
different mechanisms (Bachtrog et al. 2014), and insects capture much of this diversity (Figure
1.2), and because of the absence of hermaphroditism none of this diversity can be explained by
19
independent origins of separate sexes, unlike many other groups with diverse sex determination
systems (e.g. fish or Crustaceans).
Figure 1.2 Genus level distribution of sex determination characteristics in insects.
The exterior ring indicates male heterogametic systems followed by female heterogametic,
homomorphic systems, and haplodiploid or paternal genome elimination. The branching
structure is based on taxonomy and the figure incorporates data from 9,067 species representing
2725 genera. Data are available from treeofsex.org.
Male and female heterogamety
The phylogenetic distribution of sex determination systems indicates male heterogamety
is likely the ancestral state of insects. Male heterogamety is found in 77% of investigated
species, and within 223 of 359 families. Sex determination systems where females are the
heterogametic sex are rare in insects, and we find only two transitions to female heterogamety:
20
one at the base leading to the superorder Amphiesmenoptera (Lepidoptera and Trichoptera), and
a second transition in Diptera, within the family Tephriditae. Transitions to female heterogamety
from an ancestral state of male heterogamety might be difficult to achieve, as they result in
offspring that are homozygous for the Y (Bull 1983; Bachtrog et al. 2014). If the X contains
important genes not present on the degenerate Y, such transitions become increasingly difficult
and such constraints may underlie the rarity of female heterogamety in insects. However, in
species groups with homomorphic sex chromosomes such transitions may be easier, and there
might be several instances of undetected female heterogamety, as has been suggested for
example in Chironomidae ((Thompson 1971); but see (Martin and Lee 1984)).
Sex-limited chromosome evolution
Gain and Loss of the sex-limited chromosome is common. Many genera and famillies
have apparent recent Y or W losses, and XO or ZO appears to be the ancestral mode of sex
determination in several insect orders. In particular, parsimony ancestral state reconstruction
indicates that ZO was the ancestral state for the most recent common ancestor of Lepidoptera
and Trichoptera (Lukhtanov 2000). Likewise several other orders were likely ancestrally XO,
including Orthoptera, Odonata, Hemiptera and Blattodea. The frequency at which the sex-limited
chromosome is lost is probably largely driven by its gene content; i.e. in groups where the Y
contains genes necessary for survival or reproduction, it is unlikely to be lost unless these genes
are relocated to another chromosome first. However meiotic mechanism also appear to be
important within groups (see chapter 2 and (Blackmon and Demuth 2014a)). An indication of the
variation in the rate of loss is seen in comparing Diptera and Coleoptera, the species groups for
which we have the most data. Using the taxonomy in our database to control for phylogeny, we
infer a minimum of 70 Y chromosome losses in Coleoptera but only 12 in Diptera. After controlling
for the different number of karyotype records we have in both groups, we still have an excess of
over 30 transitions in Coleoptera. This difference in number of events is matched by a difference
in fraction of XO sex systems, with 4.2% of Diptera records and 23.3% of Coleoptera records
21
indicating XO sex chromosome systems. This suggests that the fly Y chromosome may harbor
more genes important for male function, relative to the beetle Y.
Sex-limited chromosomes can be lost, but also be regained, for example by fusions
between Z or X chromosomes with an autosome or the capture of a former B chromosome. In
fact, karyotype data from orders like Lepidoptera, Orthoptera, and Odonota indicate that sexlimited chromosomes are frequently regained after their loss in an ancestral lineage. Though the
sex chromosomes have been identified in only 40 species of Lepidoptera, we identify at least four
independent origins of W chromosomes, while in Orthoptera and Odonata we observe 12 and 10
independent origins of Y chromosomes, respectively. If fusions between autosomes and sex
chromosomes are the primary source of new sex-limited chromosomes we would expect species
with new sex-limited chromosomes to have fewer autosomes. The most recent common
ancestor of both Lepidoptera and Orthoptera had no sex-limited chromosomes. So these orders
provide an opportunity to test this expectation. The Lepidoptera genus Samia has both XO and
XY species as do the Orthoptera genera Dichroplus, Leiotettix and Scotussa in each of these four
cases the species with sex-limited chromosomes have fewer autosomes strongly suggesting that
fussion are one of the primary sources of new sex-limited chromosomes.
22
Table 1.4 Mean number of autosomes in genera with variation in either the number of sex
chromosomes. The mean number of autosome reported for species with XO/ZO, XY/ZW and
multi XY/ZW systems is shown in their respective columns. Whether the pattern is consistent
with fission or fusions and translocations is indicated in the final two columns.
Order
Genus
XO/ZO
XY/ZW
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Dermaptera
Dermaptera
Dermaptera
Diptera
Diptera
Diptera
Diptera
Diptera
Diptera
Isoptera
Lepidoptera
Lepidoptera
Neuroptera
Orthoptera
Orthoptera
Orthoptera
Plecoptera
Altica
Amystax
Anthonomus
Asphaera
Aulacophora
Botanochara
Calathus
Callosobruchus
Cassida
Chilocorus
Chondrocephalus
Chrysomela
Deporaus
Dermatoxenus
Dermestes
Diapromorpha
Disonycha
Epicauta
Euparius
Heikertingerella
Heilipodus
Heilipus
Hermaeophaga
Homoschema
Laccoptera
Lepidospyris
Monochamus
Omophoita
Opatroides
Otiorhynchus
Pityogenes
Pseudotetracha
Pyrophorus
Scarites
Scepticus
Stolas
Tanymecus
Typophorus
Forficula
Nala
Nesogaster
Anastrepha
Bacha
Hemipyrellia
Hylemya
Sepedon
Toxomerus
Cryptotermes
Orgyia
Samia
Plega
Dichroplus
Leiotettix
Scotussa
Perla
9.0
15.0
18.1
9.0
14.0
6.5
21.4
10.0
5.0
13.0
10.75
11.0
10.5
9.5
10.9
8.5
15.5
9.2
17.5
17.3
9.0
9.1
9.6
14.0
16.0
11.3
9.7
8.0
10.3
22.5
9.2
10.0
9.0
14.0
12.7
7.0
3.0
8.0
10.0
9.5
10.1
9.0
10.0
8.1
11.3
25.0
10.0
11.5
10.0
9.0
11.0
17.5
4.8
3.6
5.0
10.0
5.0
4.0
23.0
11.5
12.0
10.0
9.6
8.5
7.0
4.0
23
Multi
XY/ZW
10.5
10.0
20.0
9.9
27.6
18.8
10.0
9.0
9.0
11.0
17.0
16.0
13.0
8.5
8.0
10.0
20.0
9.0
23.0
8.3
13.0
13.0
7.0
4.0
8.0
10.0
9.0
10.0
9.0
10.0
8.0
10.0
7.0
18.0
10.0
13.0
10.0
9.0
10.3
16.0
9.0
3.5
3.0
4.0
10.0
4.0
4.0
16.4
10.0
11.0
8.0
9.0
5.5
9.0
11.2
Fusion or
Translocation
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
Fission
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Complex sex chromosomes
In some insect orders, such as Dermaptera, Plecoptera and Isoptera, a majority of the
taxa were found to harbor multiple X or Y chromosomes, and complex sex chromosomes are also
common in Mantodea, Coleoptera and Orthoptera. Multiple sex chromosomes can in principle
occur through various chromosomal mutations, including fusions, fissions or translocations.
Fusions are expected to reduce the number of autosomes between closely related species with
complex vs. simple sex chromosomes while fissions and translocations should have no effect on
the number of autosomes. To determine the source of complex sex chromosome systems, we
compared the mean number of autosomes between species with simple vs. complex sex
chromosomes within 56 genera that contain species with both types of sex chromosomes (Table
1.4). In 26 of 56 genera, species with complex sex chromosomes had fewer autosomes than
species with simple sex chromosomes, consistent with fusions generating the complex sex
chromosome system in these groups. However, in 30 of 56 genera, species with complex sex
chromosome systems have equal or more autosomes, indicating that fissions (or translocations)
are likely the source of additional X or Y chromosomes in these groups. This variation indicates
that multiple processes are creating complex sex chromosomes in insects. However, we find that
within orders, often one process dominates. For instance, within Coleoptera 26 genera show
evidence for fission or translocations while only 13 suggest fusions. While holocentric
chromosomes may be more prone to fissions than chromosomes with a defined centromere, we
find no evidence for this effect among genera that we examined. Only two orders in this analysis
have holocentric chromosomes (Dermaptera and Lepidoptera), but all 5 genera included show a
pattern of decreased number of autosomes in species with complex sex chromosomes,
consistent with fusions creating complex sex chromosomes.
Haplodiploidy
Haplodiploidy has evolved multiple times in insects, but losses are rare. Haplodiploidy,
where females develop from fertilized eggs and males from unfertilized ones, has evolved at least
6 times within insects (Normark 2003). Two insect orders, Hymenoptera and Thysanoptera
24
(thrips), are completely haplodiploid, and there are a number of smaller haplodiploid clades within
Coleoptera and Hemiptera. Aside from true haplodiploidy there are several origins of paternal
genome elimination (PGE). Species with PGE display the same transmission genetics as true
haplodiploid species, where males only transmit their maternal genome to their offspring, but in
contrast to haplodiploids, males develop from fertilized eggs and the paternal genome is lost in at
least the germline cells or in some cases is heavily condensed and not passed on to the sperm.
PGE has evolved at least 6 times within insects, once within Coleoptera, Hemiptera, Collembola
(a sistergroup to the insects), Phthiraptera and twice within Diptera. The exact number of species
with PGE is unclear as for most clades only a handful of species have been studied. However our
data suggest that PGE might be present in up to 20,000 species (2%) of insect. Haplodiploidy
and PGE are thought to have evolved as a maternal adaptation that increases the reproductive
value of females: A female’s haploid son will transmit her genes to future generations at twice the
rate of a diploid son (Brown 1964; Bull 1979). However this advantage is countered by the fact
that haploid males will probably be less viable, at least in the early stages of the evolution of
haplodiploidy, which will limit the origination of haplodiploidy (see chapter 4).
Parthenogenesis
Parthenogenesis, where females develop from unfertilized eggs, has evolved many times
across insects. Inclusion of a recently compiled list of parthenogenetic insect species to our
database shows that parthenogenesis is found in 144 families, and 1169 species (Normark, 2014;
Normark, 2003). Thus, while parthenogenesis is relatively rare when considering the total number
of species, it has evolved hundreds or even thousands of times. Hemiptera and Coleoptera
harbor many of the parthenogenetic species reported (325 and 467, respectively). Consistent
with evolutionary predictions that asexual lineages are short-lived, there are few larger clades
(genera, families etc.) that are entirely parthenogenetic, but instead parthenogenesis is found
mostly at lower taxonomic units. A wealth of theory has been published on the relative advantage
of sexual versus asexual reproduction, but much less on the factors that can explain the
phylogenetic distribution parthenogenesis (Ross et al. 2012).
25
Conclusions
Our synthesis of karyotype data provides a first step in the understanding of the evolution
of sex determination systems and chromosome number in one of the most abundant and
economically important groups of organisms on the planet. The taxonomic breadth of our analysis
allows us to make broader and more general inferences than earlier syntheses of insect sex
determination, e.g. (Cook, 2002; Kaiser & Bachtrog, 2010; Verhulst, van de Zande & Beukeboom,
2010).
In general, we find that male heterogamety is likely ancestral for insects, while female
heterogamety has evolved at least twice. Our data indicate that sex-limited chromosomes are
lost and gained more readily in some clades than others, and that some groups (i.e. Dermaptera,
Plecoptera, Mantodea, and Isoptera) exhibit a propensity for complex sex chromosomes not seen
in closely related clades. The origin of complex sex chromosomes appears to differ among
groups with both fussions and fissions playing an important role.
* Portions of this chapter were previously published as:
Tree of Sex Consortium. (2014) Tree of Sex: A database of sexual systems. Scientific
Data 1:140015.
Blackmon, Heath and Jeffery P. Demuth. (2015) Coleoptera karyotype database. The
Coleopterists Bulletin 69(1). In Press
Blackmon, Heath (2014) Coleopterist’s Corner. Entomological Society of Ontario Newsletter 19(2)
18-21.
26
Chapter 2
Estimating tempo and mode of Y chromosome turnover: explaining Y chromosome loss with the
fragile Y hypothesis
Abstract
Chromosomal sex determination is phylogenetically widespread, having arisen
independently in many lineages. Decades of theoretical work provide predictions about sex
chromosome differentiation that are well supported by observations in both XY and ZW
systems. However, the phylogenetic scope of previous work gives us a limited understanding of
the pace of sex chromosome gain and loss, and why Y or W chromosomes are more often lost in
some lineages than others, creating XO or ZO systems. To gain phylogenetic breadth we
therefore assembled a database of 4,724 beetle species’ karyotypes and found substantial
variation in sex chromosome systems. We used the data to estimate rates of Y chromosome
gain and loss across a phylogeny of 1,126 taxa estimated from seven genes. Contrary to our
initial expectations, we find that highly degenerated Y chromosomes of many members of the
suborder Polyphaga are rarely lost, and that cases of Y chromosome loss are strongly associated
with chiasmatic segregation during male meiosis. We propose the "fragile Y" hypothesis; that
recurrent selection to reduce recombination between the X and Y chromosome leads to the
evolution of a small pseudo-autosomal region (PAR), which, in taxa that require XY chiasmata for
proper segregation during meiosis, increases the probability of aneuploid gamete production, with
Y chromosome loss. This hypothesis predicts that taxa that evolve achiasmatic segregation
during male meiosis will rarely lose the Y chromosome. We discuss data from mammals that is
consistent with our prediction.
27
Introduction
Chromosomal sex determination is phylogenetically widespread, having arisen
independently in many lineages. Decades of theoretical work provide predictions about sex
chromosome differentiation that are well supported by observations in both XY and ZW
systems. However, the phylogenetic scope of previous work gives us a limited understanding of
the pace of sex chromosome gain and loss, and why Y or W chromosomes are more often lost in
some lineages than others, creating XO or ZO systems. To gain phylogenetic breadth we
assembled a database of 4,724 beetle species’ karyotypes and found substantial variation in sex
chromosome systems. We used the data to estimate rates of Y chromosome gain and loss
across a phylogeny of 1,126 taxa estimated from seven genes. Contrary to initial expectations,
We find that highly degenerated Y chromosomes of many members of the suborder Polyphaga
are rarely lost, and that cases of Y chromosome loss are strongly associated with chiasmatic
segregation during male meiosis. We propose the "fragile Y" hypothesis; that recurrent selection
to reduce recombination between the X and Y chromosome leads to the evolution of a small
pseudo-autosomal region (PAR), which, in taxa that require XY chiasmata for proper segregation
during meiosis, increases the probability of aneuploid gamete production, with Y chromosome
loss. This hypothesis predicts that taxa that evolve achiasmatic segregation during male meiosis
will rarely lose the Y chromosome. We discuss data from mammals that is consistent with our
prediction.
Chromosomal sex determination has evolved independently in many lineages (Bull
1983). In addition to their role in gender determination, sex chromosomes are fascinating
because the homologs often differ in gene content and morphology (Vallender and Lahn 2004;
Graves 2006; Arunkumar et al. 2009; Betrán et al. 2012). Their unequal distribution between
sexes also means that sex-linked genes experience and respond to evolutionary forces in
different ways compared with autosomes (Charlesworth et al. 1987; Rice 1987; Charlesworth
1991; Rice 1994). The sex chromosomes can encompass the extremes of evolutionary rate.
For example, the average divergence between human and chimpanzee X and Y chromosomes
28
are lower and higher respectively than average autosomal divergence (Mikkelsen et al. 2005).
Sex chromosomes can also play a special role in the origin of species; where the hemizygous sex
often suffers the consequences of hybridization disproportionately (Haldane’s rule)(Haldane
1922; Watson and Demuth 2012) and X-linked introgressions have larger effects on hybrid fitness
than autosomal introgressions (large X-effect)(Presgraves 2008; Phillips and Edmands 2012).
Finally, sex chromosomes that are confined to the heterogametic sex (Y or W in male or female
heterogametic species respectively) are also particularly interesting for their apparent
dispensability in some taxa but not others.
The canonical view of sex chromosome evolution assumes that a sex determining region
evolves that leads to a pair of ancestral autosomes evolving into proto sex chromosomes
(Westergaard 1958). Most models suggest that the resulting proto Y (W) will degenerate as a
consequence of reduced effective population size (as these chromosomes are only found in one
sex) and lack of recombination near the sex-determining locus. The non-recombining region can
expand to adjacent portions of the chromosome. The selective force for this is thought to be
selection to maintain linkage between sexually antagonistic loci (those polymorphic for alleles that
benefit one sex at the expense of the other) and the sex determination locus. Recombination
suppression may involve chromosomal rearrangements (e.g. inversions) that include the sex
determining locus (Charlesworth et al. 2005). Once recombination is suppressed, the Y(W)
chromosome is subject to evolutionary forces that are expected to lead to loss of the
chromosome’s genes (Charlesworth and Charlesworth 2000). The phylogenetically widespread
observation of XO (ZO) species (Makino 1951) indicates that degeneration of the Y(W) may
ultimately result in its complete loss; yet despite considerable work on the molecular evolution of
particular Y chromosomes (Lahn et al. 2001; Bachtrog et al. 2008; Hughes et al. 2012) we still
have a relatively poor understanding of the factors that govern the rates of Y(W) chromosome
gain and loss.
29
Forces promoting Y(W)-chromosome degeneration
In principle, the forces responsible for decay of these chromosomes include: Muller’s
Ratchet, background selection, Hill-Robertson effect, and genetic hitchhiking (Bachtrog 2013).
The predicted inevitable decay of Y and W chromosomes has led to the idea that they are “born
to be destroyed” (Steinemann and Steinemann 2005) and indeed these chromosomes are often
dispensable (e.g. Lepidoptera, (Traut et al. 2008); Nematodes, (Bull 1983); Orthoptera, (Castillo
et al. 2010b); and Odonata, (Kiauta 1969)). Some groups, such as Coleoptera and Diptera
exhibit multiple independent losses of the Y chromosome (White 1977). In Drosophila, the
tenuous persistence of the Y chromosome is evident in that the ancestral Y was likely lost long
ago in an ancestor of D. melanogaster while the current Y is likely a secondarily captured B
chromosome (Carvalho and Clark 2005). In fact, recent analysis indicates that the ancestral Y
may have been lost as part of a sex chromosome reversal where a formerly autosomal pair of
chromosomes became the determinants of sex allowing the ancestral X to be fixed in the
Drosophila lineage as an autosome (the dot chromosome) (Vicoso and Bachtrog 2013). In D.
pseudoobscura the existing Y is homologous with an ancestral autosome, suggesting that the sex
chromosomes fused with an autosome and the ancestral Y region was subsequently lost
(Carvalho and Clark 2005). Even among taxa with generally persistent XY chromosome systems
such as those in mammals, there is precedent for Y dispensability; both mole voles (Just et al.
1995) and spiny rats (Arakawa et al. 2002) have lost the ancestral Y chromosome.
Forces promoting Y(W)-chromosome retention
Several lines of evidence , however, suggest that the evolution of Y(W) chromosomes is
more complex than just inevitable decay. For instance, frequent turnover in the sex determining
chromosome (i.e. changes in the linkage group responsible for sex determination, so that a
chromosome is not involved long enough for gene loss to occur) and/or intermittent
recombination between sex chromosomes may play a role in persistence of the homomorphic sex
chromosomes observed among most amphibians and fish (Stein et al. 2002; Woram et al. 2003;
Van Doorn and Kirkpatrick 2007; Perrin 2009; Blaser et al. 2012; Guerrero et al. 2012). Sex30
specific gene regulation may ameliorate situations with sexually antagonistic polymorphisms
(Prince et al. 2010) and may further contribute to retention of old homomorphic sex
chromosomes, as recently suggested for the emu (Vicoso et al. 2013).
In systems that retain the Y(W) chromosome despite considerable degeneration,
selection may prevent complete gene loss and/or promote recruitment of genes from elsewhere
in the genome. For example, degeneration of the human Y-chromosome occurred in 5 waves
over 200-300 million years of mammalian evolution (Hughes et al. 2012). Linear extrapolation,
using the average rate of gene loss, predicts that the human Y would be lost within 10 million
years (Aitken and Graves 2002), however as the number of sites decline so should the rate at
which genes are lost (Bachtrog 2008). Recent analyses show that a few genes have been
conserved due to purifying selection (Hughes et al. 2012), and that new genes that are important
for male fertility have been transferred to the Y (Lahn et al. 2001). Retention of these “essential”
male genes is aided by their frequent occurrence in palindromes where intra-chromosomal gene
conversion decreases the chance of loss and may also foster fixation of new genes by adaptive
evolution (Betrán et al. 2012).
The strength of selection to retain Y(W)-linked genes should also be affected by the
evolution of dosage compensation. If X(Z)-linked genes are expressed at low levels in males, this
may lower males’ fitness, and purifying selection will then act against loss of Y(W) homologs
unless dosage compensation evolves (Ohno 1967). While chromosome-wide (global) dosage
compensation is the norm in most mammals and Drosophila, considerable data now shows that it
is incomplete in broad range of animals including trematodes (Vicoso and Bachtrog 2011),
lepidopterans (Harrison et al. 2012), birds (Itoh et al. 2007), fish (Leder et al. 2010), and
monotremes (Deakin et al. 2008). In these groups, the Y or W chromosome should decay more
slowly since loss of function mutations will not be masked by increased expression of the X or Z
copy.
31
Sex chromosome evolution in Coleoptera
The model systems for studying Y chromosome evolution, Drosophila and mammals, are
ill suited to explore hypotheses about the tempo and mode of Y chromosome turnover because
there are few transitions among sex chromosome states. Here we use comparative methods in
the order Coleoptera to explore the evolution of Y chromosomes and generate hypotheses.
Beetles are the most speciose order of eukaryotes and we have compiled karyotype data for
thousands of species (available at www.uta.edu/karyodb).
To analyze sex chromosome changes in a comparative framework, we use DNA
sequences for over 1,000 species in our karyotype database to estimate the phylogeny of
Coleoptera. There have been few explicitly phylogenetic analyses of karyotype data (Flores et al.
2008; Leache and Sites 2009; Henning et al. 2011; Maddison and Leduc-Robert 2013), and to
our knowledge, our analysis provides the first estimates of transition rates for sex chromosome
turnover, Y chromosome decay, and Y chromosome loss, over such a large number of species.
We find distinctly different patterns and rates of sex chromosome transitions between the two
main suborders of beetles (Adephaga and Polyphaga). We propose that the much lower rate of
Y chromosome loss in Polyphaga can be explained by the evolution of distance-pairing sex
chromosomes that ensure proper meiotic segregation even when no recombination occurs
between the sex chromosomes.
Methods
Data collection
Karyotypes: We performed a thorough literature search and compiled a comprehensive
record of published Coleoptera karyotypes (appendix A). To the extent possible, we reconciled
historical karyotype data with currently accepted Coleopteran taxonomy (North American species:
(Arnett and Thomas 2000; Arnett et al. 2002); outside of North America: (Beutel and Leschen
2005; Leschen et al. 2010)).
Coleoptera are male heterogametic, and in most beetles the Y chromosome is smaller
than the X (Smith and Virkki 1978). The most common sex chromosome systems in the literature
32
are XY, XO, and Xy+ (see below). Here we denote sex chromosomes that undergo synapsis
during meiosis as XY. In the vast majority of XY taxa, the synaptic chromosomes also form
chiasmata (i.e. contain at least one region that can recombine) (Smith and Virkki 1978).
However, achiasmatic male meiosis - where all chromosomes in males form synapses but do not
recombine - has evolved four times in the suborder Adephaga. Two instances of achiasmatic
male meiosis appear to involve only one or a few species and are probably of recent origin,
(Serrano 1981; Yadav and Burra 1987), while the other two instances are probably old and
appear to be synapomorphies for the clades Trechitae and Cicindelini + Colyrinae (Galian et al.
2002; Maddison and Ober 2011).
Sex chromosome systems that form distance-pairing sex bivalents are denoted as Xy+.
Such X and Y chromosomes are entirely nonrecombining. In these species the autosomes
undergo normal synapsis and crossing over in both sexes and the X chromosomes do so in
females. However, in males the X and Y chromosome pair at a distance with no synapsis and no
opportunity for crossing over. In Xy+ species the Y is usually very small often being described as
“punctiform”. We denote species that have completely lost the Y chromosome as XO.
Sequence data: Sequences for 2 mitochondrial genes (16s and COI) and 5 nuclear
genes (18s, 28s, elongation factor 1, arginine kinase, and wingless) from 1,140 operational
taxonomic units (OTUs) representing members of 47 of the 59 families with karyotype data were
downloaded from GenBank (Appendix B). The karyotype database and the sequences available
from GenBank contain overlapping, but non-identical sets of species, so the sequence data were
treated as follows. When the karyotype database and GenBank had a match at a level higher
than species, we created “chimeric OTUs” In these cases a composite branch was created by
assigning all sequences and sex chromosome states found in that clade to the single higher level
group. In this way we created 280 genus level OTUs and 14 family level OTUs. In order to
increase overlap in the matrix of gene sequences, we also created 18 anchoring OTUs In these
cases relationships among several members of a monophyletic group in the karyotype database
had sequences of a single gene, but another member of the taxon (not present in the karyotype
33
database) had sequences for additional target genes. In these cases, an arbitrarily chosen
member of the monophyletic group that was sampled for the single gene and was present in the
karyotype database was assigned sequences for all of the otherwise unrepresented target genes.
This effectively “anchors” the monophyletic group within the larger Coleoptera tree without
impacting resolution within the group. For example, in the genus Curculio we have karyotype
data for three species, but those species only have sequence data available for COI. There is an
additional species (Curculio niveopictus) with sequence data available for the 18S and 28S
genes, but it does not have karyotype data. To increase overlap in our matrix, one of the species
with COI sequence, C. nucum, was arbitrarily chosen to act as the anchor by having the 18s and
28s sequences from C. niveopictus assigned to it. This anchors the genus Curculio within the
larger Coleopteran phylogeny, while preserving resolution within the genus. Both chimeric and
anchoring OTUs are indicated as such in table A1.
All sequences were aligned in MAFFT (Katoh et al. 2009). RNA genes (16s, 18s, and
28s) were then filtered with the program Gblocks to remove ambiguously aligned sites (Talavera
and Castresana 2007). This resulted in alignments for 16s, 18s, and 28s of 544, 1964, and 404
bp in length respectively. We used MEGA to manually adjust the alignments of protein coding
genes (COI, wingless, elongation factor 1, and arginine kinase) to insure that the reading frame
was maintained (Tamura et al. 2011); these alignments were 1567, 585, 1189, and 810 bp in
length respectively. Finally all alignments were checked for poorly aligned taxa using GUIDANCE
(Penn et al. 2010); fourteen taxa were found to have few unambiguously aligned sites and were
removed from our dataset. The alignments for our seven target genes were concatenated into a
sparse supermatrix that contained 1,126 OTUs and was 7,063bp in length. Most taxa do not
have sequences for all genes, and the mean number of alignment sites with information was
1870.
Phylogenetic Inference: Inconsistency in the placement of a small subset of taxa among
trees with equally probable topologies (rogue taxa) is a common problem in phylogenetic
inference from sparse supermatrices (Thomson and Shaffer 2010). The problem is magnified by
34
the computational burden of optimizing over the large number of OTUs in our dataset (e.g. 14,000
CPU hours on CIPRES (Miller et al. 2010) servers to complete the first phase below). Therefore,
we divided our phylogenetic inference into two phases. The first phase used maximum likelihood
inferences to build a collection of trees that we used to assess taxon instability. We computed
500 maximum likelihood trees using RAxML v 7.2.8 (Stamatakis 2009). Based on the resulting
collection of trees, the instability index for all taxa was calculated (Aberer et al. 2013). High index
values indicate that a taxon’s placement is variable among trees. The distribution of instability
indices shown in figure 2.1 indicates that 92 percent of taxa have indices below 2194 but that
above this, indices increase quickly. The 84 taxa with scores above this cutoff were removed
from subsequent analyses, resulting in a dataset containing 744 Polyphaga taxa, 296 Adephaga
taxa, and 2 outgroup taxa.
35
4000
3000
2000
1000
Taxonomic Instability Index
0
200
400
600
800
1000
1200
Taxa
Figure 2.1 Taxonomic instability indices based on 500 maximum likelihood tree inferences. The
dashed line shows the chosen cutoff, an index of 2194. Most taxa, 93 percent fall below this
while above this value instability increases quickly.
The second phase of our phylogenetic inference employed Bayesian methods to produce
a posterior sampling of ultrametric trees. The best maximum likelihood phylogram from the first
phase was converted to an ultrametric tree using nonparametric rate smoothing in the R package
APE (Paradis 2011). The resulting tree was subsequently used as input for two independent
inferences in BEAST (v1.7.5; (Drummond and Rambaut 2007; Suchard and Rambaut 2009)).
We assumed a lognormal relaxed clock and used normal distributions to place priors on the age
of seven nodes. The seven nodes represent the age of the order (Coleoptera = 285 MYA), both
major suborders (Adephaga = 237.2 MYA and Polyphaga = 270.5 MYA) and four arbitrarily
36
chosen clades (Hydradephaga = 219.8 MYA, Elatridae = 139.9 MYA, Brentidae = 137.5 MYA,
and Passalidae = 121.4 MYA). The standard deviations of the priors were set to reflect the 95%
confidence interval of previous estimates (McKenna and Farrell 2009).
The two independent MCMC analyses required approximately 70 million generations to
converge on a parameter space with equal likelihood to insure that they had reached stationarity
they were allowed to run for an additional 40 million generations. The phylogeny inferred from
our sparse supermatrix is largely consistent with the previously most comprehensive family level
analysis for Coleoptera (Hunt et al. 2007). Because of the computational demands of analyzing
evolutionary rates over such large trees, parameter estimates for the evolutionary models below
were marginalized over 100 randomly selected trees from the stationary phase of our two chains
(following referred to as “sampled trees”). The sampled trees had high resolution; a maximum
clade credibility tree exhibited posterior probabilities greater than 90 percent at 76 percent of the
nodes. The 100 sampled trees were subsequently used to model sex chromosome evolution in
Polyphaga and Adephaga and are publicly available from Dryad Digital Repository:
http://dx.doi:10.5061/dryad.g8010/1.
Comparative analysis
Sex chromosome systems are reported as discrete states (e.g. XY, Xy+, XO) despite
representing a fundamentally continuous, though probably not linear, process of differentiation.
In modeling their evolution across a phylogeny we must determine which states to include in the
matrix of transition probabilities? For instance, if Xy+ is biologically equivalent to XY, then
including the additional transition probabilities for that state will only add noise to the inferred rate
of Y chromosome gain and loss. On the other hand, if Xy+ is a distinct state with different rates
of transition to and from the XO state, then allowing for independent transition probabilities should
provide a significantly better fit to the observed data. To assess how many states and rates best
describe Y chromosome evolution in Coleoptera, we estimated transition probabilities under both
2-state (XY/XO) and 3-state (XY/Xy+/XO) models (Figure 2.2). For the 2-state models, all XY
and Xy+ taxa were both coded as XY. Species where literature reports note “NeoXY” and other
37
complex sex chromosome systems (i.e. those with multiple X and/or Y chromosomes) were also
included in the analysis based on whether the sex chromosomes form a synapse during male
meiosis (XY) or not (Xy+).
Figure 2.2 Models of sex chromosome system transitions.
(A) 2-State coding model with taxa partitioned between XO and XY. Using this coding we fit
models with one rate (model 2.1) and two rates (model 2.2). (B) 3-State coding model with taxa
partitioned between XO, XY, and Xy+. Using this coding we fit models with 1, 2, 4 and 6 rate
parameters. Model 3.4 is a constrained model allowing for comparison between 2-state and 3state coding in Polyphaga. If XY and Xy+ are equivalent states model 3.4 and 3.6 should perform
equally well.
Of the 1042 OTUs in our tree 88 taxa do not possess sex chromosome data and were
coded as missing data, and do not affect our estimated rates of Y chromosome changes.
Included in these are 23 parthenogenetic taxa as well as 1 haplodiploid taxon. The remaining 64
taxa have only the chromosome number available, and while homomorphic chromosomes are not
reported in Coleoptera some of these species may have sex chromosomes of this type.
However, in most cases the investigators describe the chromosome squashes as inadequate to
38
resolve the sex chromosomes, therefore it would appear that sex chromosomes in beetles are
rarely in a homomorphic state, and this should not bias our results.
We estimated transition rates using BayesTraits, which allowed us to marginalize over
uncertainty in phylogenetic inference and uncertainty in tip states (Pagel et al. 2004). For the 2state coding we estimated rates assuming that all rates are equal (2.1) and also assuming that all
rates differ (2.2). For the 3-state coding, we again estimated rates assuming that all rates are
equal (3.1), plus a time reversible model (3.3), a 4-rate model (3.4), and a model with all six rates
different (3.6). Finally model 3.4 is a nested version of model 3.6 in which we force both states
XY and Xy+ to have a single rate of transition to XO and a single rate of transition back to XY
from XO; this is equivalent to using 2-state coding for the data. Comparing models 3.4 and 3.6
tests whether XY and Xy+ have significantly different transition rates to and from XO.
To improve computational feasibility of the rate estimates, we first used BayesTraits to
perform a preliminary maximum likelihood analysis of the sex chromosome transition rates across
all sampled trees for the two major suborders, Polyphaga and Adephaga. Since estimated
transition rates were always below 0.05, we conservatively set uniform priors between 0 and 0.1
on all transition rates for subsequent Bayesian analyses. None of our estimates were bounded
by these priors. We adjusted the RateDev parameter for each run to insure that the acceptance
rate of moves was between 20% and 40%. The marginal likelihood of each model was computed
as the harmonic mean of the post burn-in likelihoods across all sampled trees. To compare
models we used the marginal likelihoods (LS) to calculate the log Bayes Factor (LBF): LBF =
2(LS1-LS0) where LS1 is the more complex model and LS0 is the less complex model. We
interpret LBF = 2 to 6 as positive support for the more complex model, 6-10 as strong support,
and >10 as very strong support (Kass and Raftery 1995).
To further assess adequacy of our chosen models we performed posterior predictive
simulations (PPS) (Rubin 1984). PPS datasets were created in the “R” environment (R
Development Core Team 2013) using a custom function available in package evobiR. Briefly, we
extracted rate matrices and associated trees from 1500 random points during the post burn-in
39
phase of the BayesTraits MCMC runs for the best 2-state and 3-state models, each extraction
was used to create a simulated dataset by evolving sex chromosomes over the extracted tree
with the extracted rate estimates. The root state for each simulation was set so that the
distribution of roots across each set of 1500 simulations matched the distribution of root states
inferred during original parameterization of the model. We used these simulated datasets to
compare the frequency of each tip state with the observed data. This same process was
repeated on two subtrees within the suborder Adephaga to test whether specific subgroups had
significantly different patterns and rates of transitions compared to what is expected based on
rates estimated for the full suborder. Transition rates are reported as the mean probability of a
transition per 100 million years ± the standard error.
Results
Coleopteran Karyotypes: Our comprehensive database of Coleoptera karyotypes
includes 4,724 records based on 208 literature sources (Figure 2.3). The database is available
online at www.uta.edu/karyodb and can be interrogated for any combination of: suborder, family,
genus, sex chromosome system, presence of B chromosomes, and/or reproductive mode.
Tables of selected data can be downloaded from the website. Karyotypes follow the format of
Smith and Virkki (1978), the previously most comprehensive compilation of coleopteran
karyotypes. In the Coleoptera cytogenetic literature, distance-pairing sex bivalents are usually
denoted with a lower case letter that describes how they are oriented during meiosis (e.g. “p”
stands for parachute and indicates a large X chromosome with a small Y chromosome that
appears suspended from it; “r” stands for rod and indicates that the X and Y are oriented end to
end.) This format is maintained in our database, but for clarity and consistency with the broader
literature, all distance-pairing sex bivalents are denoted by Xy+ in the present study. Whenever
possible the meioformula is given in the database. For example 9+Xy+ means a haploid
autosome count of 9 and distance-pairing sex chromosomes. The meioformula is not available
for 470 species in the database where only the diploid number is reported in the literature, nor for
9 haplodiploid records, and 116 parthenogenetic records. In total, data on sex chromosomes
40
were available for 4,223 species. Since records were available for only three species in the
relatively small coleopteran suborders Archostemata and Myxophaga, we analyzed only the
major suborders Adephaga and Polyphaga.
41
Figure 2.3 Cladogram illustrating the available cytogenetic data and distribution of sex
chromosome systems in Coleoptera. Number of species in the karyotype database in each sex
chromosome (or sex determination) state. Data and references are available at
www.uta.edu/karyodb. a The number of species with chromosome number available
b Sex chromosome systems with multiple X and or Y chromosomes
c Species with parthenogenetic reproduction
d Species with haplodiploidy sex determination
42
There is a striking difference between Adephaga and Polyphaga in the number of taxa
with distance-pairing sex bivalents (Xy+). None of the 1,253 Adephaga taxa in our dataset have
Xy+ systems (Figure 3). Xy+ has been reported seven times in Adephaga, but subsequent
investigations failed to replicate the observations (Serrano and Yadav 1984; Hughes and Angus
1999; Aradottir and Angus 2004). In contrast to Adephaga, 60 percent of Polyphaga species
(2,097/3,468) exhibit Xy+.
Models of sex chromosome evolution: The difference in frequency of Xy+ systems
between Adephaga and Polyphaga suggests that different biological mechanisms may act in
each suborder. Therefore, we inferred parameters for models of sex chromosome evolution
independently for each suborder. For Adephaga, we find that Y chromosomes are gained and
lost at a rate of 0.573 ± 0.00052 gains and losses per 100 million years (Figure 2.4).
43
Figure 2.4 Sex chromosome system transition rate estimates. Rates are reported as the
probability of transition per 100 million years ± the standard error. Parentheses indicate the
mean number of transitions inferred. In Adephaga the mean number of transitions is the sum of
transitions between both states.
Since only XY and XO states are observed in Adephaga, the model comparisons reduce
to the difference between models with a single transition rate (model 2.1) and 2-rates (model 2.2;
Figure 2.2). Comparison of marginal likelihoods (Table 2.1) reveals that both models fit the data
equally well in Adephaga (LBF = 1.9, Table 2.1). Indeed, the 2-rate model estimates nearly
identical rates for XY to XO and the reverse (0.574 and 0.572 respectively).
44
Table 2.1 Marginal likelihoods and model comparisons for models of sex chromosome evolution.
Suborder
Adephaga
Model
Marginal Likelihood
Comparison
2.1
-103.5
2.1 vs 2.2
2.2
-102.6
Polyphaga
2.1
-131.2
2.1 vs 2.2
2.2
-125.3
3.1 vs 3.3
3.1
-393.8
3.3 vs 3.6
3.3
-378.6
3.4 vs 3.6
3.4
-367.8
3.6
-362.4
a Indicates very strong support for the more complex model.
Log Bayes Factor
1.9
11.8a
30.5 a
21.5 a
10.7 a
To verify that the best model is able to recapitulate the distribution of empirical data in
Adephaga we conducted PPS under model 2.1. The simulated data sets are centered on the
observed distribution of sex chromosomes (Figure 2.5A), indicating that the estimated
parameterization of model 2.1 can produce outcomes similar to the observed data.
45
Figure 2.5 Distribution of PPS datasets in the suborder Adephaga.
The black lines indicate the density of simulated datasets; the vertical red lines indicate the
number of taxa observed in the XY state. Adequate performance of model 2.1 in Adephaga is
evident by the concentration of datasets similar to the observed data. The poor performance of
model 2.1 in the subtrees composed of the clades Trechitae and Cicindelini + Colyrinae is
evidence that these clades have higher retention rates of the Y chromosome than is expected for
groups in the suborder Adephaga.
46
For the Polyphaga, the more complex of the 2-state models (2.2) is preferred in
comparison with model 2.1 (LBF= 11.8; Table 2.1). Likewise, the most complex 3-state model
(3.6; LBF = 10.7 – 30.5; Table 2.1) is preferred in comparison to all other 3-state models.
Comparison of models 3.4 and 3.6 indicates that the 3-state coding is more appropriate than 2state coding (LBF = 10.7; Table 2.1). Therefore, Xy+ and XY states differ biologically in terms of
the rates of Y chromosome changes they undergo. Changes from XY to Xy+ have the highest
estimated rate among all transitions in Coleoptera (0.65±0.00049), while the rates of transitions
from Xy+ to any other state are the lowest (to XO = 0.033±0.00007; to XY = 0.19±0.00019; Figure
2.4), i.e., , Xy+ distance-pairing sex chromosomes in Polyphaga are the most evolutionarily stable
sex chromosome state in Coleoptera in our analyses. PPS under model 3.6 show that our
phylogenies and model parameterization can produce outcomes similar to the observed data
(Figure 2.6).
47
100% XY
Root State for Simulations
XO
XY
Xy+
0.8
Observed Data
0.8
0.8
0.2
0.6
0.6
0.4
0.4
0.2
0.4
0.2
0.6
100% XO
100% Xy+
Figure 2.6 Distribution of PPS datasets in the suborder Polyphaga.
Each circle represents a simulation based on the parameter estimates from model 3.6, and are
colored to reflect the root state chosen for the simulation. The larger red circle indicates the
observed data. Axes represent the percent of terminal taxa in each of the three sex chromosome
states. The empirical observation being near the densest part of the distribution of simulation
results indicates that model 3.6 adequately predicts sex chromosome evolution in Polyphaga.
The tail of simulations with a high proportion of XO taxa arises in large part from runs where XO
was assigned as the root state, which is unlikely to be the true ancestral state in Polyphaga.
Discussion
The rates and patterns of Y chromosome turnover are distinctly different between the two
largest Coleoptera suborders, Polyphaga and Adephaga. We suggest that important differences
in the meiotic machinery, and particularly sex chromosome pairing, between these lineages
indicates that previously unappreciated evolutionary forces may influence Y chromosome
evolution.
48
In the suborder Adephaga, a Y chromosome has a 57 % chance of being lost or gained
per 100 million years. The 65 Y chromosome losses (Figure 2.4) in Adephaga, are not difficult to
explain. Presumably, these losses reflect the standard population genetic forces promoting Y
decay noted earlier, and that gene losses ultimately reach the point where the Y chromosome
becomes dispensable (Steinemann and Steinemann 2005). Our estimate of an equal number of
Y chromosome gains in Adephaga is more interesting. While some sex chromosomes are
famous for avoiding decay and remaining homomorphic (e.g. ratite birds (Adolfsson and Ellegren
2013; Vicoso et al. 2013) and anurans (Stock et al. 2011; Stock et al. 2013)), the situation in
Adephaga is different. In this group, new Y chromosomes are evolving at the same rate they are
lost.
There are two mechanisms for gaining a Y chromosome: 1) fusion of all or part of an
autosome to the X (White 1977; Charlesworth and Charlesworth 1980; Watson et al. 1991;
Veltsos et al. 2008); or 2) capture of a supernumerary (B) chromosome(Carvalho 2002). To
determine whether transitions from XO to XY are the result of fusions between the X
chromosome and a whole autosome we used stochastic character mapping (Huelsenbeck et al.
2003; Revell 2012) of chromosome number (downloaded from uta.edu/karyodb/) and sex
chromosome system, to calculate the proportion of branches where a Y chromosome gain cooccurs with a reduction in the number of autosomes. This method suggests that at least 49% of
the Y chromosome gains in Adephaga are the result of fusions between the X chromosome and a
whole autosome. This is far more frequent co-occurrence than the 2.7% expected if gaining a
new Y chromosome was independent of decreasing chromosome number (We infer that Y
chromosomes are gained on 22.7 % of branches in the Adephaga tree, and we infer chromosome
losses on 11.9% of branches; the probability of independent co-occurrence is then
0.227*0.119=0.027). This may indicate that sexually antagonistic loci are sufficiently common in
Adephaga genomes that fusions of an autosome to the X are often favored. Without data for Y
chromosome homology we cannot say whether the remaining 51% of branches are primarily due
to translocations (i.e. fusion of partial chromosomes) or B-chromosome captures.
49
In contrast to Adephaga, we estimate that Y chromosomes in the suborder Polyphaga
are more than twice as readily gained as lost (34% probability of gain per 100 my, 157 total gains;
15% probability of loss per 100 my, 69 total losses; Figure 4). To investigate the source of Y
chromosome gains, we mapped transitions from the XO state and transitions in chromosome
number as we did for Adephaga above. We find that only 27% of the Y gains in Polyphaga
coincide with reductions in chromosome number, but that this is still far more frequent than the
0.5% expected if the events were independent (We infer Y chromosomes gains on 6.8 % of
branches in the Polyphaga tree, and chromosome losses on 7.6% of branches; the probability of
independent co-occurrence is then 0.068*0.076=0.005). While fusions clearly coincide with Y
chromosome gains in both suborders, in comparison with Adephaga, a much larger proportion of
Polyphaga Y chromosome gains appear on branches where no reduction in chromosome number
is inferred. This suggests that relative to Adephaga a larger proportion of Y chromosome gains in
Polyphaga result from either B-chromosome capture or the fusion of only a portion of an
autosome to the X chromosome. The presence of an existing mechanism for segregation of
unpaired sex chromosomes in Polyphaga may facilitate the capture of B-chromosomes, and
contribute to the difference in Y chromosome origins between beetle suborders.
Rates of Y chromosome loss are also interesting in Polyphaga because they are so low.
The non-recombining Xy+ sex chromosomes in Polyphaga species do not contain a
pseudoautosomal region (PAR) and thus the entire Y chromosome is subject to the population
genetic forces promoting Y decay. If decay followed by loss is the dominant source of XO
species, as often suggested, it is surprising that Xy+ systems lose their Y 3.5-times less
frequently than XY systems with a PAR (Figure 2.4). Consequently, we suggest that some
evolutionary force(s) promoting retention must be acting in Polyphagan Xy+ systems. As noted in
the introduction, frequent turnover in the sex determining chromosome and or intermittent
recombination can promote retention of homomorphic sex chromosomes, but these mechanisms
do not apply in species with highly degenerate Xy+ sex chromosomes. Other hypotheses for
retention that could apply to the situation in Polyphaga involve purifying selection either due to
50
“essential” male genes or haploinsufficiency (Li et al. 2013). Although little is known about the
genes present on the Y chromosomes of Coleoptera, it seems unlikely that genes required for
male viability are widespread on the Y chromosome, since XO species occur in 24 of 59
Coleoptera families studied, and our estimates indicate that the Y chromosome has been
independently lost approximately 69 and 65 times in Polyphaga and Adephaga respectively
(Figure 2.4).
The argument for retention of the Y chromosome due to haploinsufficiency of X-linked
genes in males depends on whether dosage compensation occurs and to what extent. However,
it seems unlikely to explain our results. Dosage compensation has been studied in only a single
species of coleopteran, the red flour beetle Tribolium castaneum, a polyphagan beetle. In this
species, chromosome wide dosage compensation of the X occurs in males, such that, on
average expression from one X equals that from two autosomes (Prince et al. 2010). This type of
chromosome wide up-regulation should provide haplosufficiency for all the genes on the X
chromosome, reducing purifying selection to maintain Y homologs. We lack information for most
beetle taxa, but the fact that the Y in Xy+ species is typically punctiform also indicates that most X
chromosome genes must be haplosufficient so that haploinsufficiency is unlikely to be a general
explanation for the exceedingly rare loss of Y chromosomes from Xy+ species.
What then can explain the relative stability of Y chromosomes in Polyphaga? We
propose the “fragile Y” hypothesis: when proper segregation of the sex chromosomes depends
on chiasmata, recurring selection to reduce recombination between loci in the PAR (e.g. loci with
sexually antagonistic polymorphisms) and the sex determining locus (1) reduces the size of the
PAR and consequently opportunities for chiasma formation, and (2) this leads to an increased
probability of producing aneuploid gametes (Raudsepp et al. 2012) creating increased
opportunities for Y loss (hence “fragile Y”).
Our hypothesis makes two predictions: 1) as the PAR shrinks, selection should favor
segregation mechanisms that do not rely on chiasmata 2) taxa that evolve achiasmatic pairing
should have lower rates of Y chromosome loss. We tested within the suborder Adephaga for
51
evidence of this pattern. While no adephagans have distance-pairing sex chromosomes of the
type found in Polyphaga, complete achiasmatic meiosis has arisen at least 4 times independently
in Adephaga. Two of these origins involve only one or a handful of species in the genera
Egadroma and Calasoma, suggesting that they arose relatively recently (Serrano 1981).
However two of the origins involve the larger clades of Trechitae (Maddison and Ober 2011) and
Cicindelini + Colyrinae (Galian et al. 2007), and must be older. Both these clades lose the Y
more rarely than expected, consistent with our predictions. In our dataset a total of 45 Trechitae
species are represented. Within these, at most 3 changes from XY to XO have occurred; for the
clade including Cicindelini + Colyrinae we have 21 species and only a single such change. PPS
analyses for these clades suggest that both groups have fewer XO species (i.e. fewer Y
chromosome losses) than expected based on the overall transition rates for Adephaga. In
Trechitae 95% of simulations predict more XO species than we observe (mean expected by
simulation =16.5, versus 3 observed; Figure 2.5B). In Cicindelini + Colyrini 92% of the
simulations predict more XO species than the empirical observation (mean expected by
simulation = 5.8, versus 1 observed; Figure 2.5C).
Mammals: An additional opportunity to test the fragile Y hypothesis is available in
mammals. The infraclasses Eutheria (placental mammals) and Metatheria (marsupials) offer a
parallel example to Adephaga and Polyphaga in beetles. The sex chromosomes of metatherian
mammals segregate in males without the presence of a PAR or chiasmata (Page et al. 2006) and
no cases of Y loss are reported. In contrast, the Eutherians generally require a PAR region that
forms chiasmata to faithfully segregate the sex chromosomes, and in taxa with small PARs the Y
is occasionally lost (Fernández-Donoso et al. 2010). Among eutherian mammals, the rodents
have the smallest documented PAR (Raudsepp and Chowdhary 2008), and it is within the
rodents that we see multiple independent responses to the forces we ascribe to a fragile Y. First,
within the family Cricetidae, the genus Microtus exhibits at least 3 origins of achiasmatic sex
chromosomes, and as we expect there are no reported Y chromosome losses (Table 2.2)
(Borodin et al. 2012).
52
Table 2.2 Summary of mammals with achiasmatic X-Y segregation or Y chromosome losses.
Clade
Cricetidae
Microtus
X-Y Segregation
Mechanism in Males
predominately
achiasmatica
chiasmatic
achiasmatic
Taxa
29
Independent
Y losses
0
3
9b
1-2
0
Citation
(Borodin et al. 2012)
(Just et al. 1995)
(Ratomponirina et al.
1986; Ratomponirina
et al. 1989)
Tokudaia
chiasmatic
3
1
(Arakawa et al. 2002)
a 17 species are achiasmatic, 12 chiasmatic with at least 3 independent origins of achiasmatic
segregation
b 4 species of Gerbillinae have experienced autosome sex chromosome fusions, the autosomal
portion of which undergo crossover.
Muridae
Ellobius
Gerbillina
e
In contrast, the closely related mole vole genus Ellobius has not evolved achiasmatic
meiosis but shows at least one, and possibly two instances of Y chromosome loss (Just et al.
1995). Second, within the largest family of mammals, Muridae, we find additional origins of
achiasmatic male meiosis and Y chromosome loss. The subfamily Gerbillinae has evolved
achiasmatic sex chromosomes and Y losses are not reported, whereas in a related subfamily
Murinae, which has not evolved achiasmatic meiosis, the spiny rat genus Tokudaia has three
species two of which have lost the Y chromosome(Arakawa et al. 2002), and a third whose Y
chromosome is fused with an autosome, rejuvenating the PAR and escaping potential difficulty in
segregation during male meiosis (Murata et al. 2012). While this is a small sample, it is worth
noting that in mammals the Y often carries genes essential for male viability and, all else being
equal, is thus likely to be under stronger selection to be retained than in Coleoptera. The
repeated evolution of either achiasmatic meiosis, or Y chromosome loss, in eutherians with the
smallest PAR size is precisely what the fragile Y hypothesis predicts.
Conclusion
Our analysis suggests that meiotic mechanisms play an important, previously
unappreciated role in the tempo of Y chromosome gain and loss. Additionally, given the
relatively widespread loss of Y chromosomes among Coleopterans, sex determination seems
53
likely to often involve an X counting system such as in Drosophila melanogaster (Bridges 1921)
where the Y plays little role in sex determination (otherwise it would not be dispensable)
(Bachtrog 2013). Finally, despite being the largest analysis of its kind, our analysis of sex
chromosome evolution based on available karyotype data is relatively coarse. A more nuanced
understanding of sex chromosome evolution, one that tests the predictions of this study, would
benefit from genomic data that allows for assignment of chromosomal homologies. Given the
large number of novel Y chromosomes arising in Coleoptera (225, figure 2.4), many of which are
fusions or potential B-chromosome captures, it will be interesting to investigate whether some
genes (or chromosomes) are recurrently recruited to Y chromosomes.
* This chapter was previously published as:
Blackmon, H., & Demuth, J. P. (2014). Estimating tempo and mode of Y chromosome turnover:
explaining Y chromosome loss with the fragile Y hypothesis. Genetics, 197(2), 561-572.
54
Chapter 3
Assessing the impact of key ecological and phenotypic transitions on the rate of karyotype
evolution: drift drives the evolution of chromosome number.
Abstract
Chromosomal mutations such as fusions and fissions are often thought to be deleterious,
especially when heterozygous (i.e. underdominant), and consequently are unlikely to become
fixed. Yet, models of chromosomal speciation ascribe an important role to chromosomal
mutations. When effective population size (Ne) is small the efficacy of selection is weakened and
the likelihood of fixing underdominant mutations by random genetic drift is greater. Thus it is
possible that ecological and phenotypic transitions predicted to modulate Ne may facilitate fixation
of chromosome changes, increasing the rate of karyotype evolution. We synthesize all available
data on chromosome number in Coleoptera, and estimate the impact of traits expected to change
Ne on the rate of chromosome number evolution in the family Carabidae and in 12 disparate
genera from across the phylogeny of beetles. Our analysis indicates that in Carabidae, wingless
clades have faster rates of karyotype evolution. Additionally, the genus level analysis shows that
all groups with high rates of karyotype evolution exhibit multiple traits that are expected to reduce
Ne, including sibmating, oligophagy, winglessness, and island endemism. The observed
connection between reduced population size and accelerated karyotype evolution suggests that
changes in chromosome number are likely fixed by random genetic drift despite an initial fitness
cost, and that Ne is small enough for chromosomal speciation models to potentially be important
drivers of speciation in many Coleopteran clades.
Introduction
Understanding the evolutionary forces that underlie strong karyotype conservation in
some groups (Boyes and Shewell 1975; White 1978) while others are more labile (Kandul et al.
2007; Carbone et al. 2014) has remained elusive despite over 50 years of work. Historically, one
55
of the key issues has been the relative importance of natural selection versus random genetic
drift as the principal driver of change in chromosome number. However, recent analysis of the
Gibbon genome suggests that mutation rate may also be an important factor determining the rate
of karyotype evolution (Carbone et al. 2014).
Hypotheses that propose an important role for positive selection to change chromosome
number are easily allied with arguments for the costs and benefits of recombination (Muller 1932;
Fisher 1958; Muller 1964; Hill and Robertson 1966; Nei 1967; Lewontin 1971; Felsenstein 1974).
Particularly when recombination is limited to a maximum of one crossover per chromosomal arm
per meiosis, selection for increased or decreased recombination is expected to propel
corresponding changes in chromosome number. Additionally, selection has been argued to
favor increased chromosome number in social insects because having more chromosomes
increases the average relatedness within a colony - limiting the opportunity for kin-recognition
based cheating (Sherman 1979; Templeton 1979) - while simultaneously allowing for high
genotypic diversity among sibs which has been shown to benefit colony growth, efficiency, and
pathogen resistance (Tarpy 2003).
Countering the general applicability of positive selection’s role as the agent of karyotype
evolution, is the observation that most changes in karyotype are either neutral or deleterious, with
many being underdominant (i.e. deleterious when heterozygous)(Max 1995). Underdominance of
karyotype changes results from difficulties encountered in meiosis where mismatched
chromosomal types do not segregate properly. Such underdominant mutations are only expected
to fix when natural selection is overcome by random genetic drift in small populations (Wright
1941; Lande 1979, 1985). Despite the difficulties of fixing underdominant mutations, attempts to
ascribe a causal role for karyotype evolution in the speciation process have been abundant
(Lewis 1966; White 1978; Bickham and Baker 1979; Grant 1981; Templeton 1981; Baker and
Bickham 1986; Rieseberg 2001). Some models assume that karyotype changes themselves are
neutral but facilitate diverging local adaptation by sheltering some genome regions from the
homogenizing effects of gene flow and recombination. Other models posit that underdominant
56
karyotype changes that are differentially resolved between isolated populations by random
genetic drift, may then act directly as isolating barriers upon secondary contact (see Rieseberg
2001 for more detailed review of chromosomal speciation models).
Given that effective population size (Ne) governs the efficacy of natural selection in
relation to random genetic drift, one way to distinguish among these two evolutionary forces is to
see whether factors associated with differences in Ne are also associated with differences in the
rate of karyotype evolution and or the direction of change in chromosome number. More
specifically, if karyotype changes are neutral, then their fixation rate should be unrelated to Ne.
However, if karyotype changes are deleterious, then species with small Ne should have faster
rates of chromosome evolution as more changes are fixed by drift. In contrast, if selection plays
a broad role then we expect species with larger Ne to have faster rates. Furthermore, unlike
random genetic drift, which is expected to affect the rate but not the direction of karyotype
evolution, selection should be associated with a directional change (i.e. rate changes due to
selection consistently cause all species within a lineage to go up or down).
Irrespective of whether selection or random genetic drift is responsible for fixation of new
karyotypes, recent analysis of gibbon genomes suggests that an elevated rate of karyotype
evolution may also be a signature of elevated chromosomal mutation rate. In the gibbon lineage,
a transposable element that preferentially inserts into chromosome segregation genes may be
responsible for increased rates of chromosomal rearrangements, thereby helping explain why the
karyotypes of these small apes have diversified to a range of 2n=38 - 52 in the past 4 - 6 million
years (Carbone et al. 2014) while the rest of the ape lineage varies by only a single autosomal
fusion (2n=48 - 46) over more than twice that time (Stanyon et al. 2008; Locke et al. 2011). If
increased mutational input is the primary cause of faster karyotype evolution, rather than
selection or drift, then increases in the rate of change are not expected to be directional or
associated with Ne.
Efforts to relate Ne to karyotype evolution typically use ecological and phenotypic traits as
proxies for expected differences in Ne. For instance, all else being equal, winged species should
57
have larger Ne than wingless species because flight increases dispersal distances (Ne = 4πσ2δ,
where σ2 is a measure of dispersal distance;(Wright 1946). Other traits that have been used as
proxies for Ne include: mating system (inbreeding vs. outbreeding), geographic distribution (island
vs. continental), and feeding type (restricted vs varied diet). The distribution of these traits has
then been compared to the rate of karyotype change as estimated by scaling the variance in
chromosome number to a fossil date for the taxonomic group of interest (Wilson et al. 1975; Bush
et al. 1977; Bengtsson 1980; Imai et al. 1983; Larson et al. 1984; Petitpierre 1987; Olmo 2005).
These earlier studies suggest that taxa inferred to have highly structured populations also have
faster rates of karyotype evolution. In some cases there is also a negative correlation between
the estimated rate of karyotype evolution and allozyme heterozygosity levels further supporting
the hypothesis that random genetic drift in small populations drives increased rates of
chromosome change (Coyne 1984). However, previous work has been limited by not
incorporating phylogenies or evolutionary models for chromosome change and using
comparisons between highly divergent clades where the underlying mutation rates may be
different (i.e. across all vertebrates).
The present study expands previous efforts to understand Ne’s effect on karyotype
evolution in two ways. First, we incorporate ecological and phenotypic proxies for Ne into our
comprehensive database of Coleoptera karyotypes (Blackmon and Demuth 2014a). Second, we
employ a modern statistical phylogenetic framework to model the relationship between Ne and
chromosome evolution. Coleoptera are an excellent group to study the effect of Ne on
chromosome evolution because they exhibit variation in all four types of traits traditionally used as
proxies for Ne (Crowson 1981), there is a good phylogenetic scaffold, and the database of
karyotypes is extensive (4,797 species).
We show that lineages that have multiple traits associated with reduced Ne (loss of
wings, inbreeding, island endemism, restricted feeding) also have significantly faster rates of
karyotype evolution. Our findings suggest that random genetic drift is the predominant driver of
58
fast karyotype evolution, as predicted if changes to chromosome number are typically
underdominant or mildly deleterious.
Methods
Data collection
We compiled all available karyotypes from the Coleoptera Karyotype Database
(www.uta.edu/karyodb). Coleoptera karyotypes rarely include banding data and are normally
reported as the meioformula, consisting of the number of autosomes plus the sex chromosome
complement of the male. For these reasons we use the male diploid number as a surrogate for
the karyotype and for the remainder of the paper we refer to this as the chromosome number, we
describe the rate of change in chromosome number as the rate of karyotype evolution. In 19
cases where multiple values were reported for a species, the mean value was used. Since some
studies have shown that chromosome number is not normally distributed (Mank and Avise 2006),
we conducted analyses with both the raw chromosome numbers as well as log transformed
values, but we found that this does not change our conclusions and therefore we present results
based on untransformed data.
Lack of overlap in the available data for karyotype, phylogeny, and Ne related traits
resulted in our analysis being subdivided into a family level analysis of Carabidae using presence
absence of wings, and a sparser but more phylogenetically diverse analysis of 12 genera using
multiple Ne influencing traits. Data for the presence of wings in Carabidae was taken from a
previous compilation of natural history data (Larochelle and Lariviere 2003) (appendix C).
Species reported as being polymorphic for wings were scored as having equal probability of
being either winged or wingless. If wing data was not present for a species in the karyotype and
phylogenetic datasets, but other species in the genus were reported, the species was assigned a
probability reflecting available data for the genus. For instance, in the genus Calathus 64% of
species were reported as wingless, so any Calathus species not in the trait dataset were
assigned a 64% probability of being wingless and 36% probability of being winged.
59
For each of the 12 genera included in our genus level rate estimates we performed
literature searches to score them for the following traits: winged vs. wingless inbreeding vs.
outbreeding, island vs. continental distributions, and oligophagy vs. polyphagy. We use the
number of times high or low Ne traits occur in each group to classify them into high, medium, or
low expected Ne classes (Table 3.1). To account for phylogenetic uncertainty in our comparative
analysis we used 100 trees from the posterior distribution produced in an earlier study (Blackmon
and Demuth 2014a). Briefly, these trees are based on an analysis of seven genes (16s, 18s,
28s, COI, elongation factor 1, arginine kinase, and wingless) across 1042 taxa in BEAST (v1.7.5;
(Drummond and Rambaut 2007; Suchard and Rambaut 2009)). We assumed a lognormal
relaxed clock and used normal distributions to place priors on the age of seven nodes; ages were
based on previous estimates (McKenna and Farrell 2009).
Phylogenetic model based analyses
To test whether chromosome number is under selection to increase or decrease, we first
tested whether the groups in our datasets had significantly different absolute number of
chromosomes as would be expected for instance if small population size selected for more
chromosomes. For the Carabidae data we calculated phylogenetically independent contrasts for
both chromosome number and the probability that a clade was winged using the R package APE
(Paradis 2011). We then computed Pearson’s correlation coefficient to test for a correlation
between chromosome number and probability of being winged. For the genus level data we used
the R package GEIGER (Harmon et al. 2008) to compute a phylogenetically corrected ANOVA
that tests whether our three Ne classes have significantly different chromosome numbers. We
used 1000 simulations to assess significance of the F-statistic (Garland et al. 1993)
Next, for the family and genus level data we used censored rate tests based on Brownian
motion models to determine whether the data indicate multiple rates of karyotype evolution as
implemented in the R package Phytools (O'Meara et al. 2006; Revell 2012). We compare a
model where the continuous trait (chromosome number) evolves at a single rate on all branches,
to a model where each discrete state has an independent rate of chromosome number evolution.
60
In the analysis of Carabidae our discrete states were winged and wingless and in the analysis of
genera our states were low, medium and high Ne. Conducting the censored rate test required
reconstruction of the history of the discrete state across the phylogenies. In the analysis of
Carabidae, since wing loss is widely accepted as a derived state within Coleoptera we fixed the
root state of the tree as winged (Grimaldi and Engel 2005). For the genus level analyses we fixed
the root of the tree as high Ne since the last common ancestor of the included genera is expected
to have all high Ne traits. (Grimaldi and Engel 2005). We used an all rates differ Mk model
(allows for rates to be different into and out of each state) to estimate the parameters of the
transition rate matrix, and used stochastic mapping to assign a state along all branches in the
tree. To account for uncertainty in phylogenetic inference and ancestral state reconstruction we
performed five stochastic mappings on each of our 100 trees. Since previous work has shown
that different rates of chromosome evolution may be occurring in the two major suborders of
beetles we analyzed the genera in each suborder separately (Blackmon and Demuth 2014a). To
explore whether the result of censored rate tests were being driven by exceptional rates in a
single clade we also independently estimated the rate of karyotype evolution in each of the 12
genera. The R packages Geiger version 2.03 and Phytools version 0.4-21 were used to
reconstruct ancestral states and fit models of chromosome number evolution (Harmon et al.
2008; Revell 2012).
Scaled variance estimates
Since the lack of overlap between species trait data and existing phylogenetic information
causes a large reduction in the number of datapoints in our analysis, we also investigated
whether estimating the rate of karyotype evolution without incorporating phylogenies is consistent
with the phylogenetic model based approach above. We calculated time scaled coefficients of
variation by first locating the oldest available fossil record for each genus of interest in the
Paleobiology Database (http://paleodb.org). We then used the fossil ages to scale the
coefficients of variation for chromosome number in each taxon (family or genus). To assess
consistency between these “scaled variance” estimates and the phylogenetic model based rate
61
estimates, we used a non-parametric correlation analysis (Kendall’s τ). The test was one-tailed
since we expect either no significant correlation or a positive one. All tests were considered
significant at p-value < 0.05.
Results
Data collection
We downloaded 4,537 records from the Coleoptera Karyotype Database
(uta.edu/karyodb) (Appendix A). This included data for all four extant suborders of Coleoptera.
Two of these suborders are represented by only one and two karyotypes and thus we focused
our analysis on the larger suborders of Adephaga and Polyphaga. These two suborders
accounted for 1,224 and 3,310 karyotypes respectively. In Adephaga the number of autosomes
ranged from 3 to 34 (mean =15.57±0.14), while in Polyphaga the range was from 1 to 32 (mean
=10.63±0.06). Polyphaga exhibits a single mode of nine autosomes, accounting for 952 species
or 29% of all Polyphaga records. Conversely, Adephaga is bimodal with concentrations at 11 and
18 autosomes accounting for 276 and 242 species or 23% and 20% respectively (Figure 3.1).
62
Figure 3.1 Haploid number of autosomes in the two major suborders of Coleoptera.
The height of the bars reflects the number of records indicating a species with that haploid
number and is based on 4,534 records available from www.uta.edu/karyodb.
Selection on chromosome number
If chromosome number comes under directional selection it could increase the rate of
evolution in the lineage experiencing selection. However, unlike an increased rate due to random
genetic drift, we expect change in response to be directional and result in a correlation between
absolute chromosome number and traits we have associated with Ne. For instance, if wing loss
consistently selects for increased chromosome number we expect wingless species to have more
chromosomes on average. In Carabidae we find no evidence for a relationship between
chromosome number and the probability that a species has wings (r= -0.039, t=-0.46, p-value =
0.65). Likewise, in the analysis of genera we found no relationship between Ne classes and
chromosome number (F=4.06, p-value = 0.89). These results suggest that selection on
chromosome number is not the primary influence on rates of karyotype evolution in these
lineages.
63
Phylogenetic model based rate estimates
Karyotypes for 1065 Carabidae species were available, 136 of these were used in our
comparative analysis because they were included in our phylogenetic tree and had data available
on flight ability. The censored rate test supports the conclusion that chromosome number
evolves at different rates in winged and wingless clades. The single rate model was rejected on
all 500 stochastically mapped trees (max p-value < 0.01). Our analyses of Carabidae show that
wingless lineages gain and lose chromosomes 6 times faster than their winged relatives. The
mean estimate for the rate parameter σ2 in wingless clades was 9.15±0.41 while the mean for
winged clades was only 1.51±0.08 (Fig. 3.2a).
64
Figure 3.2 Phylogenetic based estimates of the rate of chromosome number evolution.
Rates are the maximum likelihood estimates across 500 stochastically mapped trees. a)
Estimates for the family Carabidae under a two-rate Brownian motion model. This dataset
included 136 species scored as winged or wingless. b) Estimates under a three-rate Brownian
motion model for the suborder Polyphaga with seven genera totaling 149 species. c) Estimates
under a three-rate Brownian motion model for the suborder Adephaga with five genera totaling
112 species.
For the analysis of genera we scored each clade for the presence or absence of traits
thought to reduce Ne. This allowed us to assign each genus to a class based on expected Ne.
Four genera posses none of the Ne reducing traits and we classify these as the high Ne class.
Three possess only one of these traits and form the medium Ne class, and five genera (Calathus,
65
Chrysolina, Cytronus, Dendroctonus, and Timarcha) posses two of the Ne reducing traits, and
these form the low Ne class (Table 3.1).
Table 3.1 Distribution of traits likely to effect population size.
Traits were scored based only on species included in the analysis and the majority state is
reported below. Expected Ne is categorized by the number of traits expected to reduce Ne. High,
medium, and low Ne categories were assigned if a clade had zero, one, or two Ne reducing traits
respectively. A (+) indicates the high Ne version of a trait and a (-) indicates the low Ne version of
Polyphaga
Adephaga
a trait.
Genus
Bembidion
Calathus
Cicindela
Harpalus
Pterostichus
Chrysolina
Cyrtonus
Dendroctonus
Diabrotica
Ips
Pimelia
Timarcha
Breeding
+
+
+
+
+
+
+
+
+
+
+
Feeding
+
+
+
+
+
+
+
-
Distribution
+
+
+
+
+
+
+
+
+
+
+
Wings
+
+
+
+
+
+
-
Expected Ne
high
low
high
high
medium
low
low
low
high
medium
medium
low
The censored rate test supports the conclusion that chromosome number evolves at
different rates in the different Ne classes. The single rate model was rejected for both datasets on
all 500 stochastically mapped trees (max p-value < 0.01). The mean estimate for the rate
parameter (σ2) was highest for the low Ne class in both suborders. In Polyphaga the low Ne class
had the highest rate of chromosome evolution (0.624±0.098); the medium Ne class exhibited an
intermediate rate (0.065±0.013) and the high Ne class had the slowest rate of chromosome
evolution (0.017±0.001) (Fig. 3.2b). In Adephaga rates of karyotype evolution are typically much
higher than in Polyphaga, but the low Ne class again had the highest rate of chromosome
evolution (24.401±8.627); unexpectedly the medium Ne class exhibited the lowest rate
(0.091±0.034) and the high Ne class had an intermediate rate (0.369±0.057) of chromosome
evolution (Fig. 3.2c).
66
Independent estimates for the rate parameter σ2 for karyotype evolution within each of
the 12 genera examined provide insights into the impact of each genus on the results described
above. The rate estimates for individual genera ranged from near zero in the genera Diabrotica
and Pimelia to as high as 24.33 (95% CI = 12.17 - 48.66) in Calathus (Table 3.2). In each
suborder the low Ne class genera exhibited the highest rates of karyotype evolution. Those
genera in the high and medium Ne groups showed similarly slow rates.
Estimates based on scaled variance
For each of the 12 genera included in our rate estimate inference, we searched PaleoDB
for fossil records. Seven of our target genera had fossil data available; if multiple dates were
available we recorded the age of the oldest available record. These ages ranged from a low of
7.24 million years for the genus Chrysolina to 150.8 million years for the genus Cicindela (Table
3.2). We calculated the scaled variance of chromosome number for each of the 7 genera by
dividing the coefficient of variation for chromosome number by the fossil based age estimates
(Table 3.2). These scaled variances ranged from 0.07 in Cicindela to 3.87 in the genus
Chrysolina. Notably, we find that there is no correlation between the scaled variance based
estimates of karyotype evolution and the phylogenetic model based rate estimates (σ2 parameter
above) for the 7 genera with overlapping analyses (τ =0.143, p-value=0.773).
67
Table 3.2 Genus level phylogenetic and scaled variance based estimates of rate of chromosome
evolution. Dashes indicate those taxa for which reliable fossil dates where unavailable and where
not included in comparison of approaches.
Clade
NMR1 MR2
95% CI3
NSCV4 CV5 Age
Bembidion
40
0.243 0.157 - 0.376
230
0.06 48.6
Calathus
16
24.33 12.16 - 48.66
36
0.12 28.4
Cicindela
27
0.462 0.271 - 0.788
83
0.10 150.8
Harpalus
14
0.515 0.246 - 1.082
31
0.05 65.5
Pterostichus 15
0.088 0.043 - 0.180
58
0.09 37.2
Chrysolina
26
0.558 0.324 - 0.961
64
0.28 7.24
Dendroctonus 13
0.323 0.150 - 0.697
16
0.32 46.2
Cyrtonus
13
1.279 0.593 - 2.758
Diabrotica
12
0.008 0.004 - 0.018
Ips
26
0.117 0.068 - 0.201
Pimelia
29
0.002 0.001 - 0.004
Timarcha
30
0.438 0.264 - 0.726
1 number of taxa used in phylogenetic model based estimate of rates
2 mean rate of chromosome change (phylogenetic approach)
3 95% confidence interval (phylogenetic approach)
4 number of taxa used in calculation of scaled variance
5 coefficient of variation.
6 scaled CV = CV/Age/100MY
SCV6
0.12
0.42
0.07
0.08
0.24
3.87
0.69
-
Fossil Reference
(Arillo and Ortuno 1997)
(Martynov 1929)
(Weyenbergh 1869)
(Birket-Smith 1977)
(Wickham 1910)
(Hopkins et al. 1971)
(Labandeira et al. 2001)
-
Discussion
Our analyses clearly show that ecological and phenotypic transitions that are expected to
reduce Ne are associated with increased rates of karyotype evolution and that this pattern holds
independently of potential differences in mutation rate between lineages. The most likely
explanation for this pattern is that chromosome number changes are predominantly deleterious
while segregating and become fixed by random genetic drift. We do not find support for a
directional trend in gain or loss of chromosome number in response to trait evolution that might
indicate a signature of selection.
The effect of reduced Ne on accelerated rates of karyotype evolution is dramatic. In
Carabidae wingless clades show a 6-fold increase in the rate of karyotype evolution. In
Polyphagan genera the difference between low and high Ne genera is >26-fold and in Adephagan
genera it is greater than 30-fold. A few notable examples serve to highlight this overall pattern in
beetles. First, within the Polyphaga family Curculionidae, our study includes the closely related
68
scolytid genera Ips and Dendroctonus. Our estimate for the mean rate of karyotype evolution in
Dendroctonus 0.323 (95% CI = 0.150-0.697) was 3-fold higher than the mean rate estimated in
Ips 0.117 (95% CI = 0.068-0.201). This marked difference matches expectations based on
breeding behavior. Dendroctonus is an inbreeding genus producing biased sex ratios and
practicing predirspersal sibmating (Grégoire 1988). Meanwhile Ips is an outbreeding genus
where both males and females disperse, and neither sib-mating nor biased sex ratios have been
documented (Kirkendall 1993). These characteristics should lead to smaller Ne in Dendroctonus
and should allow changes in karyotype to fix more easily even if they are underdominant as
theory predicts.
Second, three of the four highest rates of karyotype evolution were observed in wingless,
oligophagous genera within the family Chrysomelidae: Timarcha, Cyrtonus, and Chrysolina
(0.438, 1.279, 0.558 respectively). In all three genera the mean maximum likelihood estimate
(MLE) for the rates of karyotype evolution were higher than the MLE of Diabrotica 0.008 (95% CI
= 0.004-0.018), a chrysomelid genus lacking any of the small Ne traits.
Finally, the genus Calathus provide perhaps the best example of compound effects of
phenotype and ecological history on the tempo of karyotype evolution. First, many species in this
genus are wingless and thus may be characterized by populations composed of small demes
where fixation of karyotype changes should be more likely. However, the exceptionally high rate
estimate for karyotype evolution in this genus 24.33 (95% CI = 12.17-48.66) is likely driven by
two taxa; Calathus abaxoides and C. ascendens which have respectively the highest and lowest
chromosome number in the genus. Interestingly, C. abaxoides and C. ascendens are 2 of 24
endemic, wingless, Calathus species on the Canary archipelago. Both species occur on the
island of Tenerife which the genus has colonized in the last 12 million years (Emerson et al.
1999). These species likely experienced an initial population bottleneck during colonization, and
the continued restriction to an island has led to a sustained lower Ne than species with continental
distributions. While 17-19 autosomes is the norm for most species in this genus, C. abaxoides
has increased to 27 autosomes while C. ascendens has decreased to 10 autosomes. The
69
observation that both the lowest and highest chromosome number are the product of a single
recently colonized island further suggests that drift in small populations is responsible for rapid
karyotype evolution.
The role of mutation
Little is known about mutation rate variation in beetles, however we recently
hypothesized that differences in the mechanisms of meiosis may provide a mutational basis for
differences in the rate of sex chromosome turnover between the two main beetle suborders
Polyphaga and Adephaga (Blackmon and Demuth 2014a). The present analysis demonstrates
that two suborders also have very different overall rates of chromosome evolution (note the
difference in scales between figures 3.2b and 3.2c) that are consistent with our findings on sex
chromosome rates. It is noteworthy however, that despite a >10-fold difference in the
“background” rate of karyotype evolution, the pattern where small Ne is associated with relatively
rapid karyotype evolution holds within both suborders. Thus, while mutation rate may be a major
factor driving the baseline rate of karyotype evolution, our analysis suggests that within a given
mutational context, most changes are at least mildly deleterious and become fixed by random
genetic drift.
Comparison with previous work
While our findings are in accord with earlier studies relating Ne to variation in
chromosome number, our phylogenetic model based rate estimates are not correlated with timescaled variance estimates derived similarly to previous work. This inconsistency is worth noting
because the scarcity of reliable phylogenies is a limiting factor to conducting analyses in other
groups. The lack of consistency between approaches highlights the risk inherent in ignoring the
pattern of chromosome evolution over the phylogeny. Theoretically, a scaled variance method
could work. However its accuracy will be limited by the extent to which the ages estimated for the
groups are both accurate and correlated with the total phylogenetic branch lengths relating the
focal taxa. These requirements are unlikely to be met particularly in groups that have relatively
incomplete and highly heterogeneous fossil records such as insects. Methods not using a
70
phylogeny will also be misled when the number of records is insufficient to capture the true
variance of the groups being studied. The variance in chromosome number across families of
Coleoptera can be partly explained by the number of records available (Pearson’s correlation
coefficient between family variance and number of records =0.41, p-value=0.008). This suggests
that some families have not been sampled sufficiently to capture the true variance of extant
species. Applying an evolutionary model for karyotype evolution using a time scaled phylogeny
eliminates these issues.
Conclusion
There are almost certainly many individual cases where selection has driven a change in
karyotype. For instance it has it has often been suggested that eusocial hymenoptera may
through selection for increased recombination have evolved higher numbers of chromosomes.
However, when chromosome numbers of solitary and eusocial hymenoptera were analyzed in a
comparative framework the results suggested that selection for increased chromosome number is
variable across eusocial hymenoptera (Ross et al. 2014). Our results in concert with previous
work suggest that most chromosome changes are deleterious (at least while segregating) and
that chromosome evolution is largely governed by random genetic drift in small populations. The
association we find between factors influencing Ne and evolutionary rate also puts bounds on the
selection coefficient of mutations, suggesting that many changes are likely to be only mildly
deleterious; otherwise reduced Ne due to ecological and phenotypic transitions in Coleoptera
would not be sufficient to drive significant increases in the number of chromosome changes that
are fixed by random genetic drift. Our findings also indicate that the distribution of fitness effects
of karyotype change is independent of the mutation rate.
More broadly, our work suggests that when species evolve traits, or inhabit locations that
restrict population size, the rate of change in chromosome number increases often by orders of
magnitude relative to closely related species. By increasing the fixation rate for karyotype
changes, speciation mechanisms requiring genome rearrangements become more likely. This
should be true for models that assume underdominance of karyotype mutations such as
71
described by White (1978) and more recent models that assume karyotype changes to be neutral
such as described by Riesberg (2001). Traditionally chromosomal rearrangements are thought to
be more likely to contribute to speciation in plants than animals; possibly due to gene expression
in pollen or lack of differentiated sex chromosomes in most plants (reviewed in Rieseberg 2001).
Chromosomal speciation has also been suggested to be more likely in mammals than
invertebrates due to differences in meiosis (Coyne and Orr 2004). However, given that our results
suggest most karyotype changes in beetles are deleterious, models that invoke karyotypic
changes acting directly as reproductive barriers seem more widely plausible than they have been
considered recently. Unfortunately, the coarse nature of our karyotype data limits our analysis to
mutations such as fusions and fissions that change the number of chromosomes. Future work
incorporating genomic data would be useful to determine whether other types of mutations such
as inversions, and translocations also reflect a similar pattern.
72
Chapter 4
Genome structure and the origin of haplodiploidy
Abstract
Haplodiploid reproduction, where males are haploid and females are diploid, is
widespread among animals. Yet we understand very little about the forces responsible for the
evolution of haplodiploidy. We use a phylogenetic ANOVA and develop a novel comparative
method to test Bull’s long-standing hypothesis that the proportion of the genome that is sex linked
is important in the origin of haplodiploidy (1983). Briefly this approach tests whether
haplodiploidy originates when chromosome number has extreme values. Using simulated data
we show that our new approach has acceptable type one error rate but low power unless the
relationship between the evolution of chromosome number and haplodiploidy are strong. We use
the group Acari that has numerous independent origins of haplodiploidy to evaluate Bull’s
hypothesis, and find that the proportion of the genome that is sex linked is important in
determining if transitions to haplodiploidy occur.
Introduction
Reproduction and sex determination are remarkably variable across life. In most species
reproduction is highly symmetrical; each offspring has a father and a mother and receives an
equal share of their nuclear genes (barring sex chromosome-linked loci). However this symmetry
has frequently broken down. In a significant proportion of animals, the genetic roles of mother
and father during reproduction are highly imbalanced. In about 15% of species, mothers
monopolize the production of male offspring via haplodiploidy, either by the asexual production of
sons (arrhenotoky) or by producing sons that eliminate their father's genome from their germline
(Paternal genome elimination) (Ashman et al. 2014). The evolution of haplodiploidy remain poorly
understood, though not for lack of interest. Many authors have considered different scenarios for
its evolution, and a wealth of theory has been developed on this topic (Brown 1964; Bull 1979;
73
Borgia 1980; Bull 1983; Sabelis and Nagelkerke 1988; Haig 1993a, b; Goldstein 1994; Smith
2000; Normark 2004; Normark 2006; Immler and Otto 2014). Yet few of these ideas have been
corroborated in an empirical or comparative framework. Here, we present the first formal
comparative analysis to understanding the evolutionary dynamics of haplodiploidy.
Most theories assume that haplodiploidy arises from maternal–paternal genetic conflict.
Hartl and Brown (Hartl and Brown 1970) and Bull (Bull 1979) presented the first formal models
showing that when a mother is able to produce haploid sons (either by haplodiploidy or PGE), she
benefits, as these sons always pass on her genes to their offspring (instead of only half of her
genes in a diploid son). However selection for producing haploid sons is counterbalanced by an
expected lower viability of haploid males. As a result haplodiploidy will only spread if haploid
males are at least half as viable as diploid males. This poses a constraint on the evolution of
haplodiploidy and PGE, as most diploid species have considerable numbers of recessive lethal
mutations in their genome and therefore are unlikely to survive as a haploid. Subsequent models
for the evolution of haplodiploidy also stress the importance of viability constraints, making the
general prediction that transitions towards haplodiploidy should be difficult.
One factor that could affect the viability of haploid males is their genome structure: All
transition from diploidy to haplodiploidy took place under male heterogamety (Gardner and Ross
2014), where males are either XO or XY. Under a XO-male sex determining system (or XY with a
degenerate Y), the X chromosome shows haploid gene expression. Therefore, the frequency of
X-linked recessive deleterious alleles is expected to be low, and species who’s X-chromosome
makes up a large proportion of their genome are expected to have a lower frequency of recessive
deleterious alleles in the population. Assuming that on average autosomes and X chromosomes
have a similar size, this leads to the testable prediction that species with very few chromosomes
(e.g. one X and 2 autosomes) are more likely to evolve haplodiploidy because a larger proportion
of the genome will already be haplosufficient than in species with many chromosomes (e.g. one X
and 20 autosomes). In this manuscript we present the first formal comparative tests of this
74
hypothesis using a phylogenetic ANOVA and by estimating the chromosome number of lineages
that evolve haplodiploidy.
Our analyses focuses on the Acari (mites and ticks) which are uniquely suitable for such
an approach: In most other groups haplodiploidy is fixed across large clades (e.g. Hymenoptera)
and appears to have evolved only once in the common ancestor, but in Acari it has evolved
repeatedly. We use reproductive data on 900 species of Acari combined with a phylogeny of 770
species.
Methods
Data collection
We collected all published data on the reproduction, ploidy and karyotypes of Acari, in
particular from three important reviews of Acari reproduction (Oliver 1971; Oliver Jr 1977; Norton
et al. 1993). We supplemented this dataset further with an extensive survey of the literature, by
direct searches on Web of Science and Google scholar, as well as by forward and backward
citation searches. We scored the reproductive mode (sexual vs. parthenogenetic), the
reproductive systems (haplodiploidy vs. diplodiploidy), sex determination system (XY vs. XO) and
the number of chromosomes (diploid chromosome count in females). In total we obtained
reproductive data for 888 species, although the character matrix is not complete for every
species. All data including the references are available on the NESCent “Tree of Sex” database
(www.treeofsex.org) (Tree of Sex Consortium 2014) and Appendix D.
Phylogenetic reconstruction
We downloaded phylogenetically-informative DNA sequence clusters as FASTA files
from PhyLoTA rel 1.5 (Sanderson et al. 2008). We started with data from three mitochondrial
genes (COI, 12S, 16S) and 5 nuclear genes (EF1alpha, heat shock protein cognate 5 (Hsc70-5),
signalrecognition particle protein 54k (Srp54k), 18S, 28S) that had been sampled from 822 mite
species. We used MAFFT (Katoh et al. 2009) to align sequences from each gene. We used
Gblocks (Talavera and Castresana 2007) to purge hyper-variable regions from each ribosomal
alignment. In each Gblocks run, we set the allowed gap position to half, the minimum block length
75
to 5, and the maximum number of contiguous non-conserved positions to 12. We then used
Mesquite v2.73 (Maddison and Maddison 2001) to delimit codon positions and delete introns in
protein-coding alignments, as well as build a supermatrix of concatenated alignments. The total
length of the supermatrix was 12,132 sites. At this stage, we removed taxa from the phylogenetic
dataset that were not represented in the genetic / sexual system trait dataset. This was necessary
to make phylogeny and divergence time estimation feasible. The intersection of the phylogenetic
and trait datasets included 560 taxa. We divided the pruned supermatrix into six data partitions:
first and second nuclear codon positions, third nuclear codon positions, first and second
mitochondrial codon positions, third mitochondrial codon positions, nuclear ribosomal sites, and
mitochondrial ribosomal sites. We used BEAST v1.7.5 (Drummond and Rambaut 2007) to
estimate phylogenetic relationships and time-proportional branch lengths. The BEAST analysis
estimated the parameters of a HKY + G nucleotide substitution model independently for each of
the six data partitions. It used a birth-death model of phylogenetic branching, and an uncorrelated
lognormal relaxed clock model of among-branch substitution rate variation (Drummond et al.
2006). We calibrated divergence times with three fossil-based, exponential node priors: 1) a
minimum age of 380 MY on the stem node of Acariformes (Hirst 1923; Norton et al. 1988); 2) a
minimum age of 90 MY on the stem node of Argasidae (Klompen and Grimaldi 2001); and3) a
minumum age of 35 MY on the stem node of Ixodes (Scudder 1885). We ran the BEAST analysis
for 100 million generations and sampled trees once every 10,000 generations. We examined log
files in Tracer (Drummond and Rambaut 2007) and determined that MCMC sampling from the
stationary distribution commenced after 70 million generations. We randomly selected 100 trees
from the stationary distribution. To account for phylogenetic uncertainty, we repeated our test
over this set of 100 trees.
To maximize overlap between our trait database and phylogeny for comparative
approaches we used an iterative tip matching approach. Briefly, we built our trait matrix by first
finding exact species matches between the taxa in our tree and database. Each genus present
on our tree that had no species level matches was then collapsed to a single tip and we searched
76
for trait data for the genus. This process was repeated at the family level as well. At all
taxonomic levels if there was more than one record in our trait database that matched a tip we
used the mean chromosome number for the tip; in each of these cases the discrete traits such as
reproductive mode were the same for all matching records in the database.
Phylogenetic ANOVA
The phylogenetic ANOVA was calculated using the implementation in the R package
phytools (Revell 2012). Briefly, each taxon in our dataset was classified as haplodiploid or
diplodiploid. If more than one chromosome number was reported for a taxon the mean
chromosome number was used. Significance of the calculated F statistic was assessed using the
simulation approach (Garland et al. 1993). This test was repeated for each of the sampled trees.
Originating condition test
Our test was conceptually inspired by Maddison’s concentrated change test (Maddison
1990). Broadly, our approach determines whether there is evidence that nodes subtending the
origins of the derived state (haplodiploidy) have extreme values for the continuous trait
(chromosome number). We refer to this as the originating condition test and the four steps it
involves are described below:
1) Reconstruct ancestral states for chromosome number pruning data from those species
that exhibit haplodiploidy (fig 4.1a). By pruning chromosome number of haplodiploid taxa we
create a more conservative test that will be applicable even if transitions to haplodiploidy causes
selection for lower chromosome number.
2) Next we reconstruct the evolution of haplodiploidy with stochastic mapping to build a
distribution of possible evolutionary histories (fig 4.1b) (Huelsenbeck et al. 2003). Since there is
strong evidence that haplodiploidy is a derived state we fix the root of the tree as diplodiploid and
we allow for different rates of transition between these states.
3) We then process the stochastic mappings to classify all nodes in the tree. Some
nodes will be haplodiploid (Figure 4.1c, red nodes). The remainder will be diplodiploid and are
subdivided by whether the immediate daughter branches remain in diplodiploidy (Figure 4.1c,
77
blue nodes) or as having an immediate daughter branch that shows a transition into haplodiploidy
(Figure 4.1c, green nodes). Since these green nodes represent the origin of a haplodiploid
lineage we call these nodes “producing nodes”. Next we calculate the mean of chromosome
number at the producing nodes identified across stochastic mappings and if available a
distribution trees. This value is the estimate of the mean chromosome number of nodes that
produce haplodiploidy we refer to this value as the “observed originating condition” for
haplodiploidy.
4) To construct a null distribution we randomly sample from nodes that we have classified
as being diplodiploid (step 3; Figure 4.1c blue and green nodes). The number of nodes sampled
is the same as the number of producing nodes identified in step 3. This is repeated across
multiple stochastic mappings and multiple trees if available. Finally, we use this null distribution
to calculate an empirical p-value for the mean observed originating condition of haplodiploidy
(Figure 4.1d).
78
Figure 4.1 Four steps in the originating condition test.
A) Ancestral state reconstruction of chromosome number assuming a Brownian motion model of
evolution. B) Identification of the transition points in ploidy through stochastic mapping C)
Categorization of nodes as either diplodiploid (blue and green) or haplodiploid (red) as well as
those that subtend an origin of haplodiploidy (green). D) Depiction of the null distribution and the
observed mean chromosome number of producing nodes estimated from the data.
79
Validation testing
To test our approach we first simulated 200 trees using a birth death model with a birth
rate of 1.0 and death rate of 0.5 allowing the trees to grow until 200 extant species were reached.
Trees were simulated using the function sim.bdtree in the R package Geiger (Harmon et al.
2008). We then simulated chromosome number evolving by Brownian motion with a rate
parameter of 0.2 and a starting mean of 10.0 on each tree using the function sim.char also in
Geiger (Harmon et al. 2008). To create datasets where there is a correlation between
chromosome number and the evolution of haplodiploidy we selectively scaled portions of the
trees which had exceptionally low or high chromosome number. Briefly, we created ten versions
of each tree with transformed branch lengths. We identified those branches that had a mean
chromosome number in the upper or lower quartile. Branches in the upper quartile were scaled
by a factor of 1/x while branches in the lower quartile where scaled by a factor of x, and we
repeated this process for ten values of x ranging from 1 to 10. Since our trees begin in
diplodiploidy increasing branch lengths in portions of the tree with low chromosome number
effectively increases the rate of transition into haplodiploidy. While shortening branch lengths
reduces the rate of transition into haplodiploidy. We evolved ploidy on these scaled trees using
the sim.char function and setting the root state diplodiploid and allowing the trait to evolve under a
model where transitions to haplodiploidy were allowed but no back transitions. We evaluated a
number of rates for the transition from diplodiploid to haplodiploid; we discovered that a rate of
0.02 was sufficient to insure at least one transition occurred in all simulated datasets.
When the scaling factor (x) = 1 the trees were unchanged and there was no relationship
between the evolution of chromosome number and ploidy, allowing us to test the type I error rate
of our method. The remaining nine sets of 200 trees each allowed us to evaluate power under an
increasingly strong relationship between chromosome number and ploidy, where branches that
had low chromosome number were more likely to experience a transition into haplodiploidy while
branches inferred to have high chromosome number were less likely to experience a transition.
80
Below we show the performance of this approach with the simulated data, reporting both type I
error and power. Tests were considered significant at a p-value ≤ 0.05.
Results
Phylogenetic ANOVA
Raw data for chromosome number in diplodiploid and haplodiploid Acari suggests that
haplodiploid lineages have lower chromosome number. The mean diploid number in diplodiploid
taxa was 20 while haplodiploid taxa had a mean diploid number of 7. To test whether this
difference could be explained by the correlation structure of the phylogeny we used a
phylogenetic ANOVA. The mean F-statistic calculated across all sampled trees was 97.8. In all
cases simulations indicate that the F-statistic calculated on each phylogeny had a p-value less
than 0.01. This is indicates that the chromosome number in haplodiploid and diplodiploid clades
is significantly different.
Validation testing for originating condition test
Analysis of the 200 simulated datasets with a scaling factor of 1 (no relationship between
traits) showed a type one error rate of 5%. Analysis of the datasets with the scaling factor
ranging from 2-10 assess the power of this approach. We found that under the simulation
condition our approach had a power ranging from 15% to 76%.
81
Figure 4.2 Empirical p-values calculated for simulated datasets.
The X-axis indicates the scaling factor used to stretch trees. The values at the top of the plot
indicate the percentage of simulated datasets that resulted in significant results under our test.
Each column contains 200 results.
Chromosome number and haplodiploidy
Mapping sex determination mode to the sampled Acari trees indicates that transitions to
haplodiploidy occurred between 7 and 12 times (mean=11.2). Maximum likelihood
reconstructions of chromosome number using only data from diplodiploid taxa indicated the mean
of the diplodiploid nodes that led to transitions to haplodiploidy across all trees was 18.5. The null
distribution generated by randomly sampling the same number of diplodiploid nodes on each tree
as there were origins of haplodiploidy had a mean of 20.19 and a standard deviation of 0.78.
Only 19 of the 1000 Monte Carlo simulations produced means equal to or lower than estimated
from the observed data (Figure 4.3; empirical p-value = 0.019). This indicates that transitions to
82
haplodiploidy occur from ancestors with significantly fewer chromosomes than we would expect
0.3
0.2
0.0
0.1
Density
0.4
0.5
by chance.
17
18
19
20
21
22
23
Chromosome Number
Figure 4.3 Originating condition test results.
The red line indicates the mean estimated chromosome number in nodes that subtend an origin
of haplodiploidy. The black line indicates the distribution of simulated means produced to
calculate the empirical p-value.
Discussion
In this study we present two comparative tests of the evolutionary dynamics of
haplodiploidy. We first show that a phylogenetic ANOVA supports an absolute difference in the
chromosome number of haplodiploid and diplodiploid taxa. However this difference in
chromosome number may have evolved as a result of the transition to haplodiploidy rather than
prior to the transition to haplodiploidy. The ancestral condition test that we develop avoids this
possiblility by incorporating chromosome number data only from taxa in the ancestral condition
(diplodiploidy). We show that this approach has acceptable type one error rates but has relatively
weak power, requiring a strong relationship between the continuous trait (chromosome number)
83
and discrete trait (ploidy) (figure 4.2). While we present this method with a trait that does not
exhibit reversions, it should be straightforward to extend to traits that evolve more quickly and
experience reversions. However, we anticipate that such an extension would likely be even less
powerful since the uncertainty in the ancestral reconstruction of the discrete trait will increase.
Increasing uncertainty in identifying the nodes that subtend a transition in the discrete trait will
lead to broader sampling of nodes producing the originating value, and will lead to a lower
probability of significant results. The validation testing shows that this approach is unlikely to
produce false positives and is a conservative test.
Our analysis of the Acari data indicate that transitions to haplodiploidy are concentrated
in lineages with low chromosome number. This provides the first emperical support for Bull’s
hypothesis that haplodiploidy does arise more frequently in clades with fewer chromosomes, and
suggests that taxa with low chromosome number may have a lower viability cost when
transitioning from diplodiploidy to haplodiploidy than taxa with many chromosomes.
In this chapter we have shown that genome architecture can affect the evolution of
haplodiploidy. In doing so we were able to predict, at least in part, the phylogenetic distribution of
ploidy within mites. However the combination of male heterogametic sex determination and low
chromosome numbers occur across large clades of invertebrates that do not appear to have
evolved haplodiploidy. This suggests that these factors alone are insufficient to explain broader
phylogenetic patterns of haplodiploidy. Further theoretical and comparative work will be
necessary to determine additional evolutionary correlates of haplodiploidy, to fully unravel its
evolution.
* Portions of this chapter are in preparation for submission as:
Blackmon, H., Ross, L., Hardy, N.B. (2015) Evolutionary dynamics of haplodiploidy.
Blackmon, H., Adams, R.A. (2015) EvobiR an R package for comparative analysis and teaching
in evolutionary biology.
84
Chapter 5
Recombination, chromosome number and eusociality in the Hymenoptera.
Abstract
Extraordinarily high rates of recombination have been observed in some eusocial
species. The most popular explanation is that increased recombination increases genetic
variation among workers, which in turn increases colony performance, for example by increasing
parasite resistance. However, support for the generality of higher recombination rates among
eusocial organisms remains weak, due to low sample size and a lack of phylogenetic
independence of observations. Recombination rate though difficult to measure directly is
correlated with chromosome number. As predicted, several authors have noted that chromosome
numbers are higher among the eusocial species of Hymenoptera (ants, bees and wasps). Here,
we present a formal comparative analysis of karyotype data from 1567 species of Hymenoptera.
Contrary to earlier studies, we find no evidence for an absolute difference between chromosome
number in eusocial and solitary species of Hymenoptera. However, we find support for an
increased rate of chromosome number change in eusocial taxa. Our results support the view that
a eusocial lifestyle has either led to variable selection pressure on chromosome number or that
eusocial taxa typically have lower effective population size and fix more changes in chromosome
number that are likely mildly deliterious at least while hetrozygous.
Introduction
Sexual reproduction is near ubiquitous among multicellular life (Maynard-Smith, 1978;
Bell, 1982). Recombination -- the reshuffling of genomes during meiosis-- is thought to be the
main benefit of sex, as it increases the efficiency of selection (Otto, 2009). However, in the short
term it can also reduce organismal fitness by breaking up coadapted gene complexes (Agrawal,
2006; Otto, 2009). Theories for the evolution of recombination therefore aim to reconcile these
two opposing evolutionary forces (Otto & Lenormand, 2002). In each generation genetic variation
is produced by two separate mechanisms: independent assortment - random segregation of
85
homologous chromosomes during meiosis and crossing over between homologous
chromosomes. Interestingly, observed recombination rates can vary dramatically, even between
closely related species (White, 1973; Wilfert et al., 2007; Smukowski & Noor, 2011). Theory has
often been successful in predicting under which circumstances selection would increase
recombination rates, but statistical tests of these theories remain scarce (see however
Lenormand & Dutheil, 2005).
One factor that might promote high rates of recombination is eusociality. In eusocial
societies workers forego their own reproduction in order to help raise their siblings. The resulting
colonies can range in size from a few individuals to millions, most of them sterile workers that are
highly related, usually full- or half-siblings. High relatedness is thought to be crucial for the
evolution of eusociality (Boomsma, 2009), but high relatedness can also be problematic for those
societies. Living in a dense aggregate of close kin makes eusocial populations vulnerable to
parasites (Schmid-Hempel, 1998; Kraus & Page, 1998; Schmid-Hempel & Crozier, 1999; WilsonRich et al., 2009). Indeed, there is strong empirical evidence from ants, honeybees and
bumblebees that colonies with higher genetic diversity, are better able to resist parasites (Shykoff
& Schmid-Hempel, 1991; Baer & Schmid-Hempel, 1999; Tarpy, 2003; Hughes & Boomsma,
2004). Genetic diversity might also be important for division of labour (Oldroyd & Fewell, 2007;
Wilfert et al., 2007). Many eusocial colonies display extreme phenotypic and behavioural diversity
among and within castes. Empirical studies have shown that caste, as well as task specialization
within castes, are partly genetically determined (reviewed in Oldroyd & Fewell, 2007; Schwander
et al., 2010). Therefore, low genetic diversity might reduce colony fitness by disrupting proper
division of labour. In addition to the challenges posed by high relatedness, eusocial species are
also challenged by the decrease in effective population size due to extreme reproductive skew
(Kent & Zayed, 2013). As a result, strong linkage disequilibrium and an increased frequency of
deleterious mutations might lead to Hill-Robertson interference (Hill & Robertson, 1968), reducing
the efficiency of natural selection. Several authors have suggested that eusocial species may be
selected to increase recombination rates in responses to these challenges (Schmid-Hempel,
86
1998; Gadau et al., 2000; Wilfert et al., 2007; Sirviö et al., 2011; Gadau et al., 2012; Kent &
Zayed, 2013). Increased recombination could increase genotypic diversity within colonies thereby
helping eusocial societies to resist parasites and maintain proper division of labour (Wilfert et al.,
2007; Oldroyd & Fewell, 2007). Increased recombination rates also increase the efficiency of
selection - counteracting the effects of small effective population sizes that are built into eusocial
societies (Kent & Zayed, 2013).
Recent analyses comparing molecular estimates of recombination between eusocial and
solitary Hymenoptera found that recombination rates of eusocial Hymenoptera are indeed higher
than in solitary hymenopterans or any other metazoan that has been measured (Wilfert et al.,
2007; Sirviö et al., 2011). Unfortunately, these analyses were based on just a small number of
taxa (6 eusocial and 4 solitary Hymenoptera) and did not control for phylogenetic nonindependence. Measuring recombination rates is challenging and labour intensive (Stumpf &
McVean, 2003; Smukowski & Noor, 2011). Therefore, it is unlikely that, in the near future,
sufficient data will be available for rigorous tests of the theoretical impacts of eusocial systems on
recombination rates. On the other hand, currently it is possible to examine the relationship
between eusociality and a factor that is known to be correlated with recombination rate:
chromosome number. In fact, most earlier theory on recombination rate evolution in eusocial
species was based on comparisons between chromosome numbers in eusocial and solitary
Hymenoptera species (Sherman, 1979; Seger, 1983). As mentioned above, genetic variation is a
function of the independent assortment of chromosomes, and the number of crossing over events
between chromosomes. Increases in the number of chromosomes increase the possible
genotypes due to independent assortment during meiosis. Because the number of crossing-over
events is roughly constant (1-2 chiasmata) per chromosome (White 1973) increases in
chromosome number also affects recombination rate by increasing the total number of crossover
events.
Here we perform phylogenetically-controlled analyses to compare chromosomes
numbers among eusocial and solitary species. Although the theory on recombination rates in
87
Hymenoptera was developed to explain absolute differences in chromosome number, it can also
be used to predict differences in the variance in chromosome number between solitary and
eusocial species. Eusocial species vary in terms of their size, mating systems, and social
complexity. Each of these factors is expected to be important in shaping the evolution of
recombination rates. Therefore, we predict that the variance in recombination rates -- as
evidenced by chromosome number -- will be greater in eusocial species. As previously
mentioned, the size of eusocial societies ranges from a few to millions of individuals. Kent and
Zayed (2013) predicted that larger colonies are under stronger selection to increase
recombination, due to an increase of reproductive skew. Larger colonies are also more likely to
suffer from parasites (Schmid-Hempel 1998) and be faced with the maintenance of more
elaborate caste structures than smaller ones (Oldroyd & Fewell, 2007). We therefore expect
larger colonies to benefit more from increased genotypic diversity. Indeed Schmid-Hempel (1998)
found a positive relationship between chromosome number and colonies size across 58 ant
species. Recombination rates might also be affected by another factor that varies among eusocial
societies: polyandry (multiple mating) and polygyny (multiple queens per nest). Both reduce
reproductive skew, thereby increasing effective population size and decreasing selection on
recombination. They also increase genotypic variation within colonies and might therefore lead to
less stringent selection to increase recombination. Finally, eusocial taxa differ in their social
complexity. Eusociality can be facultative, or obligate, and among eusocial species the number of
distinct castes varies. It is currently unclear how the differences between facultative and obligate
eusociality might affect selection on recombination rates. However, the theory on genetic caste
determination predicts that species with more castes should benefit more from increased
genotypic diversity (Oldroyd & Fewell, 2007). Furthermore, colonies with many distinct castes
might be selected to increase recombination as they could benefit from breaking up linkage
between genes that are selected in opposite directions in different castes (Kent & Zayed, 2013).
In summary, we expect eusocial species to have a higher variance in chromosome
number than solitary species, because eusocial species are variable for a number of life history
88
parameters that are expected to affect recombination rate evolution and that are not applicable to
solitary species. Changes in chromosome number are often slightly deleterious and if effective
population size is reduced in eusocial lineages chromosome number should change more quickly
than in solitary species (Lande 1985; Max 1995).
Methods
Data collection
The data used for this analysis were collected from the literature between January 2012
and September 2013. We used a variety of sources, including the primary literature as well as a
number of key review papers. References were identified via Web of Science and Google
Scholar. We collected all karyotype data that has been published for species of Hymenoptera;
these data are available from the Tree of Sex Database (Tree of Sex Consortium, 2014)
(Appendix A and E). For each species with karyotype data, we recorded eusocial status
according to the definition by (Crespi & Yanega, 1995)(solitary, cooperative breeder, facultatively
eusocial, obligately eusocial, with the latter two being considered “eusocial”), and for each
eusocial species we recorded colony size, queen number, and mating numbers for single queens.
Genus-level caste number estimate for ants were based on data collected by Oster & Wilson
(1978). Several ant species are social parasites -- they do not produce any workers themselves,
but rely on those of other species. We obtained social parasite status from a recent review
(Buschinger, 2009), restricting social parasites to those species with either dulosis (slave
capturing) or inquilinism (living in a host species nest). In total we have data for 1567 species,
although the character matrix is not complete for every species. We provide all data Table S1.
Phylogeny
We estimated time-scaled phylogenetic relationships among Hymenoptera lineages using
a phyloinformatic approach. We used a Python script to download published DNA sequences
from the NCBI nucleotide database (GenBank) that were sampled from species included in our
trait dataset. We targeted nine phylogenetic markers that have been used extensively in
Hymenoptera phylogenetics: CAD, abdominal A, arginine kinase, elongation factor 1-alpha, long89
wavelength rhodopsin, wingless, COI, cytB, and the mitochondrial large ribosomal subunit 16S.
We aligned sequence clusters with MAFFT (Katoh & Toh, 2008), and filtered the 16S alignment
using Gblocks (Talavera & Castresana, 2007) to remove ambiguously aligned regions. We used
Mesquite v2.75 (W. P. Maddison & D. R. Maddison, 2013) to concatenate alignments, delete
introns and delimit codon positions. The final data matrix dimensions were 602 taxa by 5600
aligned sites. We estimated phylogenetic relationships and divergence times simultaneously
using BEAST v1.7.5 (Drummond & Rambaut, 2007). In the BEAST analysis, we estimated
nucleotide substitution model parameters independently across five partitions: nuclear codon
positions 1+2, nuclear codon position 3, mitochondrial codon positions 1+2, mitochondrial codon
position 3, and the mitochondrial ribosomal positions. We assumed an HKY site model with
among-site rate variation modeled with a gamma distribution, a birth-death model of phylogenetic
branching, and a log-normal relaxed clock model of among-lineage substitution rate variation. We
calibrated divergence time estimates with three exponential node priors: 1) 185 MY on the stem
node of Ichneumonoidea (Zessin, 1981), 2) 180 MY on the crown node of Tenthredinoidea
(Geinitz, 1887; Nel et al., 2004), and 3) 197 MY on the crown node of Vespomorpha (Heer,
1865). We ran the BEAST analysis for 50 million generations, discarding the first 42 million
generations before the stationary distribution of parameter values was reached.
Comparative analysis
For the analysis of chromosome number evolution we used haploid chromosome count.
In cases where more than one record was available we used the mean of all records. As
predictors we included eusociality (solitary vs. eusocial) as a binary trait, or degree of sociality
(0=solitary, 1= cooperative breeder, 2= facultative eusocial, 3= obligate eusocial) We combined
our data on polygyny and polyandry into one binary variable “relatedness”: species with one or
the other (or both) were scored as “low relatedness” whereas singly mated monogynous colonies
were scored as “high relatedness”. This decision was based on a previously observed negative
correlation between the two (Hughes et al., 2008). We conducted a phylogenetically corrected
90
one-way ANOVA (sensu Garland et al., 1993) comparing eusocial and solitary Hymenoptera. pvalues were calculated based on a null distribution generated from 1000 simulations.
In addition to differences in mean chromosome number, eusociality might also affect the
rate at which chromosome number evolves. We tested for a shift in the rate of chromosome
number evolution using a censored rate test, based on a Brownian motion model. This allows us
to compare models where the continuous trait (chromosome number) evolves at a single rate on
all branches to a model where each state (e.g. solitary and eusocial) has an independent rate of
evolution (O'Meara et al., 2006). Conducting the censored rate test requires a reconstruction of
the history of eusociality on our tree. Since eusociality is widely accepted as a derived state within
Hymenoptera we fixed the root state of the tree as solitary (Wilson, 1975). We used an MK (a
continuous-time Markov chain) model to estimate the parameters of the transition rate matrix, and
allowed different transition rates between states. Stochastic mapping was used to assign the
state of all branches in the tree. To account for uncertainty in ancestral states we performed our
analysis across 100 stochastically mapped trees. This analysis was repeated coding social state
into four categories (solitary, cooperative breeder, faculatively eusocial and obligately eusocial)
and two categories (solitary and eusocial). The R package Phytools version 0.3-72 was used to
both reconstruct ancestral states and fit models of chromosome number evolution. Tests were
considered significant at α=0.05.
Results
Eusociality-depended differences in chromosome number
Figure 5.1 shows a plot of the ancestral state reconstruction of chromosome number
among the 367 Hymenoptera species represented on our phylogeny. We used a phylogenetically
corrected one-way ANOVA to test for an absolute difference in chromosome number between
eusocial and solitary Hymenoptera. The phylogenetically corrected one-way ANOVA comparing
eusocial and solitary Hymenoptera also revealed no significant difference in chromosome number
(F-statistic = 70.5, p-value = 0.23).
91
Rates of chromosome number evolution
Karyotypes vary dramatically within and among the eusocial clades, with chromosome
number ranging from n=1 in the ant Myrmecia croslandi to n=53-60 in the ant Dinoponera lucida
(figure 5.1a). Here we test if this variation could be due to an increased rate of evolution of
karyotypes in eusocial compared to solitary clades. The censored rate test supports the
conclusion that chromosome number evolves more quickly in eusocial than solitary Hymenoptera.
We performed the censored rate test coding taxa into four eusocial states solitary, cooperative
breeder, facultatively eusocial, and obligately eusocial. A four-rate model where each social state
has its own rate was preferred across all stochastic mappings. The rate in the eusocial
(facultative and obligate) clades was ~3x faster than the solitary clades (1.28 and 1.10 vs. 0.374
respectively). We estimated the highest rate (7.69) in the cooperative breeder clade, but this
group is only represented by four species on the tree, and the 95% confidence interval (2.0113.37) indicated insufficient data to reliably estimate this rate. We repeated this analysis coding
taxa as eusocial or solitary. Using this coding we again found support for the more complex
model where each social state has its own rate of evolution. This two-rate model was preferred
across all 100 stochastic mappings with a p-value of less than 0.001. The rate of chromosome
number evolution in eusocial clades was again ~ 3x faster (3.17-3.25x faster across stochastic
character mappings) than in solitary clades (Figure 2b).
92
Figure 5.1 Analysis of the rate of chromosome number evolution.
In both trees the ring outside of the tree indicates the sociality state of the terminal taxa, black
indicating eusocial and gray indicating solitary. a) The branches have been painted to illustrate a
reconstruction of the evolution of chromosome number under Brownian motion according to the
scale at lower left. b) A single stochastic mapping from our analysis shows how the branches
were assigned to either eusocial or solitary states. Values for σ2 are the mean across 100
stochastic mappings, and represent the rate of chromosome number evolution.
Discussion
The observation of extraordinarily high rates of recombination in a number of eusocial
Hymenoptera has spurred the development of theories to explain the evolution of recombination
rates. Broadly speaking, these theories fall into two classes: Either eusociality leads to selection
for high recombination rates to increase genotypic diversity within a colony (“the genetic diversity
93
theory” Wilfert et al., 2007), or alternatively to alleviate the negative effects of reproductive skew
and small effective population size (“the reproductive skew theory”, Kent & Zayed, 2013).
Although the theoretical validity of these models is clear, the empirical support for a causal link
between eusociality and high recombination remains weak. In this manuscript we test the
hypothesis that eusocial behaviour has affected selection on recombination rates in the
Hymenoptera, using chromosome number as a proxy for recombination. Using phylogenetic
framework we show that, in contrast to earlier studies (Sherman, 1979; Seger, 1983; Wilfert et al.,
2007), there is no support for a higher number of chromosomes among eusocial species. We
believe that this is unlikely to be an issue of statistical power, as our study is the most
comprehensive to date and captures four out of the nine origins of eusociality. A more likely
explanation is that the differences in chromosome number/recombination rate found by earlier
analyses were due to the phylogenetic non-independence of observations; the principal
difference observed was between the Aculeata (which consist of both solitary and eusocial
species and has n=16 chromosomes on average), and the rest of the Hymenoptera (with an
average of n=10 chromosomes).
Eusocial societies are diverse. Theory predicts that selection for increased recombination
rates depends on many aspects a species’ ecology (Wilfert et al., 2007; Sirviö et al., 2011; Kent &
Zayed, 2013). We therefore decided not just to test the absolute difference between the
chromosome number of solitary and eusocial species, but also to consider the rate at which
chromosome number has evolved. Using a phylogenetic framework we found that chromosome
number indeed changes at a higher rate in eusocial hymenopterans.
Much of the published theory for recombination rate and its effect on genetic variation
were coopted from theories for the evolution of polyandry. There is a wealth of empirical support
for the importance of polyandry on within-colony diversity. However, it is unclear if these theories
make the same predictions about recombination. From the quantitative genetics literature we
know that additive genetic variance decrease with recombination rate (Falconer & Mackay, 1996).
Another issue is that for example parasite resistance could be due to a single locus, in which
94
case recombination rate would have no effect on the distribution of resistance among colony
members. The same would be true if the trait if polygenic, but the effects of each gene are
completely additive (Falconer & Mackay, 1996). A recent simulation study investigated the role of
chromosome number and recombination rate on polygenic traits (Rueppell et al., 2012). They
showed that both processes in general do not affect genetic variance, but that they can increase
the number of unique genotypes and the genotypic range in a colony (Rueppell et al., 2012).
However the effects are small, and we currently know too little about the genetic architecture of
traits determining colony fitness to access the impacts of these effects.
In this study we used chromosome number as a proxy for recombination rate. This
means that although we were able to utilize data from a larger sample of eusocial species, we
were working with only a coarse approximation of recombination rate. For example, it is possible
that recombination rate varies by means other than chromosome number, e.g. an increase of the
number of chiasmata per chromosome. Unfortunately most karyotypic studies of the
Hymenoptera do not cite the number of chiasmata (Gokhman, 2009). However Hymenoptera
have relatively small chromosomes and it is therefore unlikely that more than one chiasmata is
present per arm.
In conclusion, a theoretical link has been made between high rates of recombination,
high levels of genetic diversity, effective population size and colony performance in eusocial
species. Higher genotypic diversity within colonies could improve their performance, by
increasing resistance to parasites, and / or maintaining the genetic basis of the division of labour.
While at the same time high recombination rates can reduce linkage and interference effects
resulting from high reproductive skew. In this study, we tested these predictions using a large
comparative analysis of Hymenoptera that assumed recombination rate is correlated with
chromosome number. We found that eusociality is associated with an increased rate of
chromosome number evolution but no absolute difference in chromosome number. While the
higher rate of chromosome number evolution observed in eusocial clades may indicate varying
strengths of selection on chromosome number it may also be due to drift. Changes in
95
chromosome number are often slightly deleterious (Max, 1995). Therefore if effective population
size is reduced in eusocial lineages chromosome number should change more quickly due to
drift. However this is based on the assumption that eusociality always reduces effective
population size, which fails to take into account the ecological success of many eusocial species.
The current theory on recombination rate evolution in eusocial invertebrates fails to evaluate the
relative importance of linkage disequilibrium and colony genotypic diversity and thereby provide
clear quantitative predictions for both short-term and long-term effects of recombination rate.
Such theoretical advances, in addition to recombination rate estimates for more Hymenoptera
species, especially solitary members of the Aculeata, gamergate colonies and social parasites,
will be crucial to fully understand the processes that have produced the pattern of variation that
we have observed.
* Portions of this chapter were previously published as:
Ross, L., Blackmon, H., Lorite, P., Gokhman, V. E. and Hardy, N. B. (2015), Recombination,
chromosome number and eusociality in the Hymenoptera. Journal of Evolutionary Biology.
doi: 10.1111/jeb.12543
96
Appendix A
Cytogenetic data for invertebrates
97
Taxa
Acariformes
Acaridae
Acarus siro
Acotyledon formosani
Caloglyphus berlesei
Caloglyphus michaeli
Caloglyphus mycophagus
Rhizoglyphus echinopus
Rhizoglyphus robini
Sancassania berlesei
Sancassania michaeli
Sancassania mycophaga
Sapracarus sp.
Sapracarus tuberculalUs
Schwiebea elongata
Troglocoptes sp.1
Troglocoptes sp.2
Troglocoptes sp.3
Troglocoptes sp.4
Tyrophagus casei
Tyrophagus neiswanderi
Tyrophagus palmarum
Tyrophagus putrescentiae
Achipteriidae
Achipteria coleoptrata
Achipteria punctata
Achipteria species
Anachipteria species
Anystidae
Anystis baccarum
Anystis salicinus
Arrenuridae
Arrenurus bicuspidator
Arrenurus caudatus
Arrenurus maculata
Arrenurus pustulator
Arrenurus sp.1
Arrenurus sp.2
Arrenurus sp.3
Arrenurus sp.4
Arrenurus sp.5
Astegistidae
Cultroribula bicultrata
Cultroribula divergens
Atopochthoniidae
Atopochthonius artiodactylus
Bdellidae
Bdella capitosa
Bdella tropica
Brachychthoniidae
Brachychthonius berlesei
Brachychthonius pius
Eobrachychthonius latior
Liochthonius brevis
Liochthonius sellnicki
Liochthonius strenzkei
Poecilochthonius spiciger
Sellnickochthonius immaculatus
Sellnickochthonius lydiae
Sellnickochthonius suecia
Sellnickochthonius zelawaiensis
Synchthonius crenulatus
Camerobiidae
Neophyllobius aesculi
haplodiploid type unspecified
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
XO
18
16
16
10
XO
XO
XO
XY
18
16
16
XO
XO
XO
10
12
16
16
XY
XY
XO
XO
parth
parth
parth
parth
parth
parth
parth
parth
18
parth
parth
211
212
211
211
211
213
214
213
213
213
213
213
215
213
213
213
213
213
213
213
213
216
213
217
217
213
213
20
26
20
23
20
26
211
211
211
211
213
213
213
213
213
parth
parth
217
217
parth
217
parth
parth
213
213
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
217
217
217
217
213
217
217
217
217
217
217
217
22
98
213
Taxa
Neophyllobius elegans
haplodiploid type unspecified
Neophyllobius piniphilus
haplodiploid type unspecified
Camisiidae
Camisia aff. lapponica
Camisia carrolli
Camisia horrida
Camisia invenusta
Camisia segnis
Camisia spinifer
Heminothrus interlamellaris
Heminothrus longisetosus
Heminothrus ornatissimus
Heminothrus paolianus
Heminothrus targionii
Plarynothrus altimontanus
Plarynothrus banksi
Plarynothrus biangulatus
Plarynothrus bicarinatus
Plarynothrus brevisetosus
Plarynothrus carinatus
Plarynothrus castaneus
Plarynothrus major
Plarynothrus microclava
Plarynothrus peltifer
Platynothrus peltifer
Platynothrus punctatus
Platynothrus septentrionalis
Platynothrus sibiricus
Platynothrus skoettsbergi
Platynothrus thori
Platynothrus traversus
Carabodidae
Carabodes femoralis
Carabodes granulatus
Ceratoppiidae
Ceratoppia bipilis
Ceratoppia species
Ceratozetidae
Ceratozetes cf.
Ceratozetes cuspidatus
Ceratozetes gracilis
Ceratozetes parvulus
Fuscozetes species
Chamobatidae
Chamobates borealis
Chamobates voigtsi
Cheyletidae
Acaropsellina docta
haplodiploid type unspecified
Cheyletus malaccensis
haplodiploid type unspecified
Cheletogenes ornatus
haplodiploid type unspecified
Cheyletus eruditus
Nodele simplex
haplodiploid type unspecified
Cosmochthoniidae
Gozmanyina majesta
Nipponiella sp
Crotoniidae
Austronothrus clarki
Austronothrus curviseta
Austronothrus flagellatus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
213
20
218
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
213
213
213
213
219
219
213
213
219
216
213
213
219
219
219
213
213
213
213
213
213
217
213
213
213
213
213
213
parth
216
213
18
217
217
217
217
217
213
217
parth
parth
220
220
parth
10
213
4
213
4
213
4
4
211
213
parth
parth
213
213
221
221
221
99
Taxa
Crotonia brachyrostrum
Crotonia caudalis
Holonothrus sp.
Cunaxidae
Cunaxa capreolus
haplodiploid type unspecified
Damaeidae
Damaeobelba minutissima
Damaeus angustipes
Damaeus verticillipes
Epidamaeus species
Damaeolidae
Fosseremus laciniatus auct
Elliptochthoniidae
Elliptochthonius profundus
Eniochthoniidae
Eniochthonius minutissimus
Epidermoptidae
Myialges pari
haplodiploid type unspecified
Epilohmanniidae
Epilohmannia cylindrica
Epilohmannia pallida
Epilohmannia styriaca
Epilohmannoides jacoti
Epilohmannoides terrae
Eremobelbidae
Eremobelba gracilior
Ereynetidae
Riccardoella limacum
haplodiploid type unspecified
Eriophyidae
Abacarus hystrix
haplodiploid type unspecified
Aceria ficus
haplodiploid type unspecified
Aceria guerreronis
haplodiploid type unspecified
Aceria oleae
haplodiploid type unspecified
Aceria sheldoni
haplodiploid type unspecified
Aculops cornutus
haplodiploid type unspecified
Aculops fockeui
haplodiploid type unspecified
Aculops lycopersici
haplodiploid type unspecified
Aculops pelekassi
haplodiploid type unspecified
Aculops tetanothrix
haplodiploid type unspecified
Aculus persicae
haplodiploid type unspecified
Aculus schlechtendali
haplodiploid type unspecified
Phyllocoptruta oleivora
haplodiploid type unspecified
Eriophyidae
Artacris macrorhynchus
haplodiploid type unspecified
Epitrimerus pyri
haplodiploid type unspecified
Eriophyes laevis
haplodiploid type unspecified
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
216
216
221
22
parth
18
213
213
217
213
217
parth
213
parth
213
parth
217
16
parth
parth
parth
parth
parth
213
213
213
213
213
213
217
10
213
222
223
224
225
4
213
213
213
4
213
213
4
213
4
213
4
213
4
213
4
213
213
213
100
Taxa
Eriophyes tiliae
haplodiploid type unspecified
Leipothrix dipsacivagus
haplodiploid type unspecified
Rhyncaphytoptus ficifoliae
haplodiploid type unspecified
Tegolophus hassani
haplodiploid type unspecified
Vasates aceriscrumena
haplodiploid type unspecified
Erythraeidae
Balaustium sp.
Erythraeus sp.
Eulohmanniidae
Eulohmannia ribagai
Eupalopsellidae
Saniosulus nudus
haplodiploid type unspecified
Euphthiracaridae
Microtritia minima
Euphthiracaroidea
Rhysotritia ardua
Rhysotritia duplicata
Eupodidae
Claveupodes delicatus
Eupodes sigmoidensis
Eupodes sp.
haplodiploid type unspecified
Hawaiieupodes thermophilus
Linopodes sp.
haplodiploid type unspecified
Eutegaeidae
Eutegaeus curviseta
Euzetidae
Euzetes globulus
Galumnidae
Galumna ithacensis
Galumna sp.
Orthogalumna terebrantis
haplodiploid type unspecified
Pergalumna curva
Pergalumna emarginata
Pergalumna formiria
Gehypochthoniidae
Gehypochthonius rhadamanthus
Gehypochthonius urticinus
Gehypochthonius xarifae
Glycyphagidae
Glycyphagus domesticus
Gymnodamaeidae
Gymnodamaeus bicostaus
Jacotella species
Haplochthonlidae
Amnemochthonius taeniophorus
Haplochthonius simplex
Haplozetidae
Peloribates cf.
Protoribates capucinus
Protoribates lophotrichus
Rostrozetes foveolatus
Xylobates capucinus
Xylobates oblongus
Xylobates robuSlior
Harpirhynchidae
Harpyrhynchus brevis
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
4
226
227
225
223
228
parth
213
213
16
parth
217
6
213
parth
217
parth
parth
213
216
parth
parth
18
213
213
213
18
213
213
parth
216
18
213
18
18
217
213
213
parth
parth
217
217
217
parth
parth
parth
213
213
213
18
XO
211
217
217
parth
parth
213
213
parth
parth
parth
parth
parth
parth
217
217
217
213
213
213
213
4
101
213
Taxa
Sexual
System
Ploidy
Female
2n
haplodiploid type unspecified
Harpyrhynchus novoplumaris
4
haplodiploid type unspecified
Hemisarcoptidae
Hemisarcoptes coccophagus
14
facultative sex ratio adjustment, homomorphic chromosomes
Hermanniellidae
Hermanniella species
Hermanniidae
Hermannia gibba
16
Histiostomatidae
Anoetus laboratorium
8
haplodiploid type unspecified
Hormosianoetus laboratorium
8
haplodiploid type unspecified
Myianoetus sp.
haplodiploid type unspecified
Histiostomatidae
Hexanoetus conoidalis
haplodiploid type unspecified
Histiostoma bakeri
Histiostoma feroniarum
14
haplodiploid type unspecified
Histiostoma feroniarum.2
parth
Histiostoma formosana
haplodiploid type unspecified
Histiostoma humiditatis
parth
Histiostoma julorum
haplodiploid type unspecified
Histiostoma murchei
haplodiploid type unspecified
Humerobatidae
Humerobates rostrolamellatus
16
haplodiploid type unspecified
Hydrachnidae
Hydrachna globosa
12
Hydrachna leegei
20
Hydrachna sp.1
12
Hydrachna sp.2
12
Hydrachna uniscutata
12
Hydrodromidae
Hydrodroma despiciens
6
Hydrodroma despiciens.2
16
likely XO
Hydrozetldae
Hydrozetes dimorphus virginalis parth
Hydrozetes lacustris
parth
Hydrozetes parisiensis
parth
Hydrozetes terrestris
parth
Hydrozetes tridactylus
parth
Hydryphantidae
Hydryphantes bayeri
10
Hydryphantes clypeatus
6
Hydryphantes ruber
12
Hydryphantes sp.
10
Hydryphantes sp.1
10
Hydryphantes sp.2
10
Parathyas dirempta
18
Hygrobatidae
Hygrobates calliger
14
Hypochthoniidae
Eohypochthonius magnus
parth
Eohypochthonius travei
parth
Hypochthonius luteus
parth
Hypochthonius rufulus
18
102
Male
2n
Sex Chrom. Source
Karyotype
213
229
217
213
211
213
213
213
213
213
230
213
213
213
213
213
211
211
213
213
211
213
231
213
213
213
213
213
211
211
211
211
213
213
213
211
213
213
213
213
Taxa
Hypochthonius rufulus.2
Malacoangelia remigera
Labidostommatidae
Labidostomma luteum
Lebertiidae
Lebertia porosa
Lebertia sp.1
Lebertia sp.2
Lebertia stackelbergi
Liacaridae
Liacarus coracinus
Liacarus subterraneus
Limnesiidae
Limnesia maculata
Limnesia sp.1
Limnesia sp.2
Limnesia undulata
Limnocharidae
Limnochares aquatica
Llmnozetidae
Limnozetes amnicus
Limnozetes atmetos
Limnozetes borealis
Limnozetes guyi
Limnozetes latilamellata
Limnozetes lustrum
Limnozetes onondaga
Limnozetes palmerae
Limnozetes sphagni
Lohmanniidae
Annectacarus mucronatus
Cryptacarus promecus
Lohmannia banksi
Lohmannia lanceolata
Meristacarus sp.
Torpacarus omittens
Malaconothridae
Malaconothrus crassisetosa
Malaconothrus gracilis
Malaconothrus hauseri
Malaconothrus robustus
Malaconothrus robustus asiaticus
Malaconothrus sp.1
Malaconothrus sp.2
Trimalaconothrus glaber
Trimalaconothrus novus
Trimalaconothrus saxosus
Trimalaconothrus simplex
Mesoplophoridae
Archoplophora laevis
Archoplophora villosa
Mochlozetidae
Podoribates pratensis
Mycobatidae
Punctoribates insignis
Punctoribates punctum
Nanhermanniidae
Cyrthermalllzia guadeloupensis
Cyrthermalllzia sp.1
Masthermannia sp.1
Nanhermannia comitalis
Nanhermannia coronata
Nanhermannia dorsalis
Nanhermannia elegantula
Nanhermannia nana
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
parth
parth
217
213
parth
213
16
16
18
18
211
213
213
211
220
220
18
18
18
18
211
213
213
211
6
213
parth
parth
parth
parth
parth
parth
parth
parth
parth
213
213
213
213
213
213
213
213
213
parth
parth
parth
parth
parth
parth
213
213
216
213
213
213
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
213
216
213
213
213
217
217
213
213
213
213
parth
parth
217
213
217
parth
213
217
parth
parth
parth
parth
parth
parth
parth
parth
213
213
213
213
216
217
216
216
103
Taxa
Nehypochthoniidae
Nehypochthorum porosus
Neoliodidae
Poroliodes farinosus
Nothridae
Nothrus anauniensis
Nothrus borussicus
Nothrus macedi
Nothrus monticolus
Nothrus palustris
Nothrus pratensis
Nothrus quadripilis
Nothrus silvestris
Nothrus silvicus
Nothrus terminalis carolinae
Nothrus truncatus
Novonothrus flagellatus
Oppiidae
Graptoppia (Stenoppia)
Microppia minus
Multioppia species
Oppia cf.
Oppia nitens
Oppia nodosa
Oppia sp.
haplodiploid type unspecified
Oppiella nova
Oppioid sp.
Quadroppia circumita
Quadroppia quadricarinata
Quadroppia sp.1
Quadroppia sp.2
Quadroppia sp.3
Oribatellidae
Oribatella calcarata
Oribatella quadricornuta
Oribatella sakamorii
Oribatella sp.1
Oribatella sp.2
Oribatulidae
Oribatula sakamorii
Oribatula tibialis
Palaeacaridae
Palaeacarus hystricinus
Palaeacarus kamenskii
Parakalummidae
Protokalumma salicis
Paratydeidae
Neotydeus ardisannae
Parhypochthoniidae
Parhypochrhonius aphidinus
Pedlculochelldae
Paralycus lavoipierrei
Paralycus parvulus
Penthaleidae
Halotydeus destructor
Halotydeus destructor.2
Penthaleus major
Phenopelopidae
Eupelops hirtus
Eupelops plicatus
Eupelops torulosus
Phthiracaridae
Atropacarus striculus
Phthiracarus compressus
Sexual
System
Ploidy
Female
2n
parth
Male
2n
Sex Chrom. Source
Karyotype
216
18
213
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
219
213
213
213
219
217
213
213
217
219
216
216
parth
parth
217
217
217
217
213
213
213
parth
parth
18
parth
217
217
217
213
217
217
217
parth
parth
220
220
213
217
217
parth
parth
213
217
parth
parth
217
213
217
parth
213
parth
213
parth
parth
213
213
parth
parth
232
213
232
220
216
220
parth
217
217
104
Taxa
Phthiracarus setosellus
Steganacarus magnus
Phyllochthoniidae
Phyllochthonius aoutii
Pionidae
Piona camea
Piona coccinea coccinea
Piona nodata
Piona sp.1
Piona sp.2
Piona sp.3
Piona sp.4
Piona uncata uncata
Podapolipidae
Chrysomelobia labidomerae
haplodiploid type unspecified
Podopterotegaeidae
Podopterotegaeus tectus
Pomerantziidae
Pomerantzia benhami
Pomerantzia kethleyi
Pomerantzia prolata
Pterochthoniidae
Pterochthonius angelus
Pterygosomatidae
Geckobiella texana
haplodiploid type unspecified
Pyemotidae
Pyemotes herfsi
haplodiploid type unspecified
Pyemotes scolyti
haplodiploid type unspecified
Pyemotes tritici
haplodiploid type unspecified
Pyemotes ventricosus
haplodiploid type unspecified
Pygmephoridae
Pediculaster mesembrinae
haplodiploid type unspecified
Scutacaridae flechtmanni
haplodiploid type unspecified
Siteroptes graminum
haplodiploid type unspecified
Siteroptes reniformis
haplodiploid type unspecified
Pyroglyphidae
Dermatophagoides farinae
Raphignathidae
Neognathus terrestris
Rhagidiidae
Evadorhagidia oblikensis
Poecilophysis faeroensis
Poecilophysis pratensis
Rhagidia diversicolor
Rhagidia gigas
Robustocheles mucronata
Shibaia longisensilla
Sarcoptidae
Sarcoptes scabiei
Scheloribatidae
Scheloribates laevigatus
Scheloribates lanceoliger
Scheloribates species
Scutacaridae
Imparipes histricinus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
217
233
parth
213
22
20
8
8
20
22
211
211
211
213
213
213
213
211
20
213
parth
217
parth
parth
parth
213
213
213
parth
217
213
213
213
6
213
6
211
6
213
6
213
6
213
6
213
XO
234
parth
213
parth
parth
parth
parth
parth
parth
parth
213
213
213
213
213
213
213
18
XO
235
217
217
217
213
105
Taxa
haplodiploid type unspecified
Stigmocheylidae
Stigmocheylus sp.1
Stigmocheylus sp.2
Suctobelbidae
Allosuctobelba obtusa
Suctobelbella falcata
Suctobelbella hamata
Suctobelbella hurshi
Suctobelbella laevis
Suctobelbella palustris
Suctobelbella similis
Suctobelbella sp.1
Suctobelbella sp.10
Suctobelbella sp.11
Suctobelbella sp.12
Suctobelbella sp.13
Suctobelbella sp.14
Suctobelbella sp.15
Suctobelbella sp.16
Suctobelbella sp.17
Suctobelbella sp.18
Suctobelbella sp.19
Suctobelbella sp.2
Suctobelbella sp.20
Suctobelbella sp.21
Suctobelbella sp.3
Suctobelbella sp.4
Suctobelbella sp.5
Suctobelbella sp.6
Suctobelbella sp.7
Suctobelbella sp.8
Suctobelbella sp.9
Suctobelbella subcornigera
Suctobelbella tuberculata
Suctobelbella vera
Syringophilidae
Syringophiloidus minor
haplodiploid type unspecified
Tarsonemidae
Iponemus confusus
haplodiploid type unspecified
Iponemus radiatae
haplodiploid type unspecified
Polyphagotarsonemus latus
haplodiploid type unspecified
Tarsonemus confusus
haplodiploid type unspecified
Tarsonemus confusus.2
Tarsonemus lobosus
haplodiploid type unspecified
Tarsonemus nodosus
haplodiploid type unspecified
Tarsonemus randsi
haplodiploid type unspecified
Tarsonemus schlechtendali
haplodiploid type unspecified
Tarsonemus sp.
haplodiploid type unspecified
Tarsonemus talpae
haplodiploid type unspecified
Tarsonemus virgineus
Tarsonemus waitei
haplodiploid type unspecified
Tectocepheidae
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
parth
parth
213
213
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
217
6
213
213
213
4
213
213
parth
213
213
213
213
213
4
213
213
parth
213
213
106
Taxa
Tectocepheus cuspidatus
Tectocepheus minor
Tectocepheus sarekensis
Tectocepheus velatus
Tegoribatidae
Lepidozetes singularis
Tenuipalpidae
Brevipalpus califomicus
haplodiploid type unspecified
Brevipalpus obovatus
haplodiploid type unspecified
Brevipalpus phoenicus
haplodiploid type unspecified
Brevipalpus pulcher
haplodiploid type unspecified
Brevipalpus russulus
haplodiploid type unspecified
Brevipalpus spinosus
haplodiploid type unspecified
Oligomerismus oregonensis
Raoiella indica
haplodiploid type unspecified
Tenuipalpus inophylli
Tenuipalpidae
Dolichotetranychus summers
haplodiploid type unspecified
Tetranychidae
Anatetranychus tephrosiae
haplodiploid type unspecified
Bryobia graminum
Bryobia kissophila
Bryobia lagodechiana
Bryobia latens
Bryobia neopraetiosa
Bryobia praetiosa
haplodiploid type unspecified
Bryobia rubrioculus
Bryobia sarcothamni
haplodiploid type unspecified
Eonychus curtisetosus
haplodiploid type unspecified
Eonychus grewiae
haplodiploid type unspecified
Eotetranychus befandrianae
haplodiploid type unspecified
Eotetranychus carpini
haplodiploid type unspecified
Eotetranychus friedmanni
haplodiploid type unspecified
Eotetranychus grandis
haplodiploid type unspecified
Eotetranychus imerinae
haplodiploid type unspecified
Eotetranychus paracybelus
haplodiploid type unspecified
Eotetranychus ranoma fanae
haplodiploid type unspecified
Eotetranychus rinoreae
haplodiploid type unspecified
Eotetranychus roedereri
haplodiploid type unspecified
Eotetranychus sakalavensis
haplodiploid type unspecified
Eotetranychus tiliarium
haplodiploid type unspecified
Sexual
System
Ploidy
Female
2n
parth
parth
parth
parth
Male
2n
Sex Chrom. Source
Karyotype
213
220
220
216
217
parth
2
211
parth
2
211
parth
2
211
4
213
4
213
4
213
4
213
213
parth
parth
parth
parth
parth
parth
parth
parth
213
4
213
6
211
8
236
236
213
213
236
211
8
236
211
4
211
4
211
4
211
8
211
6
211
6
211
6
211
6
211
10
211
6
211
6
211
4
211
8
211
parth
107
Taxa
Eotetranychus tulearensis
haplodiploid type unspecified
Neotetranychus rubi
haplodiploid type unspecified
Obuloides sp.
haplodiploid type unspecified
Oligonychus andrei
haplodiploid type unspecified
Oligonychus bessardi
haplodiploid type unspecified
Oligonychus chazeaui
haplodiploid type unspecified
Oligonychus coffeae
haplodiploid type unspecified
Oligonychus gossypii
haplodiploid type unspecified
Oligonychus monsarrati
haplodiploid type unspecified
Oligonychus pratensis
haplodiploid type unspecified
Oligonychus quercinus
haplodiploid type unspecified
Oligonychus randriamasii
haplodiploid type unspecified
Oligonychus sylvestris
haplodiploid type unspecified
Oligonychus thelytokous
Oligonychus ununguis
haplodiploid type unspecified
Oligonychus virens
haplodiploid type unspecified
Panonychus ulmi
haplodiploid type unspecified
Petrobia harti
haplodiploid type unspecified
Petrobia latens
haplodiploid type unspecified
Porcupinychus insularis
haplodiploid type unspecified
Schizonobia oudemansi
haplodiploid type unspecified
Schizonobia sycophanta
haplodiploid type unspecified
Schizotetranychus australis
haplodiploid type unspecified
Schizotetranychus schizopus
haplodiploid type unspecified
Tenuipalpoides acaciae
haplodiploid type unspecified
Tetranychus atlanticus
haplodiploid type unspecified
Tetranychus cinnabarinus
haplodiploid type unspecified
Tetranychus hydrangeae
haplodiploid type unspecified
Tetranychus kaliphorae
haplodiploid type unspecified
Tetranychus ludeni
haplodiploid type unspecified
Tetranychus neocalendonicus
haplodiploid type unspecified
Tetranychus pacificus
haplodiploid type unspecified
Tetranychus panici
haplodiploid type unspecified
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
4
211
14
211
6
213
4
211
8
211
8
211
6
211
4
211
8
211
8
211
6
211
4
211
4
211
6
213
211
8
211
6
211
4
211
8
211
8
211
8
237
8
237
12
211
6
211
6
213
6
211
6
211
6
211
6
211
6
211
6
211
6
211
8
211
parth
108
Taxa
Tetranychus roseus
haplodiploid type unspecified
Tetranychus tumidus
haplodiploid type unspecified
Tetranychus urticae
haplodiploid type unspecified
Tetranychus viennensis
haplodiploid type unspecified
Tetranycopsis horridus
haplodiploid type unspecified
Thyrisomiidae
Banksinoma ovata
Trhypochthoniidae
Afronothrus giganticus
Afronothrus incisivus
Afronothrus neotropicus
Afronothrus russeolus
Afronothrus schuilingi
Allonothrus giganticus
Allonothrus neotropicus
Allonothrus russeolus
Allonothrus schuilingi
Archegozetes longisetosus
Archegozets magnus
Mainothrus badius
Mucronothrus nasalis
Pseudonothrus hirtus
Trhypochthoniellus badius
Trhypochthoniellus crussus
Trhypochthoniellus excavatus
Trhypochthoniellus setosus
Trhypochthoniellus sp.1
Trhypochthoniellus sp.2
Trhypochthonius americanus
Trhypochthonius nigricans
Trhypochthonius silvestris
Trhypochthonius tectorum
Trichthoniidae
Gozmanyina majestus
Trombiculidae
Leptotrombidium akamushi
Leptotrombidium arenicola
Leptotrombidium deliense
Leptotrombidium fletcheri
Leptotrombidium scutellare
Tydeidae
Tydeus caudatus
haplodiploid type unspecified
Unionicolidae
Neumania vemalis
Neumania vernalis
Unionicola crassipes
Winterschmidtiidae
Czenspinskia transversostriata
Ensliniella kostylevi
Ensliniella parasitica
haplodiploid type unspecified
Kennethiella trisetosa
haplodiploid type unspecified
Kurosaia jiju
Xenillidae
Xenillus tegeocranus
Acariformes
Demodicidae
Demodex caprae
Sexual
System
parth
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
8
211
12
211
6
211
6
211
4
211
217
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
213
219
213
213
213
219
213
213
213
219
213
216
216
213
216
216
216
213
213
213
216
213
213
213
parth
217
12
28
14
14
14
XY
XY
238
213
213
213
238
4
213
4
4
18
211
213
211
parth
213
239
240
213
241
18
213
4
211
109
Taxa
Sexual
System
haplodiploid type unspecified
Eriophyidae
Phytoptus tiliae
haplodiploid type unspecified
Eupalopsellidae
Eupalopsellus brevipilus
haplodiploid type unspecified
Eupalopsellus olearius
haplodiploid type unspecified
Eylaidae
Eylais mutila
Eylais rimosa
Eylais setosa
Eylais sp.1
Eylais sp.2
Oxidae
Frontipoda musculus
Stigmaeidae
Agistemus camerounensis
haplodiploid type unspecified
Agistemus exsertus
haplodiploid type unspecified
Agistemus sanctiluciae
haplodiploid type unspecified
Agistemus tranatalensis
haplodiploid type unspecified
Tarsonemidae
Phytonemus pallidus
haplodiploid type unspecified
Tenuipalpidae
Aegyptobia ephedrae
haplodiploid type unspecified
Aegyptobia sp.
haplodiploid type unspecified
Aegyptobia thujae
parth
Aegyptobia vannus
parth
Cenopalpus lanceolatisetae
haplodiploid type unspecified
Tetranychidae
Eurytetranychus buxi
haplodiploid type unspecified
Eurytetranychus madagascariensis
haplodiploid type unspecified
Eutetranychus banksi
haplodiploid type unspecified
Eutetranychus eliei
haplodiploid type unspecified
Eutetranychus grandidieri
haplodiploid type unspecified
Eutetranychus orientalis
haplodiploid type unspecified
Eutetranychus ranjatori
haplodiploid type unspecified
Eutetranychus sambiranensis
haplodiploid type unspecified
Trombidiidae
Allothrombium fuliginosum
Sericothrombium schar/atinum
Sericothrombium sp.1
Sericothrombium sp.2
Sericothrombium sp.3
Tydeidae
Homeopronematus anconai
haplodiploid type unspecified
Blattodea
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
4
213
8
213
6
213
6
4
4
4
6
211
211
211
213
213
19
211
4
213
6
213
4
213
6
213
4
213
4
213
4
213
4
4
4
242
242
213
10
211
6
211
6
211
8
211
4
211
6
211
6
211
4
211
24
18
213
211
26
22
18
211
211
213
213
110
Taxa
Blaberidae
Archimandrita tessellata
Blaberus atropos
Blaberus cranifer
Blaberus discoidalis
Blaberus giganteus
Blaberus parabolicus
Byrsotria fumigata
Capucina patula
Diploptera punctata
Epilampra abdomennigrum
Epilampra grisea
Epilampra maculicollis
Epilampra maya
Epilampra sagitta
Epilampra sp.81B
Eublaberus distanti
Eublaberus posticus
Galiblatta williamsi
Gromphadorhina brunneri
Gromphadorhina chopardi
Gromphadorhina portentosa
Hormetica scrobiculata
Hormetica ventralis
Hyporhicnoda litomorpha
Lanxoblatta emarginata
Leucophaea maderae
Macropanesthia rhinoceros
Nauphoeta cinerea
Panchlora nivea
Panchlora sp.
Panchlora viridis
Panesthia sp.
Panesthia stellata
Petasodes dominicana
Phoetalia circumvagans
Phortioica phoraspoides
Pinaconota sicca
Proscratea complanata
Pycnoscelus indicus
Pycnoscelus surinamensis
Pycnoscelus surinamensis
Rhabdoblatta annandalei
Rhabdoblatta sp.39T
Sp. sp41B
Blattellidae
Agmoblatta thaxteri
Amazonina conspersa
Amazonina n.sp.
Amazonina n.sp.38B
Attaphila fungicola
Blattella bisignata
Blattella germanica
Blattella lituricollis lituricollis
Blattella sauteri
Blattella sp.C43T
Blattella sp.D
Blattella sp.E71A
Blattella vaga
Cariblatta lutea minima
Dendroblatta cnephaia
Dendroblatta sobrina
Ectobius pallidus
Gislenia australica
Hemithyrsocera latera/is
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
46
74
74
38
74
40
48
38
48
42
50
38
36
44
40
66
44
28
64
76
64
32
32
44
66
24
80
38
36
32
38
38
38
54
54
66
34
36
36
35
53
50
32
30
45
73
73
37
73
39
47
37
47
41
49
37
35
43
39
65
43
27
63
75
63
31
31
43
65
23
79
37
35
31
37
37
37
53
53
65
33
35
35
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
49
31
29
XO
XO
XO
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
26
24
26
26
34
24
24
26
24
28
30
50
24
26
32
26
22
34
26
25
23
25
25
33
23
23
25
23
27
29
49
23
25
31
25
21
33
25
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
111
Taxa
Ischnoptera castanea
Ischnoptera deropeltiformis
Ischnoptera galibi
Ischnoptera mura
Ischnoptera panamae
Ischnoptera rufa rufa
Ischnoptera sp.93B
Loboptera decipiens
Lobopterella dimidiatipes
Lophoblatta brevis
Lophoblatta fissa
Lupparia notulata
Margattea punctulata
Nahublattella n.sp.72B
Parasymploce limbata
Parasymploce sp.25T
Parasymploce sp.30T
Parcoblatta fulvescens
Parcoblatta pennsylvanica
Pseudomops septentrionalis
Shawella couloniana
Supella longipalpa
Symploce capitata
Symploce fusca
Symplocodes marmorata
Symplocodes n.sp.50
Xestoblatta immaculata
Blattidae
Blatta orientalis
Cryptocercus punctulatus
Deropeltis erythrocephala
Duchailluia n.sp.
Eurycotis biolleyi
Eurycotis decipiens
Eurycotis floridana
Lamproblatta albipalpus
Pelmatosilpha coriacea
Periplaneta americana
Periplaneta australasiae
Periplaneta brunnea
Periplaneta fuliginosa
Periplaneta japonica
Polyphagidae
Arenivaga investigata
Cryptocercus primarius
Cryptocercus punctulatus
Cryptocercus punctulatus
Cryptocercus relictus
Latindia sp.3
Polyphaga aegyptiaca
Collembola
Bourletiellidae
Bourletiella arvalis
PGE
Bourletiella hortensis
PGE
Cyphoderidae
Cyphoderus albinus
Cyphoderus albinus
Dicyrtomidae
Dicyrtomina ornata
PGE
Ptenothrix italica
PGE
Entomobryidae
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
34
50
44
34
40
40
38
34
38
32
16
20
32
20
24
32
32
38
38
32
36
20
28
30
40
38
38
33
49
43
33
39
39
37
33
37
31
15
19
31
19
23
31
31
37
37
31
35
19
27
29
39
37
37
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
48
40
50
42
22
28
28
18
38
34
28
28
28
34
47
39
49
41
21
27
27
17
37
33
27
27
27
33
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
243
243
243
243
243
243
243
243
243
243
243
243
243
243
46
20
48
38
18
XO
XO
XO
XO
XO
64
45
19
47
37
17
22
63
XO
243
244
245
244
244
243
243
12
10
XO
246
12
10
XO
246
12
12
247
247
12
10
XO
246
14
12
XO
246
112
Taxa
Entomobrya lanuginosa
Entomobrya marginata
Entomobrya marginata
Entomobrya muscorum
Entomobrya nivalis
Heteromurus nitidus
Heteromurus nitidus
Lepidocyrtus paradoxus
Lepidocyrtus paradoxus
Orchesella bifasciata
Orchesella cincta
Orchesella cincta
Orchesella flavescens
Orchesella flavescens
Orchesella villosa
Pseudosinella sp.
Pseudosinella sp.
Sinella coeca
Sinella coeca
Sinella curviseta
chiasmatic male meiosis
Willowsia buski
Willowsia buski
Hypogastruridae
Ceratophysella bengtssoni
Ceratophysella denticulata
Ceratophysella bengtssoni
Ceratophysella denticulata
Hypogastrura adexilis
chiasmatic male meiosis
Hypogastrura manubrialis
Hypogastrura viatica
Mesogastrura ojcoviensis
Mesogastrura ojcoviensis
Xenylla grisea
Xenylla grisea
Xenylla planipila
Xenylla planipila
Isotomidae
Folsomia multiseta
Isotoma cinerea
Isotoma viridis
Isotoma cinerea
Isotoma notabilis
Isotomiella minor
lsotomiella minor
Proisotoma crassicauda
Proisotoma minuta
Proisotoma minuta
Proisotoma subminuta
Katiannidae
Stenognathellus denisi
PGE
Neanuridae
Anurida maritima
Bilobella aurantiaca
Bilobella grassei
Neanura monticola
Onychiuridae
Onychiurus armatus
Onychiurus fimatus
Onychiurus furcifer
Onychiurus pseudogranulosus
Onychiurus scotarius
Onychiurus armatus
Sexual
System
Ploidy
Female
2n
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
Male
2n
11
Sex Chrom. Source
Karyotype
XO
247
247
247
247
247
247
247
247
247
247
247
247
247
247
248
247
247
247
247
249
12
12
247
247
14
14
14
14
14
247
247
247
247
249
14
14
14
14
14
14
14
14
14
14
14
14
14
12
12
14
14
14
14
13
13
13
13
XO
XO
XO
XO
13
13
XO
XO
11
XO
247
247
247
247
247
247
247
247
247
247
247
247
247
247
247
247
247
247
247
246
8
14
12
14
14
16
14
14
248
247
247
247
13
XO
247
247
247
247
16
14
113
247
247
Taxa
Sexual
System
Ploidy
Onychiurus fimatus
Onychiurus furcifer
Onychiurus paradoxus
Onychiurus pseudogranulosus
Onychiurus scotarius
Onychiurus sibiricus
Poduridae
Podura aquatica
achiasmatic male meiosis
Podura aquatica
Podura aquatica
Sminthuridae
Allacma fusca
PGE
Caprainea marginata
PGE
Lipothrix lubbocki
PGE
Sminthurus viridis
PGE
Sminthurididae
Sminthurides aquaticus
PGE
Tomoceridae
Tomocerus flavescens
Tomocerus minor
Tomocerus minutus
Coleoptera
Anobiidae
Ernobius mollis
Gibbium psylloides
Lasioderma serricorne
Ptinus clavipes
parth
3
-automixis IIF premeiotic doubling pseudogamous
Ptinus hirtellus
-pseudogamous
parth
3
Ptinus villiger
Stegobium paniceum
Anthicidae
Notoxus calcaratus
Notoxus constrictus
Notoxus monodon
Anthribidae
Araecerus fasciculatus
Euparius marmoreus
Euparius oculatus
Habrissus longipes
Platystomos sellatus
Ptychoderes bivittatus
Ptychoderes tricostifrons
Tropideres germanus
Tropideres japonicus
Tropideres laxus
Attelabidae
Apoderus balteatus
Apoderus coryli
Apoderus erythrogaster nigriventris
Apoderus erythrogaster rufiventris
Apoderus jekeli
Auletobius irkutensis japonicus
32
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
16
14
14
14
16
14
247
247
247
247
247
247
250
22
8
247
248
12
10
XO
246
12
10
XO
246
246
12
10
XO
246
246
12
12
12
247
247
251
11
XO
22
18
18
27
22
18
18
Xyp
Xyp
Xyp
1
1
1
2
18
27
18
Xyp
2
3
18
18
18
18
18
17
XY
Xyp
XO
3
1
1
18
18
18
18
18
18
Xyp
Xyp
Xyp
1
1
1
22
22
22
22
22
22
22
24
22
22
22
22
49
22
22
22
22
22
24
22
22
Xyp
Xyp
Xyyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
1
1
1
1
1
1
22
26
28
28
26
36
38
32
22
26
28
28
26
36
38
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
1
1
1
4
1
1
1
1
114
Taxa
Sexual
System
Ploidy
Auletobius puberulus
Byctiscus launipennis
Centrocorynus aemulus
Deporaus affectatus
Deporaus harmanni
Deporaus marginatus
Deporaus minimus
Deporaus sp. 1
Deporaus sp. 2
Deporaus unicolor
Hypera viciae
Larinus sturnus
Liophloeus lentus
Liparus glabrirostris
Pissodes notatus
Rhynchites heros
Rhynchites pilosus
Rhynchites placidus
Rhynchites sp. 1
Bostrichidae
Apate monacha
Bostrychopsis bengalensis
Rhizopertha dominica
Sinoxylon anale
-B chromosomes
Brentidae
Apion abruptum
Apion collare
Apion sp. 1
Apion sp. 2
Aplemonus tuberculata
Brentus anchorago
Brentus mexicanus
Ceratapion austriacum
Ceratapion penetrans
Cylas formicarius
Cylas formicarius elegantulus
Eutrichapion melancholicum
Nanophyes marmoratus
Nemocephalus femoratus
Oxystoma cerdo
Proepisphales elegans
Pseudorychoides insignis
Rhopalapion longirostre
Squamapion flavimanum
Trachelizus sp. 1
Buprestidae
Spruce borer sp. I
Spruce borer sp. II
Acmaeodera boryi
Acmaeodera flavofasciata
Acmaeodera flavolineata
Acmaeodera gibbulosa
Acmaeodera hepburni
Acmaeodera pilosellae persica
Acmaeodera vetusta
Agrilus angustulus
Agrilus anxius
Agrilus araxenus
Agrilus liragus
Agrilus obscuricollis
Agrilus politus pseudocoryli
Agrilus sp.
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
32
Xyp
24
22
38
22
32
22
38
32
38
34
32
22
28
24
22
22
26
22
22
34
29
24
22
38
22
32
22
38
32
38
34
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
5
5
1
1
5
1
1
1
1
1
1
4
4
4
4
4
1
1
1
1
22
20
18
20
22
20
18
20
Xyp
Xyp
Xyp
Xyp
1
1
1
1
22
22
22
22
26
12
12
22
22
22
22
22
22
12
22
12
12
22
22
12
22
22
22
22
26
12
12
22
22
22
Xyp
22
22
12
22
12
12
22
22
12
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
5
1
1
6
6
1
20
22
18
18
20
18
18
20
18
22
22
20
20
20
20
20
20
22
18
18
20
18
18
NeoXY
18
22
22
20
20
20
20
20
XY
XY
Xyr
Xyr
NeoXY
Xyr
NeoXY
9
Xyr
Xyp
Xyp
XY
NeoXY
Xyp
XY
NeoXY
7
7
7
7
8
7
7
28
24
22
22
26
22
22
34
22
20
115
6
6
1
6
1
1
6
6
1
7
10
7
10
7
10
7
7
Taxa
Agrilus sp. nr. Pensus
Anthaxia amasina
Anthaxia bicolor
Anthaxia deaurata
Anthaxia hungarica
Anthaxia lgockii
Anthaxia mirabilis
Anthaxia olympica
Anthaxia podolica
Anthaxia sponsa
Anthaxia viridifrons
Buprestis fasciata
Capnodis miliaris
Capnodis tenebrionis
Castiarina adelaidae
Castiarina argillacea
Castiarina cupreoflava
Castiarina decemmaculata
Castiarina flavopicta
Castiarina grata
Castiarina rufipennis
Castiarina sexplagiata
Castiarina simulata
Castiarina subnotata
Castiarina subtincta
Castiarina triramosa
Chalcophora lacustris
Chrysobothris affinis tetragramma
Chrysobothris dentipes
Chrysobothris floricola
Coraebus rubi
Coraebus sinuatus
Dicera divaricata
Dicera prolongata
Dicera tenebrosa
Dicerca aenea validiuscula
Euchroma gigantea
-B chromosomes
Sexual
System
Ploidy
Female
2n
Male
2n
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
20
20
14
22
22
22
22
22
22
22
22
22
22
22
22
21
16
16
16
22
24
20
20
20
20
26
20
14
22
22
22
22
22
22
22
22
22
22
22
22
22
16
16
16
22
24
20
20
20
20
Sex Chrom. Source
Karyotype
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XO
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
XXXYY
7
8
7
7
7
7
7
8
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
8
7
7
10
10
7
7
7
9
11
24
XXXYYY
11
24
XXXYYY
11
32
XXXYYY
12
34
XXXYYY
12
36
XXXYYY
12
26
26
26
24
12
16
16
22
20
20
20
16
24
40
24
24
42
Xyp
NeoXY
XY
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
7
7
7
7
7
7
7
10
10
7
7
9
10
10
7
7
9
-B chromosomes
-B chromosomes
-B chromosomes
-B chromosomes
-B chromosomes
Julodella globithorax
Julodis andreae
Julodis faldermanni
Julodis whithilli
Melanophila acuminata
Melanophila drummondi
Melanophila intrusa
Meliboeus caucasicus
Ovalisia nadezhdae
Perotis cuprata
Perotis lugubris
Sphaerobothris aghababiani
Sphenoptera artemisiae
Sphenoptera glabrata
Sphenoptera mesopotamica
Sphenoptera scovitzi
Sphenoptera scpvitzii
26
26
26
24
12
16
16
22
20
20
20
16
24
40
116
Taxa
Sexual
System
Ploidy
-Male 2n = 38-46
Sphenoptera tamarisci
Stenocera sp
Sternocera laevigata
Sternocera nitidicollis
Stigmodera cancellata
Stigmodera goryi
Stigmodera gratiosa
Stigmodera macularia
Stigmodera porosa
Stigmodera roei
Themognatha alternata
Themognatha barbiventris
Themognatha bonvouloiri
Themognatha chalcodera
Themognatha chevrolati
Themognatha donovani
Themognatha heros
Themognatha mitchelli
Themognatha mniszechi
Themognatha nickerli
Themognatha parvicollis
Themognatha pubicollis
Themognatha regia
Themognatha tricolorata
Themognatha variabilis
Themognatha viridicincta
Byrrhidae
Byrrhus sp.
Cytilus alternatus
Cantharidae
Cantharis cryptica
Cantharis fusca
Cantharis lateralis
Cantharis livida
Cantharis nigra
Cantharis nigricans
Cantharis pallida
Cantharis pellucida
Cantharis rufa
Cantharis rustica
Cantharis sp.
Cantharis tenuicollis
Cantharis thoracica
Chauliognathus expansus
Chauliognathus fallax
Chauliognathus flavipes
Chauliognathus lineatus
Chauliognathus octomaculatus
Chauliognathus riograndensis
Chauliognathus sp. 1
Chauliognathus tetrapunctatus
Malthinus seriepunctatus
Malthodes dispar
Malthodes minimus
Rhagonycha fulva
Rhagonycha lignosa
Rhagonycha limbata
Rhagonycha lutea
Rhagonycha testacea
Silis ruficollis
-B chromosomes
Telephorus nobilitatus
Tylocerus barberi
14
14
14
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
30
26
26
26
22
22
22
22
22
22
20
22
22
22
22
22
22
22
22
20
22
22
22
22
22
22
30
26
26
26
22
22
22
22
22
22
20
22
22
22
22
22
22
22
22
20
22
22
22
22
22
22
NeoXY
NeoXY
NeoXY
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
8
13
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
18
18
18
18
Xyp
Xyp
1
1
14
14
20
14
14
14
14
14
14
14
14
20
20
14
14
14
14
14
13
13
14
13
12
12
12
14
14
14
14
14
14
13
XO
20
13
13
13
Xyp
XO
XO
XO
13
13
13
13
20
20
13
13
13
13
13
XO
XO
13
XO
XO
XO
XO
XO
Xyp
Xyp
XO
XO
XO
XO
XO
15
15
XO
15
14
14
4
14
14
14
14
14
14
14
14
1
1
14
15
15
15
15
11
XO
13
XO
13
13
13
13
XO
XO
XO
XO
14
14
14
14
14
14
14
14
14
14
14
13
13
XO
XO
1
1
117
15
Taxa
Carabidae
Abacetus salzamanni
Abax ater
Abax ovalis
Abax parallelepipedus
Abax parallelus
Acinopus giganteus
Acinopus gutturosus
Acinopus picipes
Acupalpus brunnipes
Acupalpus elegans
Acupalpus maculatus
Agonini sp. 1
Agonini sp. 2
Agonum bogemanni
Agonum corvus
Agonum cupripenne
Agonum dolens
Agonum ericeti
Agonum errans
Agonum extensicolle
Agonum fuliginosum
Agonum gracile
-Male 2n = 35-36
Agonum livens
Agonum marginatum
Agonum micans
Agonum moestum
Agonum moestum longipenne
Agonum muelleri
Agonum munsteri
Agonum nigrum
Agonum retractum
Agonum sexpunctatum
Agonum sp. 1
Agonum sp. 2
Agonum thoreyi
Agonum thoreyi melanocephalus
Agonum viridicupreum
Amara aenea
Amara apricaria
Amara arcuata
Amara aulica
Amara batesi
Amara communis
Amara equestris
Amara familiaris
Amara fervida
Amara fulva
Amara giganteus
Amara impuncticollis
Amara ingenua
Amara laevipennis
Amara metallescens
-B chromosomes
Amara moerens
Amara montivaga
Amara nila
Amara plebeja
Amara rotundicollis
Amara rufescens
Amara similata
Amara simplex
Amblycheila baroni
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
42
41
35
35
35
35
37
37
37
45
22
39
30
36
36
39
36
39
34
37
37
37
36
XO
43
37
35
34
XY
37
36
34
37
37
37
37
35
35
37
35
33
37
33
37
37
XO
XO
33
34
33
35
41
34
37
32
21
XO
NeoXY
XO
XO
XO
37
37
22
34
XO
XO
XY
XY
32
XY
44
XY
36
36
38
22
36
36
40
34
38
38
38
44
38
34
34
34
38
36
34
38
38
38
36
36
38
36
34
38
38
30
34
34
34
36
42
38
32
38
38
22
34
26
24
32
20
44
118
XO
XO
XO
XY
NeoXY
NeoXY
XO
XY
XO
XO
XO
XY
18
XO
XY
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
NeoXY
16
16
16
16
16
17
17
18
17
19
17
18
18
18
17
18
18
18
18
17
18
17
18
18
17
18
18
18
18
18
18
18
18
18
18
17
18
18
17
18
17
18
17
18
18
17
17
18
18
17
18
17
17
17
17
17
17
17
17
17
20
Taxa
Sexual
System
Ploidy
Amblystomus bivittatus
Amblystomus quadriguttatus
Amblytelus curtus
Amphasia interstitialis
-Male 2n = 35-37
Amphasia sericea
Anchomenidius astur
Anchomenus dorsalis
Angoleus crenatus
Angoleus nitidus
Anisodactylus binotatus
Anisodactylus hispanus
Anisodactylus signatus
Anisodactylus virens
Anomoglossus emarginatus
Anthia sexguttata
Apenes marginalis
Aptinus displosor
Arthropterus sp. 1
Asaphidion caraboides
-B chromosomes
Asaphidion curtum
Asaphidion cyanicorne
Asaphidion rossii
Baudia anomala
Bembidion aeneicolle
Bembidion aenulum
Bembidion alaskense
Bembidion ambiguum
Bembidion americanum
Bembidion anchonoderum
Bembidion andreae andreae
Bembidion andreae bualei
Bembidion antiquum
Bembidion aratum
Bembidion arcticum
Bembidion arenobilis
Bembidion arizonae
Bembidion articulatum
Bembidion ascendens
Bembidion assimile
Bembidion atrocoeruleum
Bembidion balli
Bembidion basicorne
Bembidion bellorum
Bembidion bifossulatum
Bembidion biguttatum
Bembidion bimaculatum
Bembidion bipunctatum
Bembidion bowditchi
Bembidion brachythorax
Bembidion brevistriatum
Bembidion californicum
Bembidion callosum
Bembidion carinula
Bembidion carolinense
Bembidion carpetanum
Bembidion castor
Bembidion castum
Bembidion cf anguliferum
Bembidion cf. immaturum
Bembidion chalceum
Bembidion coeruleum
Bembidion coloradense
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
28
30
24
XY
30
36
XY
30
29
43
44
37
39
37
37
37
37
37
37
35
36
32
46
24
XO
24
24
24
45
24
24
24
24
24
24
XY
24
24
24
26
24
24
XY
XY
24
24
24
24
24
24
24
24
24
24
24
23
24
24
24
24
24
24
24
24
24
24
26
24
24
XY
44
38
40
38
38
38
38
38
38
36
46
24
24
24
24
46
24
24
24
24
24
24
24
24
24
24
26
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
26
24
24
119
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XO
XY
XY
XY
XY
XY
18
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XO
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
18
21
21
18
18
18
17
16
16
16
17
18
18
18
18
18
18
18
21
18
18
18
17
18
22
22
22
18
22
22
18
22
22
22
23
22
18
17
18
18
22
22
23
22
18
22
18
22
22
22
22
18
22
22
18
22
22
22
22
22
18
22
Taxa
Sexual
System
Bembidion complanulum
Bembidion compressum
Bembidion concolor
Bembidion concretum
Bembidion confusum
Bembidion connivens
Bembidion consanguineum
Bembidion constricticolle
Bembidion contraetum-group contractum
Bembidion contraetum-group viridicolle
Bembidion convexulum
Bembidion cordatum
Bembidion coxendix
Ploidy
24
Male
2n
Sex Chrom. Source
Karyotype
24
24
24
24
24
24
24
24
24
24
24
24
24
24
22
XY
24
24
22
23
24
25
25
24
24
22
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
22
24
24
24
24
24
24
24
XY
22
24
24
24
24
24
24
24
24
XY
XY
XY
XY
XY
XY
XY
24
24
24
24
24
Bembidion cribum
Bembidion dahli
Bembidion dauricum
Bembidion decorum
Bembidion diligens
Bembidion doris
Bembidion dorsale
Bembidion dudichi
Bembidion ephippium
Bembidion fasciolatum egregium
Bembidion flebile
Bembidion fortestriatum
Bembidion fortunatum
Bembidion foveum
24
24
24
24
24
24
24
24
24
24
Bembidion frontale
Bembidion fumigatum
Bembidion gebleri turbatum
Bembidion genei
Bembidion geniculatum
Bembidion gordoni
Bembidion graciliforme
Bembidion grapei
Bembidion graphicum-Nr 1'
Bembidion graphicum-Nr 2'
Bembidion gratiosum
Bembidion grisvardi
Bembidion guadarramense
Bembidion guttula
Bembidion haruspex
Bembidion hasti
Bembidion hasurada
Bembidion hesperium
Bembidion hispanicum
Bembidion honestum
Bembidion humboldtense
Bembidion hyperboraeorum
Bembidion hypocrita
Bembidion ibericum
Bembidion impotens
Bembidion inaequale inaequale
Bembidion inaequale opaciceps
Bembidion incrematum
Bembidion indistinctum
Bembidion insulatum
Bembidion integrum
Bembidion intermedium
Bembidion interventor
Bembidion iricolor
Bembidion iridescens
Female
2n
22
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
22
24
24
24
24
24
24
24
22
22
24
24
24
24
24
24
24
24
120
22
XY
XY
XO
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
22
XY
XY
XY
XY
XY
XY
XY
XY
XY
22
22
22
22
22
22
22
22
22
22
22
22
22
18
18
22
18
22
18
22
18
18
18
22
22
18
17
22
22
18
22
18
18
22
22
22
22
22
22
18
18
18
22
22
17
17
22
18
22
22
22
18
18
22
22
22
22
22
22
22
22
18
22
Taxa
Sexual
System
Bembidion ksteri paulinoi
Bembidion kuprianovi-Nr 1
Bembidion kuprianovi-Nr 2
Bembidion lacunarium
Bembidion lampros
Bembidion lapponicum
Bembidion latiplaga
Bembidion laxatum
Bembidion lenae
Bembidion levettei carrianum
Bembidion levettei levettei
Bembidion levigatum
Bembidion littorale
Bembidion lorquini
Bembidion louisella
Bembidion lunulatum
Bembidion maculatum
Bembidion manningense
Bembidion maorinum maorinum
Bembidion mckinleyi carneum
Bembidion mckinleyi mckinleyi
Bembidion mexicanum
Bembidion millerianum
Bembidion mimus
Bembidion minimum
Bembidion modestum
Bembidion morulum
Bembidion mundum
Bembidion mutatum
Bembidion nebraskense
Bembidion nevadense
Bembidion nigripes
Bembidion nigrocoeruleum
Bembidion nigrum
Bembidion nitidulus ovalipennis
Bembidion nitidum
Bembidion nudipenne
Bembidion oberthueri-group oberthueri
Bembidion obliquulum
Bembidion obscurellum
Bembidion obtusangulum-group mormon
Bembidion obtusangulum-group obtusangulum
Bembidion obtusum
Bembidion occultator
Bembidion octomaculatum
Bembidion operosum
Bembidion patruele
Bembidion petrosum
Bembidion planatum
Bembidion planiusculum
Bembidion planum
Bembidion platynoides
Bembidion poculare
Bembidion praecinctum
Bembidion praticola
Bembidion properans
Bembidion pseudocautum
Bembidion punctatostriatum
Bembidion punctulatum
-B chromosomes
Bembidion quadrifoveolatum
Bembidion quadrimaculatum dubitans
Bembidion quadrimaculatum oppositum
Ploidy
22
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
24
24
24
24
24
24
24
24
24
22
22
24
26
24
24
24
24
23
24
24
24
24
XY
22
24
36
22
24
24
24
23
XY
XY
XY
24
26
24
24
24
24
24
24
22
24
24
24
24
XY
XY
XY
XY
XY
XO
XY
XY
XY
XY
22
XY
XY
22
24
24
24
24
24
24
24
24
24
24
24
26
24
24
24
24
24
24
22
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
26
24
121
24
24
24
24
23
24
XY
24
23
24
24
24
23
24
24
24
24
24
24
24
24
24
24
24
24
26
22
24
XY
XY
XY
XO
22
22
22
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
17
XY
XY
XY
XY
XO
XY
22
XY
XO
XY
XY
XY
XO
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
22
18
22
22
22
18
22
18
22
22
22
22
17
22
23
18
18
22
22
18
22
19
17
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
18
22
22
22
22
22
22
22
22
22
22
18
22
22
18
18
22
22
22
Taxa
Sexual
System
Bembidion quadripustulatum
Bembidion quadrulum
Bembidion rapidum
Bembidion recticolle
Bembidion ripicola
Bembidion rivulare
Bembidion roosvelti
Bembidion rothfelsi
Bembidion ruficorne
Bembidion rufinum
Bembidion rufotinctum
Bembidion rupicola
Bembidion rusticum lenensoides
Bembidion salebratum
Bembidion salinarium
Bembidion satelles
Bembidion scopulinum
Bembidion sejunctum sejunctum
Bembidion sejunctum semiaureum
Bembidion semicinctum
Bembidion semistriatum
Bembidion seudderi-group consimile
Bembidion seudderi-group flohri
Bembidion seudderi-group obtusidens
Bembidion seudderi-group scudderi
Bembidion siculum
Bembidion sierricola
Bembidion simplex
Bembidion sordidum
Bembidion sp nr ampliceps-Nr 1
Bembidion sp nr ampliceps-Nr 2
Bembidion sp nr aratum
Bembidion sp nr impotens
Bembidion sp nr mundum
Bembidion sp nr transversale-Nr 1
Bembidion sp nr transversale-Nr 2
Bembidion sp.
Bembidion stephensi
Bembidion striatum maurus
Bembidion sulcipenne hyperboroides
Bembidion tairuense
Bembidion tenellum
-B chromosomes
Bembidion tetracolum
Bembidion texanum
Bembidion thetys
Bembidion tibiale
Bembidion timidum
Bembidion transparens
Bembidion transversale
Bembidion tricolor
Bembidion umbratum
Bembidion vandykei
Bembidion varicolor
Bembidion varium
Bembidion versicolor
Bembidion versutum
Bembidion vicinus
Bembidion wickhami
Bembidion yukonoum
Bembidion zephyrum
Blemus sp
Ploidy
24
24
24
24
24
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
24
24
24
24
24
24
34
26
24
24
24
24
24
24
24
24
24
24
23
24
24
34
26
24
24
23
XY
24
24
22
24
24
24
24
24
24
XY
24
24
24
24
22
24
XY
XY
24
22
24
24
24
32
24
XY
24
24
24
XY
XY
XY
XY
XO
XY
XY
XY
XY
XY
XY
XO
22
XY
XY
23
24
26
24
24
24
24
24
24
22
24
24
24
24
23
24
24
24
24
24
22
25
XO
XY
XY
XY
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
32
24
24
24
24
24
24
24
26
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
22
122
XY
XY
XY
XY
XY
XY
22
XY
XY
XY
XY
XY
22
22
XY
XY
XY
XY
XY
XY
18
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XO
XY
XY
XY
XY
XY
XY
18
22
22
22
18
18
22
23
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
17
22
22
22
22
22
22
22
22
18
18
22
22
18
22
18
18
22
17
18
22
22
22
22
22
22
17
18
22
22
22
18
22
22
22
17
Taxa
Sexual
System
Ploidy
Female
2n
Blethsia multipunctata
Brachinus andalusiacus
Brachinus beaticus
Brachinus bodemeyeri
Brachinus cordicollis
Brachinus crepitans
Brachinus exhalans
Brachinus explodens
Brachinus humeralis
Brachinus janthinipennis
Brachinus longicornis
18
40
22
Brachinus pateri
22
28
18
Brachinus pecoudi
Brachinus plagiatus
Brachinus sclopeta
28
Brachinus sexmaculatus
Brachinus variventris
Bradybaenus festivus
Broscus cephalotes
Broscus glaber
Broscus rutilans
Calachroa sexpunctata
Calathus abaxoides
Calathus ambiguus chevrolati
Calathus amplius
Calathus angularis
Calathus angustulus
Calathus ascendens
38
36
38
37
38
36
22
Calathus asturiensis
Calathus auctus
Calathus baeticus mateui
Calathus brevis
Calathus cf fuscipes
Calathus ciliatus
Calathus circumseptus
Calathus cognatus
Calathus dejeani hispanicus
Calathus depressus
Calathus erratus
Calathus freyi
Calathus fuscipes
Calathus fuscipes fuscipes
Calathus fuscipes graecus
Calathus granatensis
Calathus hispanicus
Calathus luctosus
Calathus melanocephalus
Calathus micropterus
Calathus mollis
Calathus oreades
Calathus rectus
Calathus rotundatus
Calathus rubricus
Calathus rufocastaneus
Calathus spretus
Calathus vuillefroyi
Calosoma beesoni
Calosoma chinense
38
38
36
38
36
38
38
38
40
38
38
38
38
38
38
38
36
38
38
28
123
Male
2n
32
32
33
25
18
39
21
36
28
18
32
34
22
20
18
32
31
32
28
32
37
36
24
30
30
23
55
XO
36
37
36
24
23
39
37
37
37
38
37
43
37
37
36
37
36
37
37
37
39
37
37
37
38
37
39
37
37
39
36
37
37
37
28
28
Sex Chrom. Source
Karyotype
NeoXY
XO
XO
XY
NeoXY
XY
XO
XY
XXY
17
18
24
24
18
18
18
24
18
18
24
24
24
24
18
18
24
24
18
18
18
18
17
17
18
18
18
18
XO
XY
XYY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XO
XY
XO
XO
XY
18
17
17
18
18
18
18
17
18
17
18
18
17
18
18
18
18
18
18
18
18
18
17
17
18
18
18
18
17
18
18
18
18
17
18
18
18
Taxa
Sexual
System
Calosoma inquisitor
Calosoma maderae indicum
Calosoma maderae indigator
Calosoma orientale
Calosoma sycophanta
Carabus abbreviatus
Carabus alpestris alpestris
Carabus alpestris hoppei
Carabus arcensis arcensis
Carabus auronitens auronitens
Carabus auronitens festivus
Carabus blaptoides
Carabus blaptoides blaptoides
Carabus blaptoides lewisii
Carabus blaptoides oxuroides
Carabus blaptoides rugipennis
Carabus caelatus
Carabus caelatus schreiberi
Carabus cancellatus
Carabus cancellatus cancellatus
Carabus cancellatus celticus
Carabus carinthiacus
Carabus clathratus clathratus
Carabus coarctatus
Carabus coriaceus
Carabus coriaceus coriaceus
Carabus creutzeri kircheri
Carabus creutzeri rinaldoi
Carabus croaticus croaticus
Carabus depressus bonellii
Carabus deyrollei
Carabus dufouri
Carabus errans
Carabus faustus faustus
Carabus fruhstorferi
Carabus galicianus
Carabus ghiliani
Carabus glabratus
Carabus glabratus glabratus
Carabus granulatus
Carabus granulatus granulatus
Carabus guadarramus
Carabus hispanus
Carabus hortensis hortensis
Carabus intricatus
Carabus intricatus intricatus
Carabus irregularis irregularis
Carabus lineatus basilicus
Carabus lineatus lateralis
Carabus lineatus lineatus
Carabus lineatus troberti
Carabus lusitanicus brevis
Carabus macrocephalus
Carabus macrocephalus cantabricus
Carabus melancholicus costatus
Carabus monticola
Carabus morbillosus
Carabus nemoralis
Carabus nemoralis meridionalis
Carabus nemoralis nemoralis
Carabus nitens
Carabus olympia
Carabus problematicus
Carabus problematicus gallicus
Ploidy
28
28
28
28
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
28
28
28
28
28
28
28
28
27
28
28
28
28
28
XY
29
28
Xyr
28
28
Xyr
28
28
27
XY
28
28
XY
XY
XY
XO
XY
XY
28
30
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
26
28
18
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
28
30
28
28
28
28
28
124
28
28
28
28
28
28
28
28
28
28
28
28
28
28
25
28
17
XY
28
28
28
28
28
XY
28
28
28
28
28
28
28
XY
28
28
27
29
28
28
28
28
XY
18
XO
Xyr
18
Xyr
Xyr
18
XY
XO
18
XY
18
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
Xyr
XY
XO
XY
XO
18
XY
XY
XY
XY
XY
18
XY
XY
XY
XY
XY
18
XY
XY
XO
XO
18
XY
XY
XY
18
18
18
17
18
18
18
18
18
18
25
1
18
18
18
18
18
18
18
17
18
18
18
18
18
18
18
17
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
17
18
18
18
18
17
18
Taxa
Sexual
System
Ploidy
Carabus punctatoauratus
Carabus purpurascens
Carabus purpurascens fulgens
Carabus purpurascens purpurascens
Carabus pyrenaeus
Carabus pyrenaeus occidentalis
Carabus pyrenaeus pyrenaeus
Carabus rugosus
Carabus rutilans
Carabus serratus
Carabus silvestris silvestris
Carabus solieri
Carabus splendens
Carabus splendens splendens
Carabus violaceus germari
Carenum interruptum
Carterus fulvipes
Carterus interceptus
Carterus rotundicollis
Castelnaudi wilsoni
Ceratoderus bifasciatus
Ceroglossus chilensis sp a
Ceroglossus chilensis sp b
Ceroglossus chilensis sp c
-B chromosomes
Chlaenius aestivus
Chlaenius darlingensis
Chlaenius duvauceli
Chlaenius fraterculus
Chlaenius greyanus
Chlaenius hamifer
Chlaenius impressicollis
Chlaenius impunctifrons
Chlaenius laetiusculus
Chlaenius laticollis
Chlaenius malachinus
Chlaenius neelgheriensis
Chlaenius nitidicollis
Chlaenius olivieri
Chlaenius orbicollis
Chlaenius pallipes
Chlaenius panagaeoides
Chlaenius pennsylvanicus
Chlaenius pictus
Chlaenius praefectus
Chlaenius pretiosus
Chlaenius pulcher
Chlaenius scapularis
Chlaenius sp.
Chlaenius spoliatus
Chlaenius tricolor
Chlaenius tristis
Chlaenius velutinus auricollis
Chlaenius vestitus
Chlaenius virens
Chlaenius xanthospilus
28
28
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
30
28
28
28
29
28
28
XY
28
28
XO
XY
XY
18
28
28
28
28
30
26
28
30
30
28
24
58
52
28
30
42
XY
28
29
26
28
27
29
29
28
28
24
57
57
57
51
XY
18
18
XY
XO
NeoXY
XO
XO
XO
XY
XY
XO
XO
18
18
18
18
18
17
18
18
18
18
18
18
18
18
21
18
17
18
21
17
26
26
26
30
42
41
XY
XY
XY
XO
XO
XO
38
38
36
38
20
38
38
38
40
38
40
38
38
38
38
38
38
38
36
34
37
37
39
37
37
39
37
37
37
37
35
35
37
37
37
35
37
20
37
37
37
40
37
39
37
37
37
37
37
37
37
36
XO
XO
XO
XO
XY
XO
XO
XO
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
Xyp
18
21
18
18
21
21
18
18
18
18
18
18
18
17
18
18
18
18
18
18
18
18
18
18
18
18
18
17
18
18
17
18
18
37
37
Xyp
18
23
23
XXY
XXY
17
17
34
38
38
40
38
40
38
38
38
38
36
36
XO
XO
XO
XO
XO
XO
XO
XO
-Xyp Subsequent studies failed to replicate
-Xyp Subsequent studies failed to replicate
Cicindela agnata
Cicindela albina
125
Taxa
Sexual
System
Cicindela ancocisconensis
-Xx00 Subsequent studies failed to replicate
Cicindela angulata
Cicindela argentata
Cicindela aterrima
Cicindela aurulenta
Cicindela bigemina
Cicindela brevis
Cicindela campestris
Cicindela cardinalba
Cicindela chloris
Cicindela chloropleura
Cicindela cognata
Cicindela cyclobregma
Cicindela deserticoloides
Cicindela dorsalis
Cicindela dromicoides
Cicindela duodecimguttata
Cicindela duponti
Cicindela flohri
Cicindela formosa generosa
Cicindela fulgoris
Cicindela funerea
Cicindela gillesensis
Cicindela hemichrysea
Cicindela hispanica
Cicindela holosericea
Cicindela hybrida
Cicindela indica
Cicindela intermedia
Cicindela lemniscata
Cicindela litorea goudoti
Cicindela littoralis
Cicindela littoralis
Cicindela maroccana pseuomaroccana
Cicindela marutha
Cicindela maura
Cicindela minuta
Cicindela multiguttata
Cicindela nebuligera
Cicindela nigrocoerulea
Cicindela obsoleta
Cicindela ocellata
Cicindela oregona
Cicindela pimeriana
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
22
Xx00
18
21
21
22
22
23
23
22
24
23
23
21
21
22
22
21
22
23
22
22
22
23
26
22
21
21
22
21
21
23
22
22
22
23
23
22
23
23
21
23
22
21
22
20
22
22
22
22
21
22
22
XXY
XXY
XXXY
XXXY
XXY
XXY
XXXY
XXXY
XXY
XXY
XXY
XXY
XXXY
XXXY
XXY
XXXY
XXY
XXXY
XXXY
XXXY
XXY
XXXY
XXXY
XXY
XXY
XXXY
XXY
XXY
XXXXY
XXXY
XXY
XXXY
XXXXY
XXXXY
XXXY
XXY
XXY
XXY
XXXXY
XXXY
XXY
XXXY
Xyc
XXXY
XXOO
XXXY
XXOO
XXY
XXXY
XXOO
17
27
28
27
17
17
18
29
17
17
17
17
30
28
17
28
17
28
28
28
17
29
28
17
17
18
17
17
28
17
17
31
18
28
17
17
17
28
28
28
28
28
18
28
18
28
18
18
28
18
XXXY
22
21
22
21
21
22
21
22
22
28
XXXY
XXY
XXXY
XXY
XXY
XXXY
XXY
XXXY
XXOO
28
28
28
18
28
28
17
28
32
XXY
18
22
24
24
28
22
24
24
24
24
26
24
20
Cicindela punctulata
Cicindela purpurea
Cicindela repanda
-Subsequent studies failed to replicate
Cicindela roseiventris mexicana
Cicindela rufiventris
Cicindela rugatilis
Cicindela scutellaris
Cicindela scutellaris lecontei
Cicindela sedecimpunctata
Cicindela severa
Cicindela severini
Cicindela sexguttata
-Subsequent studies failed to replicate
Cicindela sexguttata sexguttata
22
22
22
21
126
Taxa
Cicindela silvicola
Cicindela sp. (saetigera group)
Cicindela spinolai
Cicindela splendida
Cicindela striatifrons
Cicindela striolata
Cicindela sumatrensis
Cicindela suturalis
Cicindela tranquebarica
Sexual
System
Ploidy
28
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
26
22
XXXY
21
22
23
23
23
23
22
21
22
22
40
XXY
29
XXY
XXXY
XXY
XXY
XXY
XXXXY
XXOO
XXY
Xyc
XXXY
XXXY
18
23
23
23
23
21
23
24
XXY
XXXXY
XXY
XXY
XXY
XXY
XY
44
46
30
34
34
34
18
12
37
37
37
37
XXY
XY
XY
17
17
17
17
17
17
21
21
17
17
18
18
18
18
18
18
18
17
17
18
26
22
22
42
Cicindela trifasciata
-B chromosomes
Cicindela triguttata
Cicindela trisignata
Cicindela viduata
Cicindela virgula
Cicindela viridilabris
Cicindela vittigera
Clivina australasiae
Clivina dilutipes
Clivina fossor
24
24
44
46
Clivina ypsilon
Colfax creagris
Colfax sp.
Colfax stevensi
Colliuris pennsylvanica
Craspedophorus angulatus
Cryptophonus fulvus
Cryptophonus litigiosus
Cryptophonus schaumii
Cryptophonus tenebrosus
Ctenostoma ornatum ornatum
Ctenostoma rugosum
Curtonotus alpina
Curtonotus aulica
Curtonotus giganteus
Cychrus caraboides
Cylindera cognata
Cylindera paludosa
Cylindera trisignata
Cylindra germanica
Cymindis affinis
-B chromosomes
Cymindis bedeli
Cymindis chevrolati
Cymindis cincta
Cymindis coadunata
Cymindis lineola
Cymindis scapularis
Cymindis variolosa cyanoptera
Daptus vittatus
Demetrias atricapillus
Demetrias imperialis
-Male 2n = 43-44
Dicheirotrichus obsoletus
Dicrochile brevicollis
Dinodes baeticus
Dinodes fulgidicollis
Dinodes galaecianus
Dinodes seoanei
Diplocheila latifrons
34
34
34
18
12
38
38
17
18
42
24
16
26
16
36
44
24
36
34
36
34
38
36
56
38
46
38
127
45
33
41
23
21
XY
XY
XY
NeoXY
XY
XO
XO
34
XO
XO
XXY
23
18
16
36
XXXXY
43
24
37
36
37
36
XY
37
35
44
XO
XY
37
55
37
37
37
37
XY
XY
XY
XY
17
XO
XO
XO
XO
XO
17
28
17
17
17
27
18
18
18
28
33
34
17
17
17
18
18
30
35
18
20
17
18
17
17
17
18
17
17
18
17
17
21
17
18
17
17
18
Taxa
Sexual
System
Diplocheila perscissa
Diplocheila polita
Diplocheila sp.
Diplous californicus
Ditomus calydonius
Ditomus tricuspidatus
Dixus capito obscuroides
Dixus clypeatus
Dixus sphaerocephalus
Dromius chobauti
Dromius meridionalis
Dromius piceus
Drypta argillacea
Drypta australis
-B chromosomes
Drypta dentata
Dyschirius attenuatus
Dyschirius cylindricus
Dyschirus globosus
Dyschirus importunus
Dyschirus tensicollis
Edaphopaussus favieri
Egadroma marginatum
Egadroma piceus
Egadroma smaragdulus
Egadroma smaragdulus quinquepustulatus
Elaphrus aureus
Elaphrus cupreus
Elaphrus purpurans
Elaphrus pyrenaeus
Elaphrus riparius
Elaphrus ullrichii
Ellipsoptera marginata
Elliptica lugubris
Eotachys bistriatus
Epaphius secalis
Eucryptotrichus pineticola
Eudalia macleayi
Euleptus ooderus
Eurylichnus blagravei
Euthenarus promptus
Eutrichopus canariensis
Galerita bicolor
Gehringia olympica
-Male 2n = 18-20
Graphipterus serrator
Haptoderus cantabricus
Haptoderus cf. nemoralis
Haptoderus sp
Harpalus aesculans
Harpalus affinis
Harpalus albanicus
Harpalus anxius subcylindricus
Harpalus atratus
Harpalus attenuatus
Harpalus compar
Harpalus contemptus
Harpalus decipiens
Harpalus dimidiatus
Harpalus distinguendus
Harpalus ebeninus
Harpalus faunus
Harpalus honestus
Harpalus indicola
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
40
42
38
38
58
39
42
37
37
57
59
69
45
55
25
25
25
33
34
XO
XY
XO
XO
XO
32
22
26
30
26
28
XY
XY
XY
39
XO
39
XO
31
32
33
32
32
33
31
23
22
24
28
37
33
37
35
26
34
30
19
XO
18
XO
8
37
37
37
37
37
37
XO
37
37
37
37
37
35
37
37
32
37
37
XY
XO
XO
70
46
56
26
26
34
34
32
22
26
40
50
40
38
40
39
32
34
32
32
24
24
28
38
38
36
26
30
8
38
38
38
38
38
37
38
38
38
38
38
36
38
38
32
38
38
128
XO
XO
XO
XO
XO
XO
XY
XO
XY
XO
XXXXY
XXXY
XY
XY
XO
XO
XO
XY
XY
XO
XO
18
XO
XO
XO
XO
XO
XO
XO
XO
NeoXY
XO
XO
18
18
18
17
17
17
18
18
18
17
17
18
18
21
18
18
18
17
17
17
17
18
17
18
6
17
6
17
17
17
6
28
36
17
17
18
21
17
21
21
17
18
17
18
17
17
17
18
18
17
18
18
18
18
18
18
18
18
18
18
18
Taxa
Sexual
System
Ploidy
Harpalus lethierryi aeculanus
Harpalus microthorax
Harpalus neglectus
Harpalus oblitus patruelis
Harpalus opacipennis
Harpalus pennsylvanicus
Harpalus pygmaeus
Harpalus rubripes
Harpalus rufipes
-B chromosomes
Harpalus serripes
Harpalus sp
Harpalus sp.
Harpalus sulphuripes
Harpalus tridens
Harpalus wagnerii
Laemostenus complanatus
Laemostenus oblongus
-Female 2n = 30-60
Lebia atriventris
Lebia cyanocephala
Lecanomerus sp. 1
Lestichus chloronotus
Licinus aequatus
Licinus punctatulus granulatus
Licinus punctatulus punctatulus
Lionychus albonotatus
Lionyehus albonotatus
Lophyra catena
Lophyra flexuosa
Lophyridia lunulata
Loricera pilicornis
Loxandrus sp. 1
Loxandrus sp. 2
Loxandrus sp. 3
Loxoncus microgonus
Macrocheilus trimaculatus
Mantichora amigdaloides
Masoreus wetterhallii
Mecyclothorax lewisensis
Mecyclothorax punctipennis
Megacephala brasiliensis
Megacephala cruciata
Megacephala euphratica
Megacephala rutilans
Megacephala sobrina
Melanius anthracinus
Melanius gracilis
Melanius minus
Melanius nigrita
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
38
38
38
38
38
38
37
37
37
37
37
37
39
37
37
XO
XO
XO
XO
XO
XO
38
40
38
38
38
38
30
40
45
37
39
38
38
37
37
30
40
XO
XO
XY
XY
XO
XO
XY
XY
18
18
17
18
18
18
18
17
17
32
34
34
38
38
26
26
34
34
31
33
XO
XO
37
38
XY
XY
33
33
23
22
22
19
35
35
33
38
34
38
35
23
16
12
31
31
23
25
29
37
37
37
43
51
41
40
40
46
43
46
31
31
31
31
23
22
XO
XY
18
18
XO
XO
XXY
XXXY
XXXY
18
17
21
21
17
38
38
26
26
24
24
36
36
38
34
38
36
24
16
12
32
32
24
26
30
38
38
38
42
40
40
46
Melanius rhaeticus
Microlestes abeillei
Microlestes gallicus
Microlestes luctuosos
Microlestes negrita
Myriochile fastidiosa
Myriochile melancholica
32
32
32
32
129
XO
XO
XO
XO
Xyp
XY
XY
XO
XO
XY
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XY
XY
XO
XO
XO
XO
XXY
XXXY
17
17
18
18
18
18
17
18
17
17
18
18
31
18
18
21
21
21
18
18
20
17
21
21
37
38
17
38
38
38
16
16
16
16
16
16
16
16
16
16
16
17
18
18
18
18
30
Taxa
Myriochile undulata
Mystropomus subcostatus
Nebria asturiensis
Nebria belloti
Nebria brevicollis
Nebria dilatata
Nebria expansus
Nebria fulvibarbis
Nebria gyllenhali
-Male 2n = 34-35
Nebria livida
Nebria picicornis
Nebria rubicunda
Nebria salina
Neocollyris crassicornis
Neocollyris fuscitarsis
Neocollyris redtenbacheri
Neocollyris sp. 1
Nesarpalus fortunatus
Nesarpalus sanctaecrucis
Notagonum sp. 1
Notagonum submetalicum
Notiobia germari
Notiobia melanaria
Notiobia schnusei
Notiophilus quadripunctatus
Notonomus bodeae
-B chromosomes
Notonomus flos
Notonomus hopsoni
Notonomus macoyi
Notonomus marginatus
Notonomus masculinus
Notonomus mediosulcatus
Notonomus muelleri
-B chromosomes
Notonomus obscurus
Notonomus opulentus
-B chromosomes
Notonomus peronii
Notonomus philippi
Notonomus phillipsii
-B chromosomes
Notonomus pluripunctatus
Notonomus rainbowi
Notonomus satrapa
Notonomus taylori
Notonomus triplogenioides
Notonomus variicollis
Odacantha melanura
Odontocarus cephalotes
Odontocheila confusa
-B chromosomes
Odontocheila nodicornis
Olisthopus glabratus
Omaseidius melanarius
Omaseus atterrimus
Sexual
System
Ploidy
Female
2n
38
28
42
38
Sex Chrom. Source
Karyotype
25
43
42
38
30
52
46
40
35
XXY
34
46
46
42
27
27
27
28
37
37
28
XXY
XXY
XXY
XXXY
XO
XY
18
21
17
17
18
18
17
17
17
17
17
17
18
17
17
17
20
17
18
21
21
21
21
18
18
21
41
37
37
24
37
XO
XO
35
35
35
35
35
XO
33
XO
21
21
21
21
21
21
21
30
36
29
35
XO
XO
21
21
30
36
46
30
35
45
XY
XO
XO
21
21
21
36
36
36
35
35
35
35
XO
XO
XO
35
24
40
22
XO
21
21
21
21
21
21
17
17
39
35
41
37
37
41
37
XO
37
37
36
36
19
XO
XO
XY
XY
XO
38
36
36
36
36
56
34
42
36
22
36
38
38
42
Omaseus elongatus
-B chromosomes
Omaseus nigerimus
Omophron limbatum
Omophron ovale
Omphra sp
Male
2n
38
38
36
36
20
130
XO
XO
XO
XO
XY
XO
XO
XO
39
18
16
16
16
16
16
16
17
17
17
Taxa
Sexual
System
Omus californicus
Omus dejeani
Oodes helopiodes
Oodes modestus
Oodes monticola
Oodes parallelus
Ophonus ardosiacus
Ophonus azureus
Ophonus cordatus
Ophonus diffinis
Ophonus incisus
Ophonus longicollis
Ophonus parallelus
Ophonus rufibarbis
Ophonus rufipes
-B chromosomes
Ophonus sabulicola hispanus
Ophonus similis
Ophonus stictus
Ophonus subquadratus
Ophonus subsinuatus
Orthomus barbarus
Orthomus expansus
Orthomus sp
Orthomus velocissimus pardoi
Orthotrichus indicus
Oxycheila tristis
Oxypselaphus obscurus
Pamborus alternans
Pamborus opacus
Pamborus tropicus
Paranchus albipes
Parophonus acutangulus
Parophonus hirsutulus
Penetretus andalusiacus
Penetretus rufipennis
Penetretus semipunctatus
Penetretus temporalis
Pentacomia sp. 1
Percosoma substriatum
Percus politus
Percus quiraoi
Pericompsus laetulus
Perileptus areolatus
Phaenaulax sp. 1
Pheropsophus africanus
Pheropsophus bimaculatus
Pheropsophus catoirei
Pheropsophus chaudoiri
Pheropsophus hilaris var. sobrinus
Pheropsophus lissoderus
Pheropsophus occipitalis
Pheropsophus verticallis
Philophloeus sp. 1
Platyderus alcaracinus
Platyderus lusitanicus
Platyderus muricianus
Platyderus testaceus
Platyderus varians
Platymetopus flavilabris
Platynini sp 1
Platynini sp 2
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
36
36
36
36
30
XY
XY
22
24
37
37
37
37
37
37
37
37
37
XY
XY
XO
XO
XO
XO
38
22
24
38
38
38
38
38
38
38
38
38
34
34
34
34
34
38
32
38
22
22
22
38
38
42
22
34
58
22
28
38
36
28
36
36
20
18
36
36
30
30
38
40
131
37
37
37
37
37
34
34
34
XY
37
29
37
22
22
37
37
41
33
33
33
32
21
34
63
57
22
27
37
32
35
36
27
35
35
19
17
35
36
29
29
23
23
23
40
37
40
30
36
XO
XO
XO
XO
XO
XY
XY
XY
16
XO
XXXXY
XO
XY
XY
XO
XO
XO
XO
XY
XO
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XO
XO
XO
XY
20
20
17
21
17
17
17
18
18
18
17
18
18
17
18
17
18
18
18
18
16
16
17
17
38
17
21
21
21
17
18
18
17
17
18
17
20
21
16
16
17
18
21
17
18
18
18
18
18
18
18
18
18
21
21
17
18
17
17
18
18
17
17
Taxa
Platynus assimile
Platynus chloreus
Platynus decentis
Platynus dorsale
Platynus extensicolle
Platynus longiventre
Platynus nugax
Platynus obscurus
Platynus ruficornis
Platynus sp (japan)
Platynus sp (mexico 1)
Platynus sp (mexico 2)
Platynus sp (mexico 3)
Platynus variablis
Platytarus bufo
Poecilus chalcites
Poecilus coerulescens
Poecilus crenatus
Poecilus cupreus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
38
38
26
44
38
27
37
37
26
44
37
28
37
37
37
37
37
43
37
43
30
37
37
37
39
43
44
37
40
37
37
29
45
38
38
31
27
37
37
43
32
37
41
22
22
22
22
22
30
47
45
35
35
XO
XO
XO
NeoXY
XY
XO
23
24
24
24
26
38
38
25
24
XXY
24
23
24
26
37
37
XXXXY
XXXY
18
XY
XXY
XY
XY
XO
XO
38
38
38
38
38
30
37
37
37
37
37
30
XO
XO
XO
XO
XO
XY
38
38
44
38
38
44
44
Poecilus cursorius
Poecilus decipens
Poecilus koyi ?
Poecilus kugelanni
Poecilus lepidus
Poecilus lucublandus
Poecilus nitidus
Poecilus purpurascens
Poecilus quadricollis
Poecilus sericeus
Poecilus versicolor
Poecilus vicinus
Pogonus chalceus
Pogonus gilvipes
Pogonus meridionalis
Pogonus persicus
Pogonus testaceus
Pristonychus terricola
Promecoderus mastersii
Promecoderus sp. 1
Prosopogmus chalybeipennis
Prosopogmus impressifrons
Prosopogmus oodiformis
Prothyma concinna
Prothymia sp. 1
Pseudochaetoera virgintiguttata
Pseudotetracha australis
Pseudotetracha blackburni-1
Pseudotetracha blackburni-2
Pseudotetracha whelani
Pterophilus brevipennis
Pterostichus aereipennis
-B chromosomes
Pterostichus aethiops
Pterostichus alacer
Pterostichus angustatus
Pterostichus anthracinus
Pterostichus argutor diligens
38
38
30
46
32
28
38
38
44
32
38
42
22
22
22
22
48
46
36
36
38
132
XO
XO
XO
XO
XO
XO
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XO
XO
XY
XY
XY
XY
XO
XO
XO
XO
18
18
17
18
18
18
17
17
18
18
17
17
17
17
17
17
16
16
17
17
16
16
16
16
16
18
16
16
18
16
16
16
18
17
16
16
16
16
18
17
17
17
17
18
21
21
21
21
21
36
20
40
40
40
29
16
16
16
16
18
16
18
16
Taxa
Sexual
System
Pterostichus argutor strenuus
Pterostichus aterrimus nigerrimus
Pterostichus bicolor
Pterostichus brevipennis
Pterostichus cantaber
-B chromosomes
Pterostichus cantabricus
Pterostichus cf dufouri
Pterostichus chyddaeus
Pterostichus coracina
Pterostichus coracinus
Pterostichus cristatus
Pterostichus cristatus cantabricus
Pterostichus diligens
Pterostichus dufouri
Pterostichus elongatus
Pterostichus flavofemoratus
Pterostichus ghiliani
Pterostichus globosus ebenus
Pterostichus gracile
Pterostichus herculaneus
Pterostichus impressus
Pterostichus insidiator
Pterostichus interruptus
Pterostichus madidus
Pterostichus melanarius
Pterostichus melas
Pterostichus metallicus
Pterostichus minor
Pterostichus morio
Pterostichus muehlfeldi
Pterostichus multipunctatus
-B chromosomes
Pterostichus nemoralis
Pterostichus niger
-B chromosomes
Pterostichus nigrita
-B chromosomes
Pterostichus nigrum
Pterostichus oblongopunctatus
Pterostichus patruelis
Pterostichus patruellis
Pterostichus pidanensis mikae
-B chromosomes
Pterostichus pidanensis pidanensis
-B chromosomes
Pterostichus pumicatus
Pterostichus rhaeticus
Pterostichus sp.
Pterostichus sp. Caucasucus
Pterostichus spinolae
Pterostichus strenuus
Pterostichus tuberculiger
Pterostichus vernalis
Pterostichus ziegleri
Risophilus himalayicus
Ropaloteres cinctus
Sarothrocrepis corticalis
Sarticus habitans
Sarticus monarensis
Scarites buparius
-B chromosomes
Ploidy
38
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
30
37
38
38
38
XO
37
37
37
16
18
XO
XO
XO
38
38
XO
38
37
37
37
37
37
37
XO
30
37
37
37
37
37
37
37
37
37
37
37
37
37
37
37
32
34
38
XY
18
16
18
16
16
16
18
16
16
16
16
16
18
16
18
18
16
16
16
38
38
37
37
XO
XO
16
18
38
37
XO
41
38
28
28
28
28
37
37
XY
28
28
28
XO
XO
18
XY
NeoXY
NeoXY
16
16
38
37
XO
16
38
38
38
38
38
37
37
37
37
37
37
30
37
46
34
39
27
33
35
45
37
XO
XO
XO
XO
XO
XO
XXXXY
XO
XO
XO
XXY
18
16
41
16
16
16
17
16
16
16
17
36
21
21
21
42
37
XXY
18
38
38
38
38
37
30
38
38
38
38
38
38
38
38
38
38
38
38
38
28
38
16
18
16
38
46
40
34
36
46
133
XO
XO
XO
XO
16
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
16
17
16
16
17
16
16
18
16
Taxa
Scarites eurytus
Scarites hespericus
Scarites inconspicuus
Scarites indus
Scarites laevigatus
Scarites occidentalis
-B chromosomes
Scarites planus
Scarites subterraneus
Scarites terricola
Scybalicus oblongiusculus
Sericoda bogemannii
Setalis rubripes
Siagona dejeani
Siagona europaea
Siagona fabricii
Siagona flessus
Siagona jenissoni
Siagona pumila
Siagonyx blackburni
Simodontus australis
Sphallomorpha albopicta
Sphodrus leucophthalmus
Spiralia maura
Stenolophus lecontei
Stenolophus mixtus
Stenolophus piceus
Stenolophus proximus
Stenolophus skrimshireanus
Stenolophus teutonus
Stomis pumicatus
Styracoderus atramentarius
Synchus vivalis
Syntomus foveatus
Syntomus fuscomaculatus
Syntomus obscuroguttatus
Syrdenus grayi
-B chromosomes
Tachys parvula
Taenidia circumdata
Teraphis sp. 1
Thalassotrechus barabarae
Therates sp. 1
Trechus flavolimbatus
Trechus latus
Trechus obtusus
Trechus pilisensis
Trechus pulchellus
Trechus pulpani
Trechus quadristriatus
Trichocellus godarti
Trichosternus relictus
Trichosternus vigorsi
Trichotichnus luparis
Trichus maculata
Triplectrus haplomus
Triplectrus rusticus
Trymosternus cordatus
Zabrus ambiguus
Zabrus angustatus
Zabrus castroi
-B chromosomes
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
46
54
52
16
52
62
39
45
53
52
15
52
61
41
XXY
XO
XO
XY
XO
XY
XO
XXY
18
42
42
18
18
18
42
18
41
39
37
57
37
36
37
45
46
46
44
45
46
48
33
37
36
22
25
36
XXY
XO
XO
XO
42
18
18
42
17
17
21
17
17
18
18
17
18
21
21
21
17
30
18
18
21
18
17
18
16
17
17
18
18
18
17
40
38
58
46
46
46
44
46
46
48
34
38
26
36
38
24
38
42
34
40
30
30
22
24
24
24
24
24
24
56
38
36
38
24
60
60
134
24
26
24
37
45
25
41
46
33
40
XO
XY
XY
XY
XO
XY
XO
XO
XXXY
XO
XY
XY
XO
XO
XO
XY
30
22
30
22
23
23
23
23
23
23
23
23
39
55
45
37
XY
XXXY
XY
XY
XXY
37
24
62
59
59
59
XO
NeoXY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
18
30
21
17
20
17
17
18
18
18
18
18
17
21
21
17
17
18
18
17
17
17
17
Taxa
Sexual
System
Ploidy
Zabrus coiffaiti
Zabrus consanguineus
Zabrus constrictus
Zabrus crassus
Zabrus curtus arragonensis
Zabrus curtus neglectus
Zabrus graecus
Zabrus gravis
Zabrus ignavus
Zabrus laevigatus
Zabrus marginicollis
Zabrus mateui
Zabrus obesus
Zabrus pecoudi
Zabrus seidlitzi
Zabrus seidlitzi gredosanus
Zabrus silphoides asturiensis
Zabrus spinipes
Zabrus theveneti
Zabrus urbionensis
Zabrus vasconicus
-B chromosomes
Zuphium olens
Cerambycidae
Acalolepta fraudatrix
Acalolepta luxuriosa
Acalolepta sejuncta
Acanthocinus aedilis
Acanthocinus circumflexus
Acanthocinus griseus
Acmaeops proteus
Aegosoma scabricorne
Aeloesthes holosericea
Agapanthia dahli
Agapanthia daurica
Agapanthia villosoviridescens
Anaesthetis testacea
Anaflus protensus
Anisarthron barbipes
Anoplophora chinensis macularia
Anoplophora malasiaca
Apomecyna neglecta
Apriona japonica
Asemum striatum
Batocera rubus
Callidium violaceum
Callipogon armillatus
Cerambyx scopolii
Chloridolum thaliodes
Chlorophorus annularis
Chlorophorus figuratus
Clytus arietis
Clytus lama
Clytus melaenus
Compsa textilis
Compsocerus equestris
Criocephalus rusticus
Cyllene robiniae
Cyrtoclytus caproides
Derancistrus thomae
Dihammus fraudator
Dihammus luxuriosus
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
60
58
50
XO
XO
XO
64
59
57
49
48
59
57
48
57
47
46
50
57
59
59
49
57
57
59
62
59
60
63
XO
17
17
17
18
17
17
17
17
17
18
18
43
17
43
17
18
17
17
17
17
17
17
30
30
XY
17
58
58
58
58
60
24
22
26
22
18
20
20
30
20
20
20
22
20
20
30
30
22
20
20
22
26
20
20
20
20
20
20
20
32
20
20
20
28
135
22
24
22
22
26
22
22
20
18
20
20
XY
20
20
22
20
20
XY
30
22
36
20
20
22
26
20
24
20
20
20
20
20
20
32
20
20
20
28
22
24
XO
XO
XO
XO
XO
XY
Xyp
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
XY
XY
Xyp
1
1
1
1
1
1
1
1
1
1
1
44
4
1
1
1
1
1
1
1
1
1
45
1
1
1
1
1
1
1
1
45
1
1
1
1
1
1
Taxa
Sexual
System
Ploidy
Female
2n
Male
2n
24
18
20
20
24
20
20
18
Dilus fugax
Distenia gracilis
Dorcadion anatolicum
Dorcadion scabricolle
Eburia 8-maculata
Elaphidion prob. Parallelus
-B chromosomes
Elaphidion villosum
Epiglenea comes
Ergates faber
Eutrapha 16-punctata
Eutrapha ocelota
Exocentrus adspersus
Exocentrus fasciolatus
Exocentrus galloisis
Exocentrus lineatus
Exocentrus lusitanus
Exocentrus punctipennis
Gaurotes doris
Gaurotes suvorovi
Glenea spilota
Gonocallus collaris
Graphisurus obsoletus
Lagochirus aranaeiformis
Leiopus stillatus
Leontium viride
Leptostylus sp.
Leptura dubia
Leptura ochracea
Leptura sanguinolenta
Liopus nebulosus
Macrotoma scutellaris
Mecynippus pubicornis
Mesechthistatus binodosus
Mesosa hirsuta
Mesosa longipennis
Mesosa myops japonica
Mesosa nebulosa
Mesosa senilis
Molorchus umbellatarum
Monochamus beloni
Monochamus galloprovinciallis galloprovinciallis
Monochamus latus
-B chromosomes
Monochamus marmorator
Monochamus notatus
Monochamus oregonensis
-B chromosomes
-B chromosomes
Monochamus saltuarius
Monochamus sartor
Monochamus scutellatus
-B chromosomes
Monochamus scutellatus-oregonensis comp
Monochamus sutor
Monochamus urussovi
Morimus orientalis
Nusphera madurensis
Oberea affinis
Oberea mandarina
Oberea oculata
Oberea sp. 1
20
18
18
20
Sex Chrom. Source
Karyotype
Xyp
Xyp
Xyp
1
1
46
46
46
1
1
18
20
22
20
20
26
Xyp
Xyp
22
22
22
22
20
14
20
24
24
26
20
20
20
20
22
28
22
22
20
22
20
20
20
20
22
22
21
Xyp
Xyp
XY
Xyp
Xyp
Xyp
XY
Xyp
Xyyp
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
47
1
20
20
22
Xyp
Xyp
Xyp
1
1
1
21
Xyyp
1
20
24
20
20
24
20
Xyp
Xyp
Xyp
47
47
1
20
18
20
20
Xyp
1
Xyp
1
20
20
26
22
22
22
20
14
20
24
20
20
22
28
20
20
20
20
20
22
22
20
20
22
24
22
20
20
20
20
136
20
24
24
22
20
20
20
20
Xyp
XY
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
48
1
1
1
4
1
Taxa
Sexual
System
Ploidy
Female
2n
Male
2n
20
22
18
18
20
32
20
20
20
20
20
20
22
18
18
19
32
20
20
20
20
20
20
26
10
20
20
22
20
20
20
24
20
20
20
20
20
22
20
28
28
20
20
20
Oberea tripunctata
Obrium brunneum
Olenocamptus bilobus
Olenocamptus sp. 1
Oncideres amputator
Palimna liturata
Phaea sp. 1
Phymatodes maaki
Phytoecia coerulescens
Phytoecia nigricornis
Plagionotus arcuatus
Plagionotus pulcher
Plectura metallica
Plocaederus obesus
Pogonocherus decoratus
Pogonocherus dimidiatus
Pogonocherus fasciculatus
Pogonocherus hispidulus
Pogonocherus hispidus
Pogonocherus penicellatus
Prionus coriarius
Psenocerus supernotatus
Pterolophia caudata
Pterolophia jugosa
Pterolophia leiopoda
Pterolophia rigida
Pterolophia zonata
Purpuricenus indus
Purpuricenus spectabilis
Pyrestes haematicus
Rhagium bifasciatum
Rhagium inquisitor
Rhagium mordax
Rhopalopus signaticollis
Rosalia btesi
Saperda calcarata adspersa
Saperda moesta
Saperda perforata
Saperda populnea
Saperda scalaris
Saphanus piceus
Spondylis buprestoides
Steirastoma brevis
Stenodontes spinibaris
Stenostola ferrea
Stenygra histrio
Stenygrinum 4-notatum
Strangalia 4-fasciata
Strangalia maculata
Stromatium barbatum
Tetraopes femoratus
Tetraopes tetropthamus
Tetropium castaneum
Tetropium fuscum
Tetropium gabrieli
Tetrops praeusta
Trachyderes striatus
Trachyderes thoracicus
Trichophorus interrogationes
Uraecha bimaculata
20
20
20
20
20
20
20
28
20
20
20
20
20
20
20
20
20
20
30
28
26
20
20
20
20
20
20
20
20
20
20
20
24
22
24
20
22
32
20
137
22
20
20
20
20
20
20
20
30
28
26
20
20
20
19
20
20
20
20
20
24
22
24
20
22
32
20
32
Sex Chrom. Source
Karyotype
XY
Xyp
Xyp
Xyp
XO
XY
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XO
XY
XY
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
49
1
1
1
1
4
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
50
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
51
51
45
1
Taxa
Sexual
System
Ploidy
Female
2n
Vadonia unipunctata
Vesperus xatari
54
-2 y chrom are homologous to a pair autosomes in the female
Xenolea nubila
Xylosteus spinolai
20
Xylotrechus smei
20
Chrysomelidae
Acalymma blandulum
20
-B chromosomes
Acalymma fairmairei
28
Acalymma gouldi
20
Acalymma innuba
20
-B chromosomes
Acalymma pallipe
22
Acalymma quadrivittatum
22
Acalymma s. sp. near trivittatum
39
-B chromosomes
Acalymma sp. 1
24
Acalymma thiemei
22
Acalymma trivittatum
22
-B chromosomes
Acalymma vittatum
22
Acanthoscelides obtectus
20
Aedmon eugeniae
40
-B chromosomes
Agathomerus sellatus
20
Agathomerus subfasciatus
20
Agathomerus testaceus
20
Agelastica coerulea
24
Agroiconota inedita
42
Agroiconota propinqua
38
Alagoasa acutangula
22
Alagoasa apicata
22
Alagoasa arcifera
16
-B chromosomes
Alagoasa bicolor
22
Alagoasa bipunctata
22
Alagoasa burmeisteri
22
Alagoasa burmeisteri gounellei
22
22
Alagoasa burmeisteri maculatissima
22
Alagoasa ceracollis
22
Alagoasa cruxnigra
Alagoasa decemguttata
22
Alagoasa dipus
22
Alagoasa dissepta cyaneofasciata
22
Alagoasa equestris
12
Alagoasa extrema
22
Alagoasa fasciaticollis
22
-B chromosomes
22
Alagoasa illustris
22
Alagoasa januaria
22
Alagoasa jufina
22
Alagoasa libentina
22
Alagoasa libentina tetraspilota
22
22
Alagoasa mella commutata
22
Alagoasa posticalis
22
Alagoasa rotundicollis
22
Alagoasa sp. 1
22
Alagoasa sp. 2
22
Alagoasa trifasciata
22
Alagoasa trifasciata vulgatissima
22
22
Altica aenea brasiliensis
24
Altica ambiens
24
Altica ambiens alni
24
138
Male
2n
Sex Chrom. Source
Karyotype
20
53
Xyy
46
52
20
Xyp
1
1
1
19
XO
53
27
19
19
XO
XO
XO
53
53
53
21
21
XXY
XO
XO
53
53
53
23
21
21
XO
XO
XO
53
53
53
21
20
39
XO
XY
XO
53
1
53
20
20
20
24
42
38
22
22
16
Xyp
Xyp
Xyp
Xyr
Xyp
Xyp
Xy+
Xy+
Xy+
53
53
53
53
54
55
53
53
53
22
22
22
Xy+
22
22
22
22
22
22
12
22
22
Xy+
Xy+
Xy+
53
Xy+
Xy+
53
53
53
22
22
22
22
22
Xy+
22
22
22
22
22
22
Xy+
24
24
24
Xy+
Xy+
Xy+
Xy+
Xy+
53
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
53
Xy+
Xy+
Xy+
32
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
53
53
53
53
53
53
56
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
Taxa
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
Altica amethystina
Altica ampelophaga
Altica brevicollis
Altica carduorum
Altica carinata
Altica chalybea
Altica coerulea
24
24
24
24
24
22
24
Altica coerulescens
Altica corni
Altica lazulina
-apomixis
Altica lythri
Altica oleeracea
Altica populi
Altica rosae
Altica sp. 1
Altica sp. 2
Altica sp. 3
24
24
34
24
24
24
24
24
22
24
23
24
24
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
Xyy+
Xy+
Xy+
53
53
53
53
53
53
53
53
53
53
53
24
24
24
24
40
18
24
24
24
24
22
24
24
26
24
24
24
24
40
17
Xy+
Xy+
Xy+
Xy+
Xyp
Xy+
Xy+
XXyy+
Xy+
Xy+
Xy+
Xy+
Xyp
XO
53
53
53
53
53
53
53
53
53
53
53
53
57
53
20
17
18
32
28
19
XO
18
32
28
XO
53
NeoXY
XY
XY
53
54
53
53
28
26
28
30
28
24
24
32
28
28
26
28
30
28
24
24
32
28
XY
XY
XY
XY
XY
Xy+
XY
XY
XY
53
53
53
53
53
53
53
53
53
28
30
10
12
28
50
26
38
22
28
30
10
12
Xyp
50
26
38
22
25
27
23
22
27
18
18
23
22
25
27
28
24
XY
Xyp
XY
XY
58
XY
XY
XY
Xy+
XXXXY
XXXXXXY
XXY
Xy+
XXXXXXY
Xy+
Xy+
XXY
53
53
53
53
Altica subplicata
Altica tamaricis
Altica tincta?
Altica ulmi
Ambrostoma superbum
Amphelsasma cavum
-B chromosomes
Amphelsasma cavum n. sp.
Amphelsasma cavum vicinum
Anisodera excavata
Aphthona cyparissiae
Aphthona depressa
-B chromosomes
Sexual
System
parth
Ploidy
3
24
24
24
24
22
24
24
18
Aphthona flaviceps
Aphthona herbigrada
Aphthona illigeri
Aphthona lamprocyanea
Aphthona lutescens
Aphthona nigriceps
Aphthona venustula
-B chromosomes
Apophylia sericea
Apraea portoricensis
Araucanomela wellingtonensis
Argopus punctipennis
Arrhenocoela lineata
Asiorestia impressa
Asphaera abdominalis
Asphaera abendrothi
Asphaera daniella
Asphaera limitata
Asphaera meticulosa
Asphaera nr. Weyrauchi sp. 1
Asphaera octopunctata
Asphaera pauperata
Asphaera quadrifasciata
Asphaera reflexicollis
Asphaera scutata
Asphaera semifulva
Asphaera sp. 2
28
22
18
18
32
34
139
XXXXY
XXXXXXY
XXXXXXXY
XXXY
53
53
53
53
56
56
56
53
56
53
53
56
53
56
56
56
56
Taxa
Sexual
System
Ploidy
Asphaera sp. 3
Asphaera t-album
Asphaera unicolor
Aspidimorpha difformis
Aspidimorpha dorsata
Aspidimorpha furcata
Aspidimorpha indica
Aspidimorpha miliaris
Aulacophora cincta
Aulacophora femoralis
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
27
22
18
16
18
18
18
30
58
59
30
30
59
30
57
57
59
23
18
28
28
24
18
28
32
51
47
27
41
44
XO
44
30
XXXXXXXY
XXXXXXY
Xy+
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
XXY
44
XXXY
54
53
16
22
16
22
Xyp
Xyp
20
20
22
20
20
22
20
22
20
22
38
24
20
20
22
20
20
22
20
22
20
22
37
23
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
XO
XO
53
1
60
1
1
1
1
1
1
1
1
1
1
1
53
23
XO
23
XO
24
24
24
23
23
23
23
XO
XO
XO
53
53
53
53
24
24
Xyp
53
22
18
16
18
18
18
30
58
Aulacophora fovecollis
30
30
Aulacophora intermedia
Aulacophora semifusca
Aulacoscelis melanocera
Baliosus californicus
Basilepta balyi
Basilepta fuvipes
Basilepta obliteratum
Basiprionota decemmaculata
Blaptea elguetai
Blepharida rhois
Botanochara angulata
Botanochara bonariensis
Botanochara duodecimverrucata
24
18
28
28
24
18
28
32
32
31
Botanochara sp
-B chromosomes
Bromius obscurus
parth
-apomixis syn. Adoxus obscurus
Bruchidius albizziae
Bruchidius angustifrons
Bruchidius aureus
Bruchidius dimorphous
Bruchidius mimosae
Bruchidius multilineolatus
Bruchidius saundersi
Bruchidius sp. 1
Bruchidius urbanus
Bruchus pisorum
Bruchus rufimanus
Calligrapha alni
-B chromosomes
Calligrapha alnicola
parth
Calligrapha amator
Calligrapha apicalis
parth
Calligrapha bidenticola
-B chromosomes
Calligrapha californica corepsivora
Calligrapha confluens
Calligrapha fulvipes
Calligrapha multipunctata bigsbyana
-B chromosomes
Xyp
XXY
XY
XXY
XXY
XXY
XO
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XXY
XXY
Xyy
Xyy
XXXY
54
XXXY
XXXY
24
48
24
48
24
140
56
56
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
59
53
54
54
54
54
54
54
54
53
53
53
53
Taxa
Calligrapha ostryae
Calligrapha philadelphica
-B chromosomes
Calligrapha pnirsa
Calligrapha polyspila
Calligrapha pruni
Calligrapha rowena
-B chromosomes
Calligrapha scalaris
Calligrapha verrucosa
Calligrapha vicina
Calligrapha virginea
Callosobruchus analis
Callosobruchus chinensis
Callosobruchus maculatus
Callosobruchus subinnotatus
Capraita egleri
Capraita sp. 1
Capraita strichocephala
Capraita trinidadensis
Capraita virkkii
Caryedon acaciae
Caryedon gonagra
Caryedon lineatonota
Caryedon longus
Caryedon opacus
Caryedon prosopius
Caryedon serratus
Caryedon sp. 1
Caryedon sp. 2
-Male 2n = 20-26
Cassida algirica
Cassida azurea
Cassida bergeali
Cassida circumdata
Cassida deflorata
Cassida enervis
Cassida flaveola
Cassida fuscorufa
Cassida hemisphaerica
Cassida hexastigma
Cassida inquinata
Cassida leucanthemi
Cassida margaritacea
Cassida nebulosa
Cassida pannonica
Cassida panzeri
Cassida prasina
Cassida pusilla
Cassida rubiginosa
Cassida rufovirens
Cassida sanguinolenta
Cassida sanguinosa
Cassida sp. 1
Cassida subreticulata
Sexual
System
parth
parth
parth
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
48
24
23
XO
53
53
24
24
24
24
24
23
23
24
23
23
XO
XO
Xyp
XO
XO
53
53
53
53
53
23
23
XO
XO
20
20
20
19
20
21
20
19
20
20
20
22
22
22
22
XY
Xyp
XY
XO
Xyp
Xyyp
XY
XO
XY
Xyp
XY
Xy+
Xy+
Xy+
Xy+
18
18
20
18
18
18
18
20
23
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
53
53
53
53
53
61
1
61
1
62
62
61
1
1
1
61
53
53
53
53
53
1
1
1
1
1
1
1
1
1
18
18
18
18
18
18
18
18
20
18
18
18
30
18
20
18
18
18
18
18
18
18
20
21
30
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyyp
Xyp
63
63
63
53
63
53
53
53
63
63
53
63
53
4
63
63
53
53
63
63
63
63
53
53
63
44
24
24
48
48
20
20
20
20
20
20
20
20
20
20
22
22
22
22
22
18
18
20
18
18
18
18
20
23
18
18
18
18
18
18
18
20
18
18
18
30
18
20
18
18
18
18
18
18
18
20
30
141
Taxa
Sexual
System
Cassida varians
-B chromosomes
Cassida vibex
Cassida virdis
Cassida virdis 1
Cassida Virdis 2
Cassida Virdis 3
Cassida vittata
Cassidinae sp
Cerotoma atrofasciata
-B chromosomes
Cerotoma ruficornis
Cerotoma sp. 1 in variegata complex
Chaetocnema chlorophana
Chaetocnema concinna
Chaetocnema conducta
Chaetocnema depressa
Chaetocnema major
Chaetocnema tibialis
Chalcophana cincta
Chalcophana sp.
Chalcophana sp. 1
Chalcophana verecunda
Chalepus dorsalis
Chalepus inequalis
Chalepus sanguinicollis
Chalepus sp. 1
Charidotella immaculata
Charidotella quadrisignata
Charidotella sexpunctata
Charidotella sexpunctata
Chelymorpha cassidea
Chelymorpha cribraria
Chelymorpha indigesta
Chelymorpha nigricollis
Chelymorpha varians
Chirida sp. 1
Chiridopsis bupunctata
Chlamisus straminea
Chlamydocassis metallica
Chrysochus asclepiadeus
Chrysochus auratus
Chrysochus cobaltinus
Chrysodina sp.
Chrysolina affinis
Chrysolina americana
Chrysolina analis
Chrysolina aurichalcea
Chrysolina banksi
Chrysolina bicolor
Chrysolina bigorrensis
Chrysolina carnifex
Chrysolina cerealis
Chrysolina cf. Subcostata
Chrysolina coerulans
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
18
Xyp
64
20
18
24
30
16
24
30
20
22
34
20
18
24
30
16
24
30
20
22
33
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
XY
XO
53
4
63
63
4
4
4
53
54
53
34
22
20
22
22
26
22
22
26
26
26
26
26
16
18
18
18
22
24
22
22
22
22
22
22
22
36
18
33
21
20
22
22
26
22
22
26
26
26
26
26
16
18
18
18
22
24
22
22
22
22
22
22
22
36
18
21
32
24
26
29
12
24
24
40
40
31
41
32
45
40
46
23
24
26
40
40
24
26
24
XO
XO
Xyp
XY
Xyp
Xyr
Xyp
Xyr
Xyp
NeoXY
NeoXY
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XXY
Xyp
Xyp
Xyc
XO
Xyp
Xyp
Xyp
Xyp
Xyp
53
53
53
53
53
53
53
53
53
45
53
53
45
54
53
55
53
54
55
53
55
53
55
53
53
53
53
53
53
53
53
53
53
45
53
53
65
53
53
53
53
53
53
53
53
53
65
65
53
53
65
4
32
24
26
30
12
24
24
40
40
32
42
46
40
46
24
24
26
40
40
24
26
24
142
XO
Xyp
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Taxa
Sexual
System
Ploidy
Chrysolina colasi
Chrysolina colasi
Chrysolina curvilinea
Chrysolina diluta
Chrysolina exanthematica
Chrysolina fastuosa
Chrysolina femoralis
Chrysolina fragariae
Chrysolina gebleri
Chrysolina gemina
Chrysolina geminata
Chrysolina globosa
Chrysolina graminis
Chrysolina grossa
Chrysolina gypsophilae grossepunctata
Chrysolina haemoptera
Chrysolina helopioides
Chrysolina herbacea
Chrysolina interstincta
Chrysolina janbechynei
Chrysolina kuesteri
Chrysolina lucida
Chrysolina marcasitica
Chrysolina marginata
Chrysolina marginata
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
24
40
36
24
24
24
24
26
24
24
24
40
36
23
24
24
24
26
24
38
26
24
24
30
40
47
24
40
40
22
24
42
40
40
40
47
23
40
46
24
23
24
24
24
40
23
23
23
23
22
47
47
30
32
24
34
34
Xyp
34
34
35
34
34
34
24
24
24
24
24
24
24
40
27
28
Xyp
Xyp
Xyp
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
26
24
24
30
40
48
24
40
40
22
24
42
40
40
40
48
24
40
46
24
24
24
24
24
40
Chrysolina obscurella
Chrysolina obsoleta
Chrysolina oricalcia
Chrysolina peregrina
Chrysolina polita
Chrysolina purpurascens
Chrysolina purpurascens crassimargo
Chrysolina purpurascens purpurascens
Chrysolina pyrenaica
Chrysolina rufa
24
Chrysolina rutilans
Chrysolina staphylea
Chrysolina timarchoides
Chrysolina umbratilis
Chrysolina varians
Chrysolina veridana
Chrysomela interrupta
Chrysomela lapponica
Chrysomela mainensis interna
Chrysomela populi
Chrysomela schaefferi
Chrysomela sp. 1
Chrysomela sp. 2
Chrysomela tremulae
Chrysophtharta aurea
Chrysophtharta decolorata
Chrysophtharta nobilitata
Chrysophtharta variicollis
Clytra laeviscula
Clytra laeviuscula
Clytra quadripunctata
Clytra succinata
Colaphus sophiae
Colaspidema barbarum
34
24
22
48
48
30
32
24
34
34
34
34
34
34
34
34
24
24
24
24
24
24
40
28
28
143
Xyp
Xyp
Xyp
Xyp
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XO
XO
Xyp
Xyp
Xyp
XO
Xyp
Xyp
Xyp
Xyp
XO
XO
Xyp
XO
XO
Xyp
Xyp
Xyp
Xyp
Xyp
53
Xyp
Xyp
Xyyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
XO
Xyp
57
53
65
53
53
4
53
53
57
53
53
65
4
53
53
53
65
53
65
53
53
53
53
57
65
53
65
53
53
53
4
65
53
65
65
53
65
53
53
53
53
65
53
65
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
4
53
53
44
57
Taxa
Colaspis greyi
Colaspis tricolor
Colasposoma auripenne
Colasposoma lividipes
Colasposoma metallicum
Colasposoma ornatum
Colasposoma rufipes
Colasposoma semicostatum
Colasposoma sp. 1
Conchyloctenia nigrovittata
Conicobruchus indicus
Coptocephala floralis
Coptocephala unifasciata
Coptocycla adamantina
Coraia clarki
Coraia maculicollis
Cosmogramma decora
Craspedonta leayana
Crepidedera aurea
Crepidedera plutus
Crioceris asparagi
Crioceris duodecimpunctata
Crioceris macilenta
Crioceris paracenthesis
Crosita altaica
Crosita rugulosa
Crosita salviae
Cryptocephalus alboscutellatus
Cryptocephalus analis
Cryptocephalus aureolus
Cryptocephalus bipunctatus
Cryptocephalus capucinus
Cryptocephalus crassus
Cryptocephalus espanoli
Cryptocephalus fulvus
Cryptocephalus globicollis
Cryptocephalus hypochoeridis
Cryptocephalus lusitanicus
Cryptocephalus moraei
Cryptocephalus ocellatus
Cryptocephalus ochroleucus
Cryptocephalus octopilosus
Cryptocephalus oppositus
Cryptocephalus pominorum
Cryptocephalus primarius
Cryptocephalus quadruplex
Cryptocephalus rugicollis
Cryptocephalus sexmaculatus
Cryptocephalus sexpunctatus
Cryptocephalus sexpustulatus
Cryptocephalus sexsignatus
Cryptocephalus sp. 1
Cryptocephalus sp. 2
Cryptocephalus sp. 3
Cryptocephalus sulphureus
Cryptocephalus triangularis
Cryptocephalus violaceus
Cryptocephalus vittula
Cteisella confusa
Cyrsylus volkameriae
Cyrtonus arcasi
Cyrtonus contractus
Cyrtonus cupreovirens
Cyrtonus cylindricus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
30
16
16
16
16
16
16
16
16
18
20
22
30
16
16
16
16
16
16
16
16
18
20
24
24
18
36
36
20
18
18
26
16
16
16
16
30
30
24
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
24
18
36
36
20
18
18
26
16
16
16
16
30
30
24
30
30
30
30
30
32
30
30
30
30
30
30
32
30
40
24
30
28
16
30
30
30
30
30
32
32
30
28
18
40
28
28
28
144
30
30
30
30
32
30
32
30
30
30
30
30
30
32
30
30
40
24
30
28
16
30
30
30
30
30
32
32
30
28
18
30
40
28
28
28
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
66
Xyp
Xyr
Xyr
Xyr
Xyr
Xyr
Xyr
Xyr
Xyr
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
Xyp
Xyp
Xyr
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
Xyp
Xyp
Xxyy
Xyp
Xyp
Xyp
Xyp
53
53
53
53
53
53
53
53
53
54
54
1
66
4
53
53
53
45
53
53
53
53
53
53
53
57
57
53
53
53
53
53
53
53
53
53
53
66
53
53
53
53
53
66
53
53
53
53
53
53
53
53
53
53
53
53
53
53
54
53
67
67
67
67
Taxa
Sexual
System
Ploidy
Cyrtonus dufouri
Cyrtonus elegans
Cyrtonus fairmarei
Cyrtonus majoricensis
Cyrtonus pardoi
Cyrtonus plumbeus
Cyrtonus puncticeps
Cyrtonus punctipennis
Cyrtonus rotundatus
Cyrtonus ruficornis
Cyrtonus sp. 1
Dactylispa atkinsoni
Dactylispa brevispinosa
Dactylispa humeralis
Deloyala cruciata
Deloyala guttata
Desmogramma nigripes
Deuteronoda suturalis
Diabrotica adelpha
Diabrotica balteata
Diabrotica bioculata
Diabrotica capitata
Diabrotica circulata
Diabrotica cortezi
Diabrotica cristata
Diabrotica dissimilis
Diabrotica flaviventris
Diabrotica graminea
Diabrotica lemniscata
-B chromosomes
Diabrotica limitata 15-punctata
Diabrotica longicornis
-B chromosomes
Diabrotica longicornis barberi
-B chromosomes
Diabrotica longicornis nigricornis (undesc.)
-XX B chromosomes
Diabrotica marginata
Diabrotica nummularis
Diabrotica ochreata
Diabrotica octoplagiata
Diabrotica porracea
Diabrotica rufolimabata
Diabrotica scutella
-B chromosomes
Diabrotica sexmaculata
Diabrotica speciosa
Diabrotica tibialis
-B chromosomes
Diabrotica trifasciata
Diabrotica undecimpunctata
-B chromosomes
Diabrotica undecimpunctata duodecimnotata
Diabrotica undecimpunctata howardi
-B chromosomes
-B chromosomes
Diabrotica undecimpunctata tenella
-B chromosomes
Diabrotica variablis?'
Diabrotica virgifera
Diabrotica virgifera zeae
Diabrotica viridula
20
20
20
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
28
40
28
28
46
28
40
28
28
28
28
16
16
16
18
18
20
24
16
20
20
20
20
20
20
20
20
20
20
20
28
28
40
28
28
46
28
40
28
28
28
28
16
16
16
18
18
20
24
16
19
19
19
19
19
19
19
19
19
19
19
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
53
67
67
67
53
67
67
67
53
67
67
67
53
53
53
54
54
55
53
53
53
53
53
53
53
53
53
53
53
53
53
19
20
XO
19
53
XO
53
19
XO
53
20
XXOO
53
20
20
20
20
20
20
20
19
19
19
19
19
19
19
XO
XO
XO
XO
XO
XO
XO
53
53
53
53
53
53
53
20
22
20
19
21
19
XO
XO
XO
53
53
53
20
20
19
19
XO
XO
53
53
19
20
XO
19
53
XO
53
20
19
XO
53
20
19
XO
53
20
20
20
20
19
19
19
19
XO
XO
XO
XO
53
53
53
53
145
Taxa
Diabrotica viridula rufobasis
Diapromorpha melanopus
Diapromorpha pallens
Diapromorpha quadripunctata
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
19
22
24
22
XO
22
24
Xy+
23
22
23
32
40
16
16
40
30
30
30
50
63
65
34
35
46
44
34
48
29
29
30
30
30
30
28
28
30
12
26
26
22
18
26
26
26
18
22
22
22
22
20
18
18
31
24
44
44
39
53
XY
XY
53
Xyy+
Xy+
Xyy+
NeoXY
Xyp
Xyp
Xyp
46
38
38
40
44
34
34
34
34
34
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
22
Diapromorpha turcica
22
Dibolia penstemonis
Dibolia timida
Dicladispa occator
Dicladispa testacea
Diorhabda lusca
Diphaulaca arguta
Diphaulaca sp. 1
Diphaulaca sp. 2
Disonycha alternata
Disonycha bicarinata
32
40
16
16
Disonycha brevilineata
Disonycha glabrata
Disonycha laevigata
Disonycha nigrita
Disonycha sp. 1
Disonycha sp. 2
Disonycha spilotrachela
Donacia andalusica
Donacia bicolor
Donacia biimpressa
Donacia clavipes
Donacia hirticollis
Donacia subtilis
Donacia vulgaris
Doryphora quadrisignata
Entomoscelis adonidis
Entomoscelis sacra
Epimela indet. sp.
Epistictina viridimaculata
Epitrix cucumeris
Epitrix fasciata
Erganoides sp. 1
Estignema chinensis
Eumolpus sp.
Eumolpus sp. 1
Eumolpus sp. 2
Eumolpus surinamensis
Eurypedus thoni
Eurypepla jamaicensis
Exema canadensis
Exora encaustica narensis
Forsterita sp. 1
Galeruca angusta
Galeruca artemisiae
Galeruca canigouensis
-Male 2n = 38-40
Galeruca indica
Galeruca miegi
Galeruca monticola
Galeruca tanaceti
Galerucella bicolor
Galerucella bimarnica
Galerucella calmariensis
Galerucella cavicollis
Galerucella lineola
Galerucella nymphae
30
30
30
50
46
44
48
30
30
30
30
30
28
28
30
12
26
26
18
26
26
26
18
22
22
22
22
20
18
18
32
24
44
44
46
38
38
40
44
34
34
34
34
34
146
Xyp
Xyp
Xyp
NeoXY
XXY
XXY
XXOO
XXXXXXY
Xyp
NeoXY
XXOO
XY
XO
XXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
XO
Xyp
Xyp
Xyp
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
45
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
54
53
53
53
53
45
53
53
45
45
55
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
Taxa
Sexual
System
Ploidy
Galerucella placida
Galerucella pusilla
Galerucella sp. 1
Galerucella tenella
Galerucella vaccinii
Gastrophysa cyanea
Gastrophysa poligoni
Gastrophysa viridula
Genus nr. Stenophyma sp. 1
Glyphocassis trilineata
Glyptina sp. 1
Glyptoscelis chontalensis
Glyptoscelis sp. 1
Gonioctena americana
Gonioctena linnaeana
Gonioctena nivosa
Gonioctena quinquepunctata
Gonioctena sibirica
Gonioctena springlovae
Gonioctena variabilis
Gonioctena viminalis
Gratiana lutescens
Gratiana pallidula
Gratiana spadicea
Gynandrobrotica lepida
Gynandrobrotica nigrofasciata
-B chromosomes
Gynandrobrotica sp. 1
-B chromosomes
Gynandrobrotica variabilis
Heikertingerella brevitarsis
Heikertingerella sp. 1
Heikertingerella sp. 2
Heikertingerella sp. 3
Helocassis clavata
Hemipyxis sp. 1
Henicotherus porteri
Hermaeophaga cicatrix
Hermaeophaga ruficollis
Hilarocassis exclamationis
Hispa armigera
Hispa atra
Homoschema nigriventre
Homoschema obesum
Hydrothassa glabra
Hyphasoma sp. 1
Hypocassida subferruginea
Hypolampsis bryanti
Hypolampsis sp. 1
Isotes multipunctata
Isotes quadrispilota
Jolivetia obscura
Kanarella sp. 1
Labidomera clivicollis
-Subsequent studies failed to replicate
Labidomera suturella
Laccoptera quadrimaculata
18
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
38
16
34
26
34
30
24
28
32
16
18
28
20
16
24
24
24
24
24
40
24
24
18
18
20
18
17
38
16
34
26
34
30
24
28
32
16
18
28
20
16
24
24
24
24
24
40
24
24
18
18
20
17
XO
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xy+
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XO
53
53
53
53
53
53
53
53
53
53
54
53
53
53
53
53
53
53
53
53
53
53
53
53
53
54
53
14
13
XO
53
18
34
22
38
20
22
24
38
28
26
33
17
24
20
18
18
18
18
24
28
17
16
34
34
12
18
16
8
11
34
22
38
20
22
23
37
28
26
33
XO
XXYY
NeoXY
XXYY
XXYY
XY
Xyp
XY
Xyp
XXY
XY
Xyp
Xyc
XY
XY
Xyp
XY
XXY
Xyp
NeoXY
NeoXY
XY
Xyp
XO
XO
Xyp
Xyp
XY
53
53
53
53
53
53
53
53
59
53
68
53
55
53
53
53
53
53
53
53
53
53
53
53
53
69
53
53
34
32
18
18
33
31
18
18
XO
XO
XY
Xyr
70
70
54
54
18
18
Xyp
53
20
18
18
24
28
16
34
34
12
18
16
8
-B chromosomes
147
Taxa
Lachnaia pubescens
Lasiochila excavata
Lema cerea
Lema cirsicola
Lema coromandeliana
Lema cyanea
Lema cyanella
Lema dorsalis
Lema n.sp.
Lema nigripes
Lema nigrrofrontalis
Lema praeusta
Lema rufotestacea
Lema semifulva
Lema terminata
Lema trilineata
Leptinotarsa behrensi
Leptinotarsa belti
Leptinotarsa decimlineata
Leptinotarsa defecta
Leptinotarsa flavitarsis
Leptinotarsa haldemani
Leptinotarsa heydeni
Leptinotarsa juncta
Leptinotarsa lineolata
Leptinotarsa peninsularis
Leptinotarsa rubiginosa
Leptinotarsa signaticollis
Leptinotarsa texana
Leptinotarsa tumamoca
Leptinotarsa typographica
Leptinotarsa undecimlineata
Leptispa filiformis
Leucocera laevicollis
Lilioceris impressa
Lilioceris lilii
Linaeidea aenea
Lochmaea capreae
Longitarsus australis
Longitarsus ballotae
Longitarsus candidulus
Longitarsus codinai
Longitarsus exsoletus
Longitarsus holsaticus
Longitarsus kleiniiperda
Longitarsus luridus
Longitarsus lycopi
Longitarsus melanocephalus
Longitarsus nanus
Longitarsus nervosus
Longitarsus niger
Longitarsus nigrofasciatus
Longitarsus oakleyi
Longitarsus obliteratoides
Longitarsus ochroleucus
Longitarsus ordinatus
Longitarsus pellucides
Longitarsus pratensis
Longitarsus reichei
Longitarsus rubiginosus
Sexual
System
Ploidy
Female
2n
24
18
14
16
24
16
18
16
18
16
16
14
16
16
32
36
24
34
36
36
36
36
36
36
36
36
36
36
34
36
36
36
34
32
24
20
22
34
28
32
30
30
32
32
30
30
28
32
28
30
26
26
28
28
30
30
28
30
26
28
30
148
Male
2n
Sex Chrom. Source
Karyotype
19
24
18
14
18
16
24
16
18
16
18
16
16
14
16
16
32
35
23
33
35
36
35
35
35
35
35
35
35
35
33
35
35
35
33
32
24
20
20
22
34
34
28
32
30
30
32
32
30
30
28
32
28
30
26
26
28
28
30
30
28
30
26
28
30
Xyyp
Xy+
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XO
XO
XO
XO
Xyp
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
Xyp
Xyr
Xyp
Xyp
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
71
53
53
53
53
53
53
68
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
Taxa
Sexual
System
Ploidy
Longitarsus succineus
Longitarsus tabidus
Longitarus aeruginosus
Luperodes praeustus
Luperus discrepans
Lysathia ludoviciana
Lysathia occidentalis
Lysathia sp. 1
Macrocoma rufotibialis
Macrohaltica jamaicensis
Macrohaltica mexicana salvadorensis
Macrohaltica transversa
Macrolenes dentipes
Mantura rustica
Merilia sublunata
Mesoplatys cincta
Metallactus patagonicus
Metriona elatior
Microrhopala vittata
Microtheca ochroloma
Monolepta erythrocephala
Monolepta sexsignata
Monolepta signata
Monomacra caprai
Monomacra sp. 1
Monoxia batisia
Monoxia sp. 1
Neocrepidodera transversa
Nodonota aridicola
Nodonota lateralis
Nodonota puncticollis
Nodonota virdis
Notosacantha maculipennis
Octotoma scabripennis
Odontota dorsalis
Oedionychus cinctus
Oedionychus limbatus
Oedionychus sp.
Oedionychus sp. 1
Oedionychus umbratica
Oides bipunctata
Oides sp.
Omophoita aequatorialis
Omophoita aequinoctialis
Omophoita albicollis
Omophoita angustolineata ?
Omophoita annularis
Omophoita clerica
Omophoita cyanipennis
Omophoita cyanipennis octomaculata
Omophoita latitarsis
Omophoita lunata
Omophoita magniguttis
Omophoita octoguttata
Omophoita personata
Omophoita quadrinotata
Omophoita sexnotata
Omophoita sp.
Omophoita superba
Oocassida pudibunda
-Male 2n = 38-40
Ophraella communa
Ophraella conferta
Ophraella cribata
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
30
28
24
32
XY
XY
XY
NeoXY
XNeoXY
Xy+
Xy+
Xy+
Xyp
Xy+
Xy+
Xy+
22
39
30
28
24
32
33
24
24
24
16
24
26
24
22
30
22
24
15
18
32
28
26
18
18
18
30
33
36
18
38
12
12
12
16
16
20
16
16
16
22
22
22
17
49
22
22
22
Xy+
22
23
22
22
22
24
22
22
22
22
22
22
22
39
24
24
34
24
24
24
24
24
16
24
26
24
22
22
24
16
18
32
28
26
18
18
18
30
34
36
18
38
12
12
12
16
16
20
16
16
16
22
22
22
18
50
22
22
22
22
22
22
22
22
24
22
22
22
22
22
149
Xy+
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
Xyp
NeoXY
XO
XY
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xy+
Xy+
XY
Xy+
Xy+
XO
XO
Xy+
Xy+
Xy+
53
Xy+
Xyy+
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
Xy+
53
53
68
53
53
53
53
53
53
53
53
68
66
53
53
53
45
54
53
53
53
53
53
53
53
53
53
53
4
53
53
53
53
53
53
53
68
68
45
53
53
53
53
53
53
53
Xy+
Xyp
53
53
53
53
53
53
72
53
53
53
72
72
53
53
NeoXY
NeoXY
66
53
53
Taxa
Ophraella notata
Ophraella notulata
Ophraella pilosa
Ophraella sexvittata
Ophrida marmorea
Oreina alpestris
Oreina cacaliae
Oreina elongata
Oreina fairmairiana
Oreina ganglbaueri
Oreina intricata
Oreina variabilis
Oreina virgulata
Oulema astrosuturalis
Oulema downesi
Oulema gallaeciana
Oulema hoffmannseggi
Oulema melanopus
Oulema tristis
Oxylepus deflexicollis
Pachybrachis anoguttatus
Pachybrachus azuereus
Pachybrachus bivittatus
Pachybrachus catalonicus
Pachybrachus melanostrictus
Pachybrachus peccans
Pachybrachus petitpierrei
Pachymerus chinensis
Paranaita bilimbata
Paranaita opima
Paranaita sanguinipes
Paranapiaca connexa
Paranapiaca significata
Paranapiaca tricincta
Parasyphraea sp. 1
Paratriarius curtisii
Paratriarius dorsata
Paria sp. 1
Paria sp. 2
Paridea sp. 1
Paropsis aegrota
Paropsis atomaria
Paropsis porosa
Paropsis rubidipes
Paropsisterna beata
Paropsisterna nucea
Paropsisterna octosignata
Pataya nitida
Phaedon affine
Phaedon cochleariae
Phaedon consimile
Phaedon cyanopterum
Phaedon tumidulus
Phanaeta aida
Phenrica aequinoctialiformis
Phenrica austriaca
Philhydronopa sp. 1
Phratora laticollis
Phratora polaris
Phratora tibialis
Phratora vitellinae
Phratora vulgatissima
Phyllotreta diademata
Phyllotreta fallax
Phyllotreta nemorum
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
36
36
34
24
32
24
24
24
24
24
24
24
24
20
22
16
16
16
16
18
36
36
34
24
32
24
24
24
24
24
24
24
24
20
22
16
16
16
16
18
16
16
16
16
16
16
16
20
22
22
22
19
19
23
20
23
19
16
16
40
24
24
24
24
24
24
24
38
34
34
34
34
34
20
41
48
24
34
34
34
34
34
30
32
32
XY
XY
XY
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
16
16
16
16
16
16
20
22
22
22
20
20
24
20
24
20
16
16
40
24
24
24
24
24
24
24
38
34
34
34
34
34
20
24
34
34
34
34
34
30
32
32
150
Xyr
Xyr
Xyr
Xyr
Xyr
Xyr
Xyp
Xy+
Xy+
Xy+
XO
XO
XO
Xyp
XO
XO
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyy
Xyyy
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
53
53
53
53
53
53
53
53
57
53
53
53
53
53
53
53
53
4
4
63
66
53
53
53
53
53
53
1
53
73
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
69
53
53
53
58
53
53
53
53
53
53
57
53
57
57
53
53
53
Taxa
Plagiodera atritarsis
Plagiodera erythroptera
Plagiodera rufescens
Plagiodera versicolora
Platycorinus modestus
Platycorinus peregrinus
Platycorinus sp. 1
Platyphora aulica
Platyphora spectablis
Platypria hystrix
Platyprosopus rubidus
Podagrica fuscicornis
Podagrica fuscipes
Podagrica malvae
Podagrica menestriesi
Podontia quatordecimpunctata
Polyconia caroli
Prasocuris junci
Psylliodes affinis
Psylliodes algirica
Psylliodes chrysocephalus
Psylliodes circumdata
Psylliodes cupreus
Psylliodes dulcamarae
-B chromosomes
Psylliodes marcidus
Psylliodes napi
Psylliodes obscuroaeneus
Psylliodes thlaspis
Pyrrhalta viburni
Rhytidocassis indicola
Sagra femorata
Sceloenopla mantecada
Sermylassa halensis
Smaragdina longicornis
Smaragdina mungphuensis
Specularius maindroni
Spermophagus latescenta
Sphaeroderma rubidum
Sphaeroderma testaceum
Spintherophyta sp. 1
Sterromela interlita
Sterromela lineata
Stolas chalybeus
Stolas festiva
Stolas lacordairei
Stolas sp. 1
Strabala ambulans puertoricensis
Strichosa eburata
Syphraea cubana
Syphraea cylindrica
Syphraea quintanillai
Systena basalis
Talurus rugosus
Timarcha affinis
Timarcha aurichalcea
Timarcha balearica
Timarcha calceata
Timarcha catalaunensis
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
30
30
30
28
28
30
28
28
30
26
24
24
16
22
40
40
40
40
36
16
30
30
30
28
28
30
28
28
30
26
24
24
16
22
40
40
40
40
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
53
Xyp
20
30
36
32
34
28
34
34
32
34
36
18
16
18
44
24
24
22
32
32
52
52
12
24
24
24
24
22
30
24
24
24
20
32
16
20
22
30
20
151
16
34
20
30
36
32
34
28
34
34
32
34
36
18
16
18
44
24
24
22
32
32
52
52
12
24
24
24
24
22
30
30
31
33
24
24
24
19
32
16
20
18
22
30
20
NeoXY
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
XY
Xyp
XY
Xyp
XY
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XXNeoXYp
XXY
XXY
Xyp
Xy+
Xy+
XO
NeoXY
XY
Xyp
Xyp
Xyp
Xyp
53
53
53
53
53
53
53
53
53
53
53
57
53
53
53
53
53
53
53
66
53
53
53
53
53
74
74
53
68
53
53
53
53
55
53
53
53
1
1
1
53
53
53
53
53
54
53
45
53
53
53
53
58
53
53
53
53
53
53
75
53
53
53
Taxa
Sexual
System
Timarcha cornuta
Timarcha cyanescens
Timarcha erosa vermiculata
Timarcha espanoli
Timarcha fallax
Timarcha geniculata
Timarcha goettingensis
Timarcha goettingensis normanna
Timarcha gougeleti
Timarcha gr. Perezi
Timarcha granadensis
Timarcha hispanica
Timarcha insparsa
Timarcha intermedia
Timarcha interstitialis
Timarcha intricata
Timarcha laevigata
Timarcha lugens
Timarcha marginicollis
Timarcha maritima
Timarcha metallica
Timarcha monserratensis
Timarcha perezi
Timarcha pimelioides
Timarcha pratensis
Timarcha recticollis
Timarcha rugosa
Timarcha scabripennis
Ploidy
20
20
20
26
20
20
20
20
20
20
20
44
28
20
20
20
20
20
20
28
26
20
26
24
20
20
28
26
22
26
30
30
30
32
28
20
Timarcha sicelidis
Timarcha sinuatocollis
Timarcha strangulata
Timarcha tangeriana
Timarcha tenebricosa
Timarcha ventricosa
Trirhabda canadensis
Trirhabda virgata
Typophorus nigritus
Uroplata girardi
Uroplata nigritarsis
Walterianella eugenia
Walterianella humilis
Walterianella nr. humilis sp. 1
Walterianella nr. Opthalmica sp. 1
Walterianella opthalmica
Walterianella sp. 1
Walterianella sp. 2
Walterianella venusta
Xanthogaleruca luteola
Yingaresca obliterata
Yingaresca variicornis
Zabrotes subfasciatus
Zygogramma bigenera
Zygogramma tetragramma
Ciidae
Cis fuscipes
Cis impressa
parth
Octotemnus aevis
Sulcasis lengi
Cleridae
Enoclerus moestus
Female
2n
22
20
20
22
22
22
22
22
22
22
46
30
40
26
26
24
28
Male
2n
18
20
Xyp
26
20
20
20
20
20
20
22
20
20
20
20
44
28
20
20
20
20
20
20
28
26
20
26
28
24
20
20
28
26
22
26
29
30
30
32
28
20
21
20
20
22
22
Xy+
22
22
22
22
46
30
40
26
26
23
27
Sex Chrom. Source
Karyotype
Xyp
53
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XO
XY
Xyp
Xyp
Xyp
Xyyp
Xyp
Xyp
Xy+
Xy+
53
Xy+
Xy+
Xy+
Xy+
Xy+
XY
NeoXY
Xyp
Xyp
XO
XO
14
75
53
53
53
53
53
53
75
75
75
75
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
75
53
53
53
53
53
53
53
53
53
53
53
53
53
53
53
71
53
53
53
53
53
53
53
76
53
53
77
53
53
14
22
22
22
22
Xyp
Xyp
1
1
1
1
18
18
Xyp
1
152
Taxa
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
Enoclerus nigripes rufiventris
Enoclerus sp. 1
Enoclerus sp. 2
Priocera spinosa
Thanasimus dubius
Thanasimus formicarius
Thanasimus undulatus
Trichodes apiarius
Trichodes nutalli
Trichodes ornatus
Coccinellidae
Adalia bipunctata
18
18
18
18
18
18
18
18
18
18
18
Xyp
18
18
18
18
18
18
18
18
18
1
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Adalia frigida melanopleura
Adonia variegata
Afissa parvula
Aiolocaria mirablis
20
20
20
20
20
20
Xyp
20
18
XY
Xyp
1
17
18
18
18
XY
20
Xyp
20
Xyp
16
16
Xyp
20
18
20
20
20
19
20
20
20
20
20
Xyp
20
Xyp
Xyp
18
18
24
24
22
18
XO
NeoXY
NeoXY
NeoXY
1
Xyp
1
Xyr
1
Xyp
Xyr
1
XY
Xyr
Xyp
Xyr
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
1
Xyp
1
1
NeoXY
NeoXY
NeoXY
NeoXY
1
1
18
16
NeoXY
NeoXY
16
18
18
20
20
18
NeoXY
1
1
1
78
78
1
24
22
22
22
24
22
22
22
NeoXY
NeoXY
NeoXY
NeoXY
1
1
1
1
Anatis 15-punctata
Anatis mali
Anatis rathvoni
Anisocalvia duodecimmaculata
20
Anisocalvia quatuordecimguttata
20
Anisocalvia quatuordecimguttata similis
Aphidecta axyridis
20
Aphidecta axyridis spectablis
Aphidecta bruchii
16
Aphidecta californica
Aphidecta crotchi
Aphidecta novemnotata
Aphidecta obliterata
Aphidecta repanda
Aphidecta septempunctata
Aphidecta septempunctata bruckii
Aphidecta transversalis
Aphidecta transversoguttata
Aphidecta transversoguttata quinquenotata
Aphidecta trifasciata
Aphidecta trifasciata juliana
Aphidecta trifasciata perplexa
Axion pilatei
Axion plagiatum
Azya luteipes
Azya pontbrianti
Azya trinitatis
Brachyacantha 2-3-pustulata
-B chromosomes
Brachyacantha felina
Brachyacantha ursina ursina
Brachyacantha ursina uteella
Brumus parvicollis
Brumus septentrionis
Brumus suturalis
Calvia decemguttata
Calvia quatruordecimguttata
Chilocorus angolensis
-B chromosomes
Chilocorus bilineatus
Chilocorus bipustulatus
Chilocorus cacti cacti
Chilocorus cacti confusor
20
20
20
18
20
18
18
18
18
20
20
20
20
20
16
16
16
20
18
20
20
20
20
20
20
20
20
20
20
20
20
20
18
18
24
22
18
16
20
16
16
20
18
18
153
Xyp
NeoXY
NeoXY
NeoXY
NeoXY
1
1
1
1
1
1
1
1
1
1
1
78
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
45
1
1
1
Taxa
Sexual
System
Ploidy
Chilocorus circumdata
Chilocorus discoideus
Chilocorus distigma
-B chromosomes
Chilocorus erythrocephalus
Chilocorus fraternus
Chilocorus hauseri
Chilocorus hexacyclus
Chilocorus kuwanae
Chilocorus n. sp.
Chilocorus nigritus
Chilocorus orbus
Chilocorus renipustulatus
Chilocorus rubidus
-B chromosomes
Chilocorus schioedtei
Chilocorus similis
Chilocorus stigma
-B chromosomes
Chilocorus tricyclus
Chnoodes sp.
Cladis nitidula
Coccidula rufa
Coccidula scutelata
Coccinella septempunctata
Coccinella transversalis
Coelophora inaequalis
Coleomegilla innotata
Coleomegilla maculata lengi
Cryptognatha nodiceps
Cryptognatha simillima
Cryptolaemus montrouzieri
Curinus coeruleus
Cycloneda polita
Cycloneda sanguinea
Egius platycephalus
Epilachna 12-stigma
20
Epilachna borealis
Epilachna cacica
Epilachna chrysomelina x E. capensis
Epilachna nipponica
Epilachna obscurella
Epilachna orientalis
Epilachna paenulata
Epilachna pustulosa
Epilachna septima
Epilachna sp.
Epilachna varivestis
Epilachna vigintioctomaculata
Epilachna vigintioctomaculata nipponica
Epilachna vigintioctomaculata x pustulosa
Epilachna vigintioctopunctata
18
20
20
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
24
24
22
24
24
NeoXY
NeoXY
NeoXY
1
1
1
24
22
22
14
20
18
22
22
20
18
24
22
22
14
20
18
22
22
20
18
NeoXY
NeoXY
NeoXY
NeoXY
NeoXY
NeoXY
NeoXY
NeoXY
NeoXY
NeoXY
1
1
1
1
1
1
1
1
1
1
24
20
26
24
20
25
NeoXY
NeoXY
XXY
1
1
1
26
20
18
16
20
20
XXY
NeoXY
Xyp
NeoXY
Xyp
Xyp
1
1
1
1
79
79
78
13
1
1
20
18
18
18
20
20
20
18
20
18
20
20
20
20
18
18
25
20
18
16
20
20
20
20
20
20
Xyp
14
14
22
16
20
20
20
20
14
18
10
18
20
18
18
18
20
20
20
18
20
XY
20
20
Xyp
Xyp
18
18
18
20
18
20
Xyp
Xyp
1
83
14
14
18
14
14
18
NeoXY
NeoXY
NeoXY
1
1
1
20
20
20
20
14
14
22
16
20
20
20
20
14
18
10
Xyp
Xyr
Xyp
1
NeoXY
NeoXY
Xyp
NeoXY
Xyp
Xyp
NeoXY
Xyp
Xyr
XY
Xyp
Xyr
Xyp
XY
Xyp
XY
Xyp
Xyp
Xyp
Xyp
1
XY
Xyp
1
1
XY
Xyp
1
1
1
1
1
80
1
1
1
1
45
1
1
1
1
81
1
1
1
1
1
1
1
1
82
-B chromosomes
Eriopis connexa
-B chromosomes
Exochomus bisbinotatus
Exochomus flavipes
Exochomus jamaicensis
154
Taxa
Sexual
System
Ploidy
Exochomus marginipennis californicus
Exochomus marginipennis childreni
Exochomus marginipennis fasciatus
Exochomus metallicus
Exochomus Olituratus sp. 1
-B chromosomes
Exochomus orbiculus
Exochomus Ouropygialis sp. 1
Exochomus Ouropygialis sp. 2
-B chromosomes
Exochomus OuropygialisO sp. 3
Exochomus quadrapustulatus
-B chromosomes
Exochomus quadrapustulatus floralis
Exochomus quadripustulatus
-B chromosomes
Harmonia arcuata
Hippodamia convergens
Hippodamia parenthesis
Hippodamia quinquesignata ambigua
Hippodamia quinquesignata punctulata
Hippodamia sinuata
Hippodamia sp.
Hippodamia tibialis tibialis
Hippodamia tridecimpunctata
Hyperaspis 4-oculata
Hyperaspis annexa
Hyperaspis bigeminata
Hyperaspis billoti
Hyperaspis binotata
Hyperaspis cincta
Hyperaspis congressis
Hyperaspis dissoluta dissoluta
Hyperaspis donzeli
Hyperaspis jucunda
Hyperaspis lateralis lateralis
Hyperaspis lateralis montanica
Hyperaspis notatula
Hyperaspis paspalis
Hyperaspis sp.
Hyperaspis trilineata
Illeis cincta
Illeis indica
Menochilus 6-maculatus
Micraspis cardoni
Microweisea marginata
Microweisea sp.
Mulsantina hudsonica
-B chromosomes
Mulsantina n. sp.
Mulsantina picta minor
Mulsantina picta picta
Neda aequatoriana
Neomysia caseyi
Neomysia oregona
Neomysia pullata pullata
Neomysia pullata randalli
Olla abdominalis
Orcus chalybeus
Pharoscymnus sp.
Phyllobora vigintiduopunctata
Prodilis indagator
16
16
18
20
14
14
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
16
18
NeoXY
NeoXY
18
18
16
18
18
18
18
18
18
16
16
18
18
NeoXY
NeoXY
NeoXY
NeoXY
1
1
18
14
NeoXY
14
1
NeoXY
1
14
14
14
14
14
14
NeoXY
NeoXY
NeoXY
1
1
84
16
20
20
20
20
20
20
20
20
14
16
20
20
20
20
20
20
20
Xyp
14
14
14
16
14
14
14
Xy+
14
16
Xy+
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
Xy+
13
1
1
1
1
1
1
1
14
16
14
14
14
14
14
16
14
14
14
14
14
14
20
20
20
20
20
20
12
12
18
20
20
20
20
20
20
20
20
18
20
20
24
155
14
14
14
14
20
20
20
20
20
20
12
12
18
20
20
20
20
20
20
20
20
18
20
24
NeoXY
NeoXY
Xy+
Xy+
Xy+
Xy+
Xy+
1
Xy+
Xy+
1
1
Xy+
Xy+
Xy+
Xy+
Xyp
Xyp
Xyp
Xyp
XY
XY
NeoXY
NeoXY
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
78
NeoXY
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Taxa
Propylaea japonica
Propylea quatuordecimpunctata
Psyllobora taedata
Rhyzobius chrysomeloides
Rhyzobius litura
Rhyzobius ventralis
Rodolia cardinalis
Scymnus binaevatus
Scymnus marginicollis
Scymnus nubilis
Scymnus sp. 1
Scymnus sp. 2
Scymnus sp. 3
Scymnus sp. 4
Scymnus sp. 5
Solanophila paenulata
Sumnius cardoni
Synonycha grandis
Thea bisoctonotata
Tythaspis sedecimpunctata
Various undet. Spp.
Verania allardi
Verania discolor
Cupedidae
Distocupes varians
Curculionidae
Acalles camelus
Acalles commutatus
Acalles echinatus
Acalles fallax
Acalles petryszaki
Acallopistus sp. 1
Acicnemis alni
Acicnemis nigra
Adexius scrobipennis
Adosomus melanogrammus
Alcides collaris
Alcides sp. 1
Alcidodes affaber
Alcidodes sp. 1
Amblyrrhinus poricollis
Amblyrrhinus sp. 1
Amblyrrhinus subrecticollis
Amystax fasciatus
Anobleptus sp. 1
Anthonomus bisignifer
Anthonomus corvulus
Anthonomus grandis
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
20
18
18
16
18
18
16
16
16
16
18
20
20
16
18
18
20
20
20
XY
78
NeoXY
Xyp
NeoXY
XO
XO
NeoXY
NeoXY
NeoXY
NeoXY
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyr
Xyp
20
20
20
20
18
18
16
17
17
16
16
16
16
18
20
20
16
18
18
20
20
24
20
20
20
20
20
30
28
30
30
30
30
24
24
22
24
36
46
28
40
34
32
32
22
22
22
16
22
1
XY
XY
Xyp
Xyp
1
79
79
1
1
1
1
1
1
1
1
1
1
45
1
1
1
78
1
1
1
1
19
XO
85
22
24
20
30
32
34
36
26
44
42
30
28
30
30
30
30
24
24
22
24
36
46
28
40
34
32
32
22
22
22
16
22
23
22
24
19
30
32
34
36
26
44
42
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyyp
Xyp
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
6
6
6
86
86
86
5
1
87
1
87
1
13
1
87
87
87
87
87
87
87
87
87
87
1
1
1
1
1
1
1
88
87
32
28
34
43
32
28
34
XXY
XY
Xyp
Xyp
87
1
1
1
-B chromosomes
Anthonomus haematopus
Anthonomus scutellatus
Anthonomus signatus
156
Taxa
Sexual
System
Ploidy
Anthonomus sp. 1
Anthonomus sp. 2
Anthonomus sp. 3
Apiophorus pictus
Aplotes roelofsi
Apodrusus wolcotti
Apotomorhinus cribatus
Arrhines languides
Astycus lateralis
Astycus sp.
-B chromosomes
Atactogaster sp. 1
Athesapeuta sp. 1
Athesapeuta sp. 2
Barioscaphus cordiae
Baris arthemisiae
Baris ezoana
Barynotus obscurus
Barynotus squamosus
Barypeithes albinae
Barypeithes chevrolati
Barypeithes formaneki
Barypeithes interpositus
Barypeithes liptoviensis
Bothrometopus fasciatus
Brachyderes incanus
Brachysomus dispar
Brachysomus echinatus
Brachysomus hirtus
Brachysomus setiger
Byctiscus congener regalis
Byctiscus venustus
Byrsopages kiso
Byrsopages sp(Mt. Asahidake)
Byrsopages sp(Mt. Niseko)
Byrsopages sp(Mt. Proshiri)
Caenocryptorrhynchus frontalis
Calendra australis australis
Calendra bruchi
Calendra costipennis
Calendra linearis
Calendra parvula
Sex Chrom. Source
Karyotype
34
26
32
30
22
34
26
32
30
22
20
26
22
22
22
XY
Xyp
Xyp
Xyp
Xyp
48
32
32
22
38
32
34
32
18
22
32
54
Xyp
48
22
38
32
34
32
18
22
32
54
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
38
parth
parth
parth
parth
3
5
4
3
Barypeithes mollicomis
Barypeithes pellucidus
-B chromosomes
Barypeithes purkynei
Blosyrus asellus
Blosyrus falcatus
Blosyrus japonicus
Male
2n
26
22
22
22
Baris sp.
Baris striata
Barynotus moerens
Female
2n
parth
parth
parth
parth
parth
parth
3
4
5
6
3
3
33
55
44
33
22
22
22
22
26
28
22
22
22
22
24
22
33
44
55
66
22
22
22
22
33
33
22
22
18
22
22
22
22
34
26
26
30
24
26
157
22
22
22
22
26
28
22
22
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
22
22
24
22
Xyp
Xyp
Xyp
Xyp
22
22
22
22
Xyp
Xyp
Xyp
22
22
18
22
22
22
22
34
26
26
30
24
26
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
1
5
87
1
87
5
5
5
5
87
89
89
1
87
87
1
1
1
1
1
1
1
1
90
90
90
90
90
87
90
90
1
90
87
1
91
91
91
91
91
87
90
90
90
90
90
1
1
1
1
1
1
87
1
45
1
1
1
Taxa
Calendra zeae
Callirhopalus bifasciatus
Callirhopalus minimus
Callirhopalus obesus
Callirhopalus setosus
-Female 2n = 41-44
Calosirus albosuturalis
Canonopsis sericea
Carcilia strigicollis
Catagmatus japonicus
Catapionus gracilicornis
Sexual
System
Ploidy
parth
parth
parth
parth
parth
parth
parth
parth
3
4
5
3
4
parth
parth
parth
parth
parth
parth
Catapionus modestus
Catapionus obscurus
Catapionus viridimetallicus
Cathormiocerus aristatus
parth
Centricnemus leucogrammus
Ceutorrhynchus ancola
Ceutorrhynchus costatus
Ceutorrhynchus diffusus
Ceutorrhynchus erysimi
Ceutorrhynchus lewisi
Ceutorrhynchus punctiger
Ceutorrhynchus sp. 1
Chlorophanus grandis
Chlorophanus viridis
Chokkirius truncatus
Christensenia antarctica
Cidnorrhinus 4-maculatus
Cionus championi
Cionus ganglbaueri
Cionus hortulans
Cionus longicollis-montanus
Cionus nigritarsis
Cionus olivieri
Cionus tamazo
Cionus tuberculosus
Cirrorhynchus kelecsenyi
Cleonus japonicus
Colobodes konoi
Colobodes matasumurau
Colobodes matsumurai
Colobodes sp. 1 Ov-alum n.sp.O
Colobodes sp. 2
Corigetus chandigarhensis
Corigetus sp. 1
Cosmopolites sordidus
Cryptorhynchus sp. 1
-Male 2n = 22-53
Cryptorrhynchus erectus
Cryptorrhynchus lapathi
-Male 2n = 33-34
Cryptorrhynchus sp. 1
Curculio aino
4
2
3
4
5
6
3
48
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
33
44
55
33
44
44
65
43
26
Xyp
1
87
87
87
1
1
1
1
1
34
22
22
32
22
33
44
55
66
44
22
22
20
33
22
34
20
24
28
28
24
24
24
22
22
22
28
40
38
38
40
38
42
40
44
30
28
48
48
34
22
22
32
XY
Xyp
Xyp
Xyp
22
22
20
Xyp
Xyp
NeoXY
22
34
20
24
28
28
24
24
24
22
22
22
28
40
38
38
40
38
42
40
44
30
28
48
48
48
36
22
22
30
Xyp
36
22
22
30
38
Xyp
Xyp
Xyp
XY
XY
Xyc
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
XY
Xyp
1
Xyp
Xyp
Xyp
Xyp
1
87
1
1
87
87
87
87
87
92
87
93
1
94
6
1
1
1
1
1
1
1
1
87
1
87
1
5
87
87
87
87
87
1
87
95
1
1
1
87
32
34
32
34
Xyp
Xyp
1
1
34
36
32
38
28
34
36
32
38
28
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
158
1
87
1
96
87
Taxa
Sexual
System
Ploidy
Curculio arakawai
Curculio convexus
Curculio dentipes
Curculio distinguendus
Curculio elaeagni
Curculio ficusi
Curculio flavescens
Male
2n
Sex Chrom. Source
Karyotype
26
26
26
26
26
26
26
26
26
26
26
26
28
26
32
26
34
26
26
26
22
26
32
26
24
26
28
26
32
28
32
22
22
22
Xyp
Xyp
Xyp
Xyp
Xyp
22
22
22
22
22
24
20
20
22
22
20
22
22
22
22
22
22
22
28
34
26
30
22
22
22
22
22
22
22
22
22
28
24
22
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
26
28
26
32
26
34
26
26
26
22
26
32
26
24
26
28
26
Curculio funebris
Curculio koreanus
Curculio longirostris
Curculio minutissimus
Curculio murakamii
Curculio nucum
Curculio obtusus
Curculio pictus
Curculio quercivorus
Curculio roelofsi
Curculio sp. 1
Curculio sp. Nov.
Cycnotrachelus roelofsi
Cylindrocopturus sp. 1
Cyphicerinus tectonae
Cyphicerus aceri
Cyrtepistomus bardus
Cyrtepistomus castaneus
-Female 2n = 30-32
Cyrtepistomus gucindus
Cyrtepistomus jucundus
Cyrtepistomus sp. 1
Deiradolcus pubescens
Deiradolcus sp. 1
Demimaea mori
Dermatoxenus caesicollis
Female
2n
parth
Dermatoxenus clathratus
Dermatoxenus sp. 1
Desmidophorus hebes
Desmidophorus sp. 1
Diaprepes abbreviatus
Diaprepes marginicollis
Diaprepres abbreviata
Diatropus marshalli
Dicranthus elegans
Dicranthus majzlani
Didothis melancholica
Didothis sp. 1
Diocalandra sp. 1
Diplogrammus quadrivittatus
Disker tenuicornis
Dodecastichus atripes
Dodecastichus aurosignatus vlasuljensis
Dodecastichus dolomitae dryadis
Dodecastichus geniculatus
Dodecastichus inflatus
Dodecastichus obsoletus as D. Speiseri
Donus comatus carpathicus
Donus oxalidis
Dorytomus mongolicus
Dorytomus shikotanus
Dusmoecetes marioni
3
28
32
22
22
22
31
22
22
22
22
22
24
20
22
22
20
22
22
22
22
22
22
22
28
34
26
30
22
22
22
22
22
22
22
22
22
28
24
22
159
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
5
1
1
1
1
5
1
1
87
1
1
1
1
87
87
87
87
1
5
1
1
5
1
5
1
87
87
87
5
5
1
1
1
1
87
1
5
96
96
1
87
87
87
87
87
1
45
87
95
95
95
95
95
95
1
1
1
1
87
Taxa
Dusmoecetes richtersi
Dyslobus verrucifer
Dystropicus dorsalis
Echinocnemus marmoreus
Echinocnemus sp. 1
Echinocnemus squameus
Ectatorhinus adamsi
Ectemnorrhinus angusticollis
Ectemnorrhinus drygalski
Ectemnorrhinus viridis
Episomus lacerta
Episomus turritus
Esamus albomarginatus
Esamus circumdatus
Esamus plurisetosus
Esamus sciurus
Eudiagogus episcopalis
Eugnathus distinctus
Eugnathus distincyus
Euops punctatostriata
Euops splendida
Eupagoderes sp. 1
Eusomus ovulum
-Female 2n = 28-32
Sexual
System
parth
parth
Euthyrhinus yakushimanus
Exophthalmus famelicus
Exopthalmus quadritaenia
Foucartia liturata
Foucartia squamulata
-Female 2n = 31-33
Gasteroclisus aethiops
Gasteroclisus binodulus
Gelus calafornicus
-B chromosomes
Gymnetron antirrhini
Gymnetron smreczynski
Gymnetron tetrum
Heilipodus erythropus
Heilipodus scaber
-XpneoXneoYyp
Heilipodus scabripennis
-neoXpneoXneoYyp
Heilipodus sp.
Heilipus tremolerasi
-neoXpneoYyp
Heilipus wiedemanni
Hemerus sp. 1
Hilaus sp. 1
Hydrophilus olivaceus
Hylobius abietis
Hylobius abietis haroldi
Hylobius albosparsus
Hylobius aliradicis
Hylobius congener
Hylobius cribripennis
Hylobius desbrochersi
Hylobius elongatus
parth
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
22
56
18
16
22
38
22
22
22
26
32
22
22
22
22
34
22
22
26
22
22
30
22
22
56
18
16
22
38
22
22
22
26
32
22
22
22
22
34
22
22
26
22
22
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
XY
Xyp
33
16
22
22
22
22
32
16
22
22
22
22
NeoXY
Xyp
Xyp
XY
Xyp
90
87
96
96
90
1
98
36
36
38
25
36
36
38
26
Xyp
Xyp
Xyp
XO
87
87
87
99
32
32
32
30
32
32
32
30
30
Xyp
Xyp
Xyp
Xyp
XXYyp
1
87
87
87
87
30
XXYyp
87
28
30
28
30
29
Xyp
Xyp
Xyyp
100
87
87
22
30
22
18
20
22
24
30
22
22
40
22
22
36
36
48
22
30
22
18
20
22
24
29
22
22
40
22
22
36
36
48
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
87
45
87
87
87
87
87
13
1
1
1
1
1
1
1
1
160
87
1
5
1
87
87
1
87
87
87
13
1
5
5
5
5
45
1
87
1
1
1
97
Taxa
Sexual
System
Hylobius exculptus
Hylobius freyi (=exculptus)
Hylobius galloisi
Hylobius gebleri
Hylobius gigas
Hylobius insularis
Hylobius montanus
Hylobius pales
Hylobius perforatus
Hylobius piceus
Hylobius pinastri
Hylobius pinastri karafutoensis
Hylobius pinicola
Hylobius radicis
Hylobius sp. 1
Hylobius warreni
Hypera elongata
Hypera medicaginis
Hypera mongolicus
Hypera nigrirostris
Hypera postica
Hypera punctata
Ploidy
22
Hypera rumicus
Hypera viciae
Hypermias sp. 1
Hypolixus truncatulus
Hypomeces squamosus
Indocurculio minutus
Indomecus lectus
Indomias acutipennis
Ixalma dentipes
Larinodontes obtusus
Larinus griseopilosus
Larinus latissimus
Larinus meleagris
Larinus saussureae
Lepidospyris demissus
Lepropus chrysochlorus
Lepropus flavovittatus
Lepropus lateralis
Leptomias angustatus
Leptomias waltersi
Lepyrus alternans
Lepyrus japonicus
Lepyrus konoi
Lepyrus palustris
Limnobaris jucunda
Liophloeus gibbus
Liophloeus lentus
Liophloeus tessulatus
-Female 2n = 29-31
parth
3
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
28
36
22
36
50
20
22
36
40
22
22
40
22
24
32
22
22
22
22
22
28
28
36
22
36
50
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
22
36
40
22
Xyp
40
22
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
32
22
22
22
22
22
20
22
22
22
22
44
22
24
22
22
22
40
40
40
40
36
22
20
22
22
22
22
44
22
24
22
22
22
40
40
40
40
36
22
23
22
22
22
22
22
22
30
30
30
30
24
30
22
22
Xyp
Xyp
Xyp
XY
Xyp
Xyp
XY
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
22
22
22
22
22
22
30
30
30
30
24
30
22
22
30
22
Liparus glabrirostris
Listroderes costirostris
-Female 2n = 25-30
22
32
Xyp
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
87
87
1
1
87
1
87
1
1
87
87
1
5
5
5
87
1
87
1
1
1
5
87
87
5
87
87
87
5
5
1
1
1
1
1
1
87
87
1
parth
3
28
1
87
1
parth
3
35
1
parth
3
30
1
parth
3
30
1
-Female 2n = 28-41
-Female 2n = 29-30
161
Taxa
Lixus acutipennis
Lixus cardui
Lixus divaricatus
Lixus impressiventris
Lixus maculatus
Lixus subtilis
Lobotrachelus sp. 1
Macrocorynus griseoides
Magdalis alutacea
Magdalis gentilis
Magdalis leconti
Sexual
System
Ploidy
Female
2n
parth
3
33
46
44
46
40
46
40
28
33
32
30
parth
3
Magdalis perforata
Magdalis piceae
Magdalis sp. 1
Mechistocerus fumosus
Mechistocerus nipponicus
Mechistocerus sp. 1
Mechistocerus sp. 2
Mechistocerus sp. 3
Mechistocerus sp. Nov.
Mecopomorphus griseus
Mecyslobus flavosignatus
Mecyslobus westermanni
Mecysmoderes sp. 1
Mecysolobus erro
Mecysolobus flavosignatus
Mecysolobus trifidus
Meotiorrhynchus querendus
Metamasius hemipterus
Metapocyrtus yonagunianus
Metialma scenica
Metialma sp
Miccotrogus picirostris
Myllocerus 11-pustulatus maculosus
Myllocerus angulatipes
Myllocerus blandus
Myllocerus conspersus
Myllocerus dentifer
Myllocerus discolor
Myllocerus discolor canescans
Myllocerus dorsatus
Myllocerus fumosus
parth
Myllocerus griseus
Myllocerus kashmiriensis
Myllocerus leativirens
Myllocerus lefroi
Myllocerus nipponicus
parth
Myllocerus paetus
Myllocerus pauper
Myllocerus sabulosus
Myllocerus sp. 1
Myllocerus sp. 2
Myllocerus subfasciatus
22
3
3
30
32
32
30
34
36
28
22
32
40
20
32
40
22
32
40
24
34
32
38
34
34
30
22
26
22
42
32
32
34
30
34
22
22
22
22
22
22
22
22
33
22
22
22
22
33
22
22
22
22
30
22
22
162
Male
2n
Sex Chrom. Source
Karyotype
46
44
46
40
46
40
28
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
32
30
16
30
32
32
30
34
36
28
22
32
40
20
32
40
22
32
40
24
34
32
38
34
34
30
22
26
22
42
32
32
34
30
34
22
22
22
22
22
22
Xyp
22
Xyp
Xyp
22
22
22
22
Xyp
Xyp
Xyp
Xyp
22
22
22
22
30
22
22
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
5
Xyp
1
1
87
1
1
1
1
87
87
1
1
1
1
1
1
1
1
87
1
87
87
87
1
1
1
87
87
87
87
87
5
1
1
1
1
1
96
87
87
87
87
87
1
1
1
87
87
87
87
87
87
87
1
87
87
87
1
87
87
87
1
1
87
1
Taxa
Myllocerus transmarinus
Myllocerus tusicollis
Myllocerus viridanus
Myllocerus viridens
Myosides seriehispidus
-Female 2n = 30-33
Naupactus bruchi
Sexual
System
parth
Ploidy
3
Naupactus peregrinus
Naupactus xantographus
Nedyus quadrimaculatus
Neocanonopsis dreuxi
Neomecopus sp. 1
Niphades sp. 1
Niphades variegatus
Notaris acridulus
Odoiporous longicollis
Oedophyrus hilleri
Okikuruminus oblongus
Okikuruminus roelofsi
Orobitis apicalis
Orochlesis takaosanus
Otiorhynchus alpicola
Otiorhynchus alpicola atterimus
Otiorhynchus anthracinus
-Female 2n = 54-55
Otiorhynchus apenninus
Otiorhynchus arcticus
parth
parth
parth
4
6
22
5
Otiorhynchus armadillo
Otiorhynchus atripes
Otiorhynchus atroapterus
Otiorhynchus aurosignatus vlasuljensis
Otiorhynchus austriacus
Otiorhynchus dryadis
Otiorhynchus dubius
Otiorhynchus equestris
Otiorhynchus fuscipes
Otiorhynchus gemmatus
Otiorhynchus geniculatus
Otiorhynchus inflatus
Otiorhynchus inflatus salebrosus
Otiorhynchus kollari
Otiorhynchus koritnicensis
Sex Chrom. Source
Karyotype
22
22
22
22
32
22
22
22
22
Xyp
Xyp
Xyp
Xyp
87
87
87
1
1
22
22
22
22
Xyp
Xyp
22
22
28
22
30
32
34
30
52
78
26
26
36
36
26
40
22
22
55
22
22
28
22
30
32
34
30
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
26
36
36
26
40
22
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
87
87
45
101
87
45
87
87
1
87
1
1
102
102
102
1
1
103
1
87
95
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
22
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XXOO
Xyp
Xyp
XY
20
XXOO
22
22
22
22
28
Xyp
22
22
22
95
Xyp
XY
Xyp
22
22
44
22
22
22
33
22
22
22
22
22
22
22
22
Xyp
Xyp
22
22
22
Xyp
XY
XY
22
22
22
22
22
22
Xyp
Xyp
Xyp
XY
XY
Xyp
parth
3
parth
4
44
Otiorhynchus chrysops
-XX00
Otiorhynchus coecus as O. niger
Otiorhynchus cornicinus
Otiorhynchus corvus
Otiorhynchus croaticus
-Female 2n = 22-33
Male
2n
22
22
22
33
Otiorhynchus bisulcatus
Otiorhynchus carmagnolae
Otiorhynchus chrysocomus
-Female 2n = 32-33
Female
2n
22
parth
parth
parth
4
3
163
1
95
1
95
95
87
104
87
95
1
104
95
95
1
1
1
95
95
87
87
95
87
1
95
1
1
1
95
1
1
1
95
95
Taxa
Otiorhynchus ligustici
-Female 2n = 33-35
Otiorhynchus mendax
Otiorhynchus meridionalis
Sexual
System
Ploidy
parth
34
parth
44
22
22
22
22
22
22
22
22
22
Otiorhynchus minutesquamosus
Otiorhynchus morio
Otiorhynchus multipunctatus
-B chromosomes
Otiorhynchus niger
Otiorhynchus obsidianus
Otiorhynchus obtusus
Otiorhynchus opulentus
Otiorhynchus ovatus
-Female 2n = 30-34
Otiorhynchus pauxillus
-Female 2n = 29-38
parth
parth
3
Otiorhynchus salicicola psudonotus
Otiorhynchus salicis
parth
-Female 2n = 28-35
parth
-Female 2n = 29-34
parth
-Female 2n = 32-33
parth
33
22
22
22
22
22
32
Male
2n
Sex Chrom. Source
Karyotype
105
22
22
XY
22
22
22
22
22
XY
Xyp
95
Xyp
XY
Xyp
XY
Xyp
22
22
22
22
22
XY
Xyp
XY
XY
Xyp
33
34
parth
parth
Otiorhynchus praecellens bosnarum
Otiorhynchus proximus
parth
Otiorhynchus raucus
parth
Otiorhynchus repletus
Otiorhynchus retifer
Otiorhynchus rotifer
Otiorhynchus rugifrons
parth
Otiorhynchus scaber
-Female 2n = 30-34
Female
2n
87
1
95
87
1
95
1
95
1
1
1
95
95
87
1
1
1
3
33
22
33
33
22
22
22
33
22
22
32
3
32
1
3
33
1
3
22
Xyp
22
22
22
XY
Xyp
Xyp
22
22
Xyp
XY
1
95
1
106
95
87
95
1
1
1
1
parth
3
32
22
32
parth
3
33
1
parth
4
43
1
parth
4
44
1
parth
3
22
Xyp
1
1
1
-Female 2n = 32-33
-Female 2n = 40-45
-Female 2n = 43-44
Otiorhynchus sensitivus
Otiorhynchus singularis
Otiorhynchus speiseri
Otiorhynchus strumosus
Otiorhynchus subdentatus
-Female 2n = 43-44
Otiorhynchus sulcatus
-Female 2n = 32-33
Otiorhynchus tenebricosus
Otiorhynchus vehemens
Oxydema sp. 1
Pachylobius picivorus
parth
4
33
22
33
22
22
44
parth
parth
3
3
33
33
1
1
parth
parth
3
3
33
34
22
22
30
1
1
95
1
87
1
parth
3
164
22
Xyp
22
22
Xyp
Xyp
22
22
30
36
XY
XY
Xyp
1
95
1
87
95
1
Taxa
Pachyrhynchus infernalis
Pachytychius sp. 1
Pagiophloeus sp. 1
Paophilus afflatus
Parafoucartia squamulata
Paramecopus farinosus
Paraplapoderus ulmi
Parascaphus sp. 1
Paratrachelophorus longicornis
Peritelus hitricornis
Sexual
System
parth
Ploidy
3
30
parth
parth
Phloeophagosoma takenouchii
Phyllobius annectans
Phyllobius arborator
Phyllobius argentatus
Phyllobius armatus
Phyllobius brevis
Phyllobius brevitarsis
Phyllobius caucasicus
Phyllobius galloisi
Phyllobius incomptus
Phyllobius intrusus
Phyllobius longicornis
Phyllobius maculicornis
Phyllobius mundus
Phyllobius nigritus
Phyllobius oblongus
Phyllobius prolongatus
Phyllobius pyri
Phyllobius rotundicollis
Phyllobius sp. 1
Phyllobius urticae
Phyllobius viridicollis
Phymatapoderus latipennis
Phymatapoderus pavens
Phytobius sp. 1
Phytoscaphus inductus
-B chromosomes
Phytoscaphus sp. 1
Pissodes affinis
Pissodes affinis curriei
Pissodes approximatus
-Male 2n = 30-34
Pissodes approximatus canadensis
-Male 2n = 30-34
Pissodes dubius
Pissodes dubius fraseri
Pissodes dubius piperi
Pissodes fasciatus
Pissodes fiskei
-XYp III+10 haploid
Pissodes gyllenhali
Pissodes memorensis
Pissodes nitidus
Pissodes notatus
Pissodes obscurus
Pissodes OP27O
Pissodes OYT3O
Pissodes radiatae
Pissodes rotundatus
Pissodes rotundatus alascensis
Pissodes rotundatus nigrae
Pissodes schwarzi
32
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
32
28
22
33
32
30
22
20
30
33
44
32
22
22
22
22
22
22
22
22
22
22
22
24
22
22
22
22
22
22
22
22
22
22
32
24
24
22
22
32
28
22
Xyp
Xyp
Xyp
Xyp
32
30
22
20
Xyp
Xyp
XY
Xyp
Xyp
1
Xyp
22
22
22
22
22
22
22
22
22
22
22
24
22
22
22
22
22
22
22
22
22
22
32
24
24
22
1
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
20
30
30
20
30
30
32
Xyp
Xyp
Xyp
1
1
32
30
87
87
87
90
90
87
1
5
87
1
87
87
1
1
87
107
1
1
87
87
1
87
1
1
1
1
87
1
1
87
1
1
1
1
87
87
1
1
1
1
30
30
30
30
24
30
30
30
30
25
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
32
30
30
30
28
30
34
30
30
30
30
28
32
30
30
30
28
30
34
30
30
Xyp
30
28
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
1
1
1
1
1
1
1
1
1
165
1
1
Taxa
Pissodes schwarzi yosemite
Pissodes similis
Pissodes similis utahensis
Pissodes strobi
Pissodes strobi engelmanni
Pissodes strobi sitchensis
Pissodes terminalis
-XYp III+12 haploid
Pissodes validirostris
Pissodes webbi
Platymycterus himalayanus
Platymycterus moestus
Platymycterus sjostedti
Platymycterus sp.
Plintini sp. 1
Podapion gallicola
Podeschrus abietis
Polydrusus amoenus
Polydrusus atomarius
Polydrusus calabricus
Polydrusus impressifrons
Polydrusus inustus
Polydrusus marginatus
Polydrusus mollis
-Female 2n = 30-31
Sexual
System
Ploidy
Male
2n
Sex Chrom. Source
Karyotype
28
30
30
34
34
34
28
28
30
30
34
34
34
29
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
108
1
1
1
1
30
34
22
22
22
22
22
22
22
30
34
22
22
22
22
22
22
22
22
22
20
20
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
22
Xyp
1
1
87
87
5
87
1
1
1
1
87
87
1
106
87
1
22
20
parth
parth
3
33
22
31
parth
3
33
parth
2
22
22
22
22
22
22
22
22
24
Polydrusus pilosus
Polydrusus ruficornis
Polydrusus sericeus
Polydrusus sicanus
Polydrusus undatus
Polydrusus viridicintus
Procas biguttatus
Proctorus decipiens
Protacallinus sp. 1
Pseudoanthonomus hamamelidis
32
Pseudphytoscaphus sp. 1
Ptochus oyulum
Pycnodactylus hypocritia
Rhadinomerus annulipes
30
32
38
22
22
34
36
44
20
22
20
28
24
28
Rhadinomerus maebarai
Rhinocerus sp. 1
Rhinomias forticornis
Rhinoncus pagnus
Rhinoncus sibiricus
Rhynchaenus mutabilis
Rhynchaenus niger
Rhynchaenus pallicornis
Rhynchaenus populi
Rhynchaenus sp. 1
Rhynchaenus stigma
Rhynchophorus ferrugineus
Rhyssomatus marginatus
Ruteria hypocrita
Rynchophorus palmarum
Ryssematus lineaticollis
Scepticus griseus
Scepticus insularis
Female
2n
parth
2
28
26
26
28
22
22
22
30
22
24
22
22
166
Xyp
Xyp
22
22
22
22
XY
Xyp
Xyp
Xyp
22
22
24
22
30
XY
38
22
22
38
34
36
44
20
22
20
28
24
28
26
28
26
25
28
22
22
22
30
22
24
22
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
XY
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
106
109
1
1
87
87
87
1
87
1
1
1
1
1
5
5
5
87
87
1
87
87
5
1
1
1
1
1
1
1
87
1
87
45
6
96
1
1
1
Taxa
Sexual
System
Ploidy
Female
2n
parth
parth
3
5
33
55
22
22
Scepticus tigrinus
Sciaphilus asperatus
-bisexual
Sciopithes obscurus
Scythropus japonicus
Scythropus ornatus
Scythropus sp. 1
Shirahoshizo insidiosus
Simo variegates
Simulatacalles simulator
Sipalus hypocrita
Sitona crinitus
Sitona cylindricollis
Sitona flavescens
Sitona hispidulus
Sitona humeralis
Sitona lepidus
Sitona lineata
Sitona lineatus
Sitona macularis
Sitona scissifrons
Sitona sp. 1
parth
parth
parth
Sitona suturalis
Sitophilus granarius
Sitophilus oryzae
Sitophilus sasakii
Sitophilus zeamais
-B chromosomes
Sphenophorus parvula
Strophosoma capitatum
Strophosoma faber
Strophosoma melanogrammum
Strophosoma melanogrammus
-Female 2n = 31-35
3
4
3
33
44
22
22
20
38
33
24
22
22
22
22
22
22
22
22
22
22
22
22
24
26
22
12
24
12
22
22
22
22
22
parth
parth
30
22
22
33
33
parth
34
parth
34
22
22
22
22
Male
2n
22
23
22
Sex Chrom. Source
Karyotype
Xyp
Xyyp
Xyp
22
22
20
38
Xyp
Xyp
Xyp
Xyp
24
22
22
22
22
22
22
22
22
22
22
22
22
24
26
22
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
24
11
22
22
22
22
22
Xyp
XO
XY
NeoXY
XY
Xyp
NeoXY
30
22
22
Xyp
XY
Xyp
1
1
1
1
1
98
90
1
1
1
1
1
94
87
1
5
1
1
1
6
1
1
6
6
1
1
1
1
6
1
110
111
1
1
1
1
1
1
90
90
90
1
1
-Female 2n = 33-34
Sympiezomias cribicollis
Sympiezomias cribricollis
Sympiezomias lewisii
Sympiezomias sp. 1
Syrotelus septentrionalis
Syrotelus umbrosus
Tachypterellus 4-gibbus
Tanymecus cephalotes
Tanymecus feae
Tanymecus hispidus
Tanymecus indicus
Tanymecus longulus
Tanymecus palliatus
Tanymecus sciurus
Tanymecus sp. 1
Tanysphyrus lemnae
16
34
22
22
22
22
22
22
22
16
167
22
22
22
22
16
16
34
22
22
22
22
22
22
22
23
22
16
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyyp
Xyp
Xyp
1
1
87
1
1
87
87
1
87
87
5
87
87
87
87
87
1
1
Taxa
Sexual
System
Ploidy
Tanysphyrus major
Telephae konoi
Thlipsomerus glebosus
Trachyphloeus aristatus
Trachyphloeus bifoveolatus
-Female 2n = 32-33
Trachyphloeus parallelus
Trachyrinus sp. 1
Transptochus sp.
Trichalophus albonotatus
Tropiphorus carinatus
-Female 2n = 32-33
Tropiphorus cucullatus
-Female 2n = 40-41
Tropiphorus elevatus
Tropiphorus terricola
Tychius aureolus femoralis
Tychius ginsuji
Tychius sp. 1
Tychius stephensi
Tylotus chrysops as O. chrysops
Xanium vanhoeffenianum
Xanthochelus sp.
Zacladus geranii
Dermestidae
Anthrenus fasciatus
Anthrenus flavipes
Anthrenus sp.
Anthrenus verbasci
Attagenus elongatulus
Attagenus megatoma
Dermestes ater
Dermestes caninus
Dermestes frischii
parth
parth
3
3
parth
parth
parth
3
4
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
16
22
26
22
33
33
18
16
22
26
22
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
87
1
1
parth
3
33
44
33
22
28
33
parth
4
41
1
parth
parth
3
4
33
44
40
34
34
40
22
22
22
28
94
1
1
1
1
1
Dermestes haemorrhoidalis
Dermestes lardarius
Dermestes maculatus
-1-4 y chromosomes
Dermestes peruvianus
Dermestes talpinus
Dermestes vulpinus
Dermests signatus
Orphilus subnitidus
Trogoderma glabrum
Trogoderma inclusum
Trogoderma parabile
Trogoderma variabile
Dytiscidae
Agabus biguttatus
Agabus bipustulatus
22
28
22
22
28
Xyp
Xyp
Xyp
Xyp
95
Xyp
Xyp
Xyp
18
18
18
18
18
18
18
18
18
18
18
18
19
18
18
18
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyyp
Xyp
Xyp
Xyp
1
1
1
1
1
1
1
1
1
1
1
1
1
18
18
18
18
18
20
18
20
18
18
20
18
18
18
18
18
18
20
18
20
18
18
20
18
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
1
1
1
1
1
1
1
44
44
42
43
43
42
XO
XO
Xyp
112
113
18
43
42
40
XO
XY
XY
113
18
18
43
43
XO
XO
113
113
18
18
18
18
18
18
18
18
18
-Xyp Subsequent studies failed to replicate
Agabus confinis
44
42
Agabus conspersus
40
-material studied was from India and likely is a different species
44
Agabus ramblae
44
168
40
34
34
40
Xyp
NeoXY
1
94
1
5
1
1
87
87
87
Taxa
Sexual
System
Ploidy
Agabus sturmi
-Xyp Subsequent studies failed to replicate
Boreonectes alpestris
Boreonectes griseostriatus
Boreonectes ibericus
Boreonectes macedonicus
Boreonectes multilineatus
Boreonectes riberae
Colymbetes adpressus
Colymbetes affinis
Colymbetes biguttatus
-reported as 22 in text likely a misprint
Colymbetes binotatus
Colymbetes bipustulatus
Colymbetes bipustulatus var. solieri
Colymbetes confinis
-only females examined
Colymbetes congener
Colymbetes conspersus
Colymbetes fuscus
Colymbetes fuscus
-Male 2n = 35-37
Colymbetes infuscatus
Colymbetes labiatus
Colymbetes lapponicus
Colymbetes melanarius
Colymbetes nebulosus
Colymbetes nevadensis
Colymbetes paykulli
Colymbetes paykulli
-Xyp Subsequent studies failed to replicate
Colymbetes piceus
Colymbetes ramblae
Colymbetes serricornis
Colymbetes striatus
Colymbetes striatus
Colymbetes sturmii
Colymbetes thomsoni
Colymbetes unguicularis
Colymbetes wollastoni
Cybister japonicus
Cybister lateromarginalis
Cybister limbatus
Cybister sugillatus
Cybister tripunctatus
Deronectes angusi
Deronectes costipennis costipennis
Deronectes costipennis gignouxi
Deronectes ferrugineus
Deronectes latus
Deronectes platynotus
Deronectes wewalai
Dytiscus circumcinctus
Dytiscus dauricus
Dytiscus marginalis
-Male 2n = 36-41
28
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
44
42
43
42
XO
Xyp
113
18
56
62
54
54
58
56
44
44
44
55
61
53
53
57
55
44
43
43
XO
XO
XO
XO
XO
XO
XY
XO
XO
114
114
114
114
114
114
115
115
115
44
44
44
44
43
43
43
XO
XO
XO
115
115
115
115
44
44
42
43
43
41
36
XO
XO
XO
115
115
115
18
44
44
44
44
44
44
42
38
36
44
43
43
43
43
43
41
37
36
NeoXY
XO
XO
XO
XO
XO
XO
XO
Xyp
115
115
115
115
115
115
115
18
18
42
44
44
42
40
42
44
44
44
44
44
41
43
43
41
40
42
44
43
43
43
43
43
22
43
43
43
50
28
NeoXY
62
50
28
62
38
37
39
XO
XO
XO
XO
XY
XY
XY
XO
XO
XO
XO
112
115
115
115
18
18
18
115
115
115
115
116
18
18
18
18
117
117
38
37
39
XY
44
44
44
50
28
28
62
50
28
62
38
40
38
Dytiscus marginalis czerskii
Dytiscus sp.
-Male 2n = 38-40
169
XO
XO
XO
XY
NeoXY
117
XY
XY
NeoXY
XY
XY
117
117
117
117
18
116
18
18
18
116
18
Taxa
Sexual
System
Ploidy
Eretes sp.
Eretes strictus
Hydaticus decorus
Hydaticus fabricii
Hydaticus leander
Hydaticus luczonicus
Hydaticus vittatus
Hydroporus humilis
Hydrovatus cuspidatus
Ilybius aenescens
Ilybius albarracinensis
Ilybius ater
Ilybius chalconatus
Ilybius erichsoni
Ilybius fenestratus
Ilybius fuliginosus
Ilybius guttiger
Ilybius montanus
-Male 2n = 30-35
Ilybius neglectus
Ilybius quadriguttatus
Ilybius subaeneus
Ilybius vittiger
Ilybius wasastjernae
Nebrioporus amicorum
Nebrioporus assimilis
Nebrioporus baeticus
Nebrioporus bucheti
Nebrioporus canaliculatus
Nebrioporus canariensis
Nebrioporus carinatus
Nebrioporus ceresyi
Nebrioporus croceus
Nebrioporus crotchi
Nebrioporus depressus
Nebrioporus depressus elegans int.
Nebrioporus elegans
Nebrioporus fabressei
Nebrioporus insignis
Nebrioporus lanceolatus
Nebrioporus lanceolatus
Nebrioporus martinii
Nebrioporus sardus
Nebrioporus walkeri
Platambus maculatus
Rhantus exsoletus
Rhantus exsoletus
-Xyp Subsequent studies failed to replicate
Rhantus frontalis
Rhantus grapii
Rhantus notatus
-Xyp Subsequent studies failed to replicate
Rhantus suturalis
Rhantus suturellus
Scarodytes fuscitarsis
Scarodytes halensis
Scarodytes malickyi
Scarodytes nigriventris
Stictotarsus bertandi
Stictotarsus duodecimpustulatus
56
Female
2n
Male
2n
44
44
42
42
46
42
42
46
36
34
36
36
36
36
36
36
36
36
34
34
40
43
43
41
41
45
41
41
45
35
33
35
35
35
35
35
35
35
35
33
36
36
36
36
38
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
50
20
42
35
35
35
35
37
49
49
49
49
49
49
49
49
49
49
49
49
49
49
49
49
49
49
49
49
19
41
42
42
46
46
45
45
44
46
42
56
56
56
56
56
56
170
Sex Chrom. Source
Karyotype
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
18
18
18
112
112
18
18
18
18
18
112
112
18
113
18
113
113
18
18
18
113
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
Xyp
113
113
18
113
113
117
117
117
117
117
117
117
117
117
117
117
117
117
117
112
117
112
117
117
112
18
115
118
18
44
XO
XO
XO
Xyp
115
115
118
18
45
41
55
55
55
55
56
NeoXY
XO
XO
XO
XO
XO
XO
NeoXY
117
115
115
117
117
117
117
117
Taxa
Stictotarsus enexpectatus
Stictotarsus griseostriatus strandi
Stictotarsus procerus
Trichonectes otini
Unidentified dytiscid
Elateridae
Adelocera colonicus
Adelocera modesta
Adelocera murina
Adelocera rectangularis
Adelocera sp.
Agriotella bigeminata
Agriotes lineatus
Agriotes mancus
Agriotes obscurus
Agriotes sputator
Agrypnus fuscipes
Agrypnus sp.
Ampedus apicatus
Ampedus deletus
Ampedus fusculus
Ampedus luctuosus
Ampedus melsheimeri
Ampedus pullus
Ampedus sp., nr. deletus
Ampedus sp., nr. Deletus
Ampedus sp., nr. miniipennis
Athous rufiventris
Cardiophorus cardisce
Cardiophorus convexulus
Cardiophorus convexus
Cardiophorus fenestratus
Cardiophorus gagates
Cardiophorus haridwarensis
Cardiophorus limbatus
Cardiophorus togatus
Cardiorhinus rufilateris
Chalcolepidius silbermanni
Chalcolepidius zonatus
Colaulon lezeleuci
Conoderus dimidiatus
Conoderus fuscofasciatus
Conoderus malleatus
Conoderus pilatei
Conoderus rodriguezi
Conoderus rufidens
Conoderus scalaris
Conoderus sp.
Conoderus stigmosus
Conoderus ternarius
Ctenicera aenea
Ctenicera aeripennis aeripennis
Ctenicera aeripennis destructor
Ctenicera appressa
Ctenicera appropinquans
Ctenicera arata
Ctenicera bombycina
Ctenicera hieroglyphica
Ctenicera inflata
Ctenicera mediana
Ctenicera nitidula
Ctenicera ochreipennis
Ctenicera propola columbiana
Sexual
System
Ploidy
62
20
20
20
22
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
60
61
56
50
38
59
XO
56
49
37
XO
120
NeoXY
XO
XO
18
18
22
22
18
18
20
20
20
20
20
18
12
20
20
20
20
20
20
20
22
19
20
22
22
20
22
22
22
22
22
20
12
4
18
18
18
18
18
18
18
18
18
16
14
18
20
19
19
22
20
18
22
22
20
22
18
22
21
17
17
22
22
17
17
19
19
19
19
20
17
11
19
19
19
19
19
19
19
21
XO
19
22
22
20
22
22
21
21
22
19
12
4
17
17
17
17
17
17
17
17
17
16
14
17
19
XO
XO
22
19
17
22
21
20
22
17
22
XO
XO
XO
Xyp
Xyp
XO
XO
XO
XO
XO
XO
Xyp
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
121
XO
Xyp
Xyp
Xyp
Xyp
Xyp
XO
XO
Xyp
XO
XY
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
NeoXY
NeoXY
XO
XO
121
121
Xyp
XO
XO
Xyp
XO
Xyp
Xyp
XO
Xyp
121
171
119
120
117
18
121
121
4
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
122
121
123
121
121
121
121
121
123
121
123
121
123
121
121
121
121
121
121
121
121
121
121
Taxa
Sexual
System
Ploidy
Ctenicera propola propola
Ctenicera rufopleuralis
Ctenicera semimetallica
Ctenicera splendens
Ctenicera tarsalis
Ctenicera tessellata
Dicrepidius politus
Dicrepidius ramicornis
Eanus estriatus
Eanus maculipennis
Elater sp.1
Elater sp.2
Hemirrhipus lineatus
Heterocrepidius mendex
Heteroderes lenis
Heteroderes macroderes
Heteroderes modestus
Heteroderes sericeus
Lacon profusa
Limonius aeger
Limonius griseus
Melanotus fissilis
Melanotus kamaunensis
Melanotus leonardi
Melanotus longicornis
Melanotus oregonensis
Melanotus sp.
Melanotus sp., nr. Communis
Melanotus tenebrosus
Melanotus trapezoideus
Monocrepidius sp.
Monocrepidus sp.
Pomachilius sp.2
Prosternon tesselatum
Pyrearinus candelarius
Pyrophorus divergens
Pyrophorus luminosus
Pyrophorus phosporescens
Pyrophorus punctatissimus
Pyrophorus radians
Erotylidae
Triplax thoracica
Geotrupidae
Anoplotrupes balyi
Anoplotrupes stercorosus
Bolboceras quadridens
Bolboceras sp.
Bolboceratops indicum
Ceratophyus hoffmannseggi
Cnemotrupes splendidus miarophagus
Geotrupes hypocrita
Geotrupes mutator
20
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
18
20
20
20
22
24
24
20
20
20
20
10
XO
XO
XO
XO
Xyp
Xyp
XO
XO
Xyp
Xyp
XO
XO
NeoXY
16
16
12
21
17
19
19
20
22
23
23
20
20
19
19
10
22
17
17
19
17
17
14
20
17
19
12
19
19
19
19
XO
19
19
17
17
20
22
15
15
17
15
15
11
18
18
NeoXY
1
22
22
20
20
20
20
16
22
22
20
20
20
20
16
22
22
22
20
22
22
20
22
20
22
22
22
22
20
XY
XY
Xyp
Xyp
Xyp
Xyp
XY
125
126
127
125
128
125
129
125
125
125
127
125
129
127
125
127
129
129
129
129
127
18
18
20
18
18
14
20
18
20
12
20
20
20
20
19
20
20
18
18
20
22
16
16
22
20
Geotrupes spiniger
22
20
Geotrupes stercocarius
Geotrupes stercorarius
Sericotrupes niger
Thorectes geminatus
Thorectes intermedius
Thorectes lusitanicus
Thorectes punctatissimus
20
22
22
22
22
20
172
XO
XO
XO
XO
XO
NeoXY
Xyp
XO
XO
XY
XO
XO
XO
XO
121
XO
XO
XO
XO
Xyp
XY
XO
XO
XXY
XO
XO
XO
XY
XY
XY
Xyp
XY
XY
XY
XY
XY
XY
121
121
121
121
121
121
121
121
121
121
121
121
121
124
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
121
45
121
121
121
121
121
121
121
121
Taxa
Trypocopris pyrenaeus
Trypocopris vernalis
Typhaeus typhoeus
Glaphyridae
Lichnanthe rathvoni
Glaresidae
Glaresis sp.
Gyrinidae
Dineutes americanus
Dineutes horni
Gyrinus caspius
Gyrinus distinctus
Gyrinus paykulli
Gyrinus substriatus
Gyrinus suffriani
Haliplidae
Brychius elevatus
Haliplus confinis
Haliplus flavicollis
Haliplus fluviatilis
Haliplus fulvus
Haliplus immaculatus
Haliplus laminatus
Haliplus lineatocollis
Haliplus lineatus
Haliplus mucronatus
Haliplus obliquus
Haliplus ruficollis
Haliplus sibricus
Haliplus variegatus
Peltodytes caesus
Helophoridae
Helophorus aequalis
Helophorus aquaticus
-B chromosomes
Helophorus atlantis
Helophorus brevipalpis
Helophorus calpensis
Helophorus grandis
-B chromosomes
Helophorus maritimus
Helophorus minutus
Helophorus oblongus
Helophorus occidentalis
Helophorus orientalis
Helophorus paraminutus
Histeridae
Hister sp.
Saprinus sp.
Hybosoridae
Hybosorus orientalis
Hydrophilidae
Anacaena bipustulata
Anacaena globulus
Anacaena limbata
Anacaena lutescens
Anacaena rufipes
Berosus indicus
Helocombus bifidus
Hydrobius fuscipes
Hydrophilus indicus
Hydrous acuminatus
Sexual
System
parth
parth
parth
parth
Ploidy
3
3
2
3
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
22
20
20
22
20
Xyp
XY
Xyp
127
129
127
20
20
Xyp
125
20
20
Xyp
125
28
28
44
40
27
27
27
27
27
XO
18
18
18
18
18
18
18
40
22
30
24
36
24
24
24
24
20
24
24
24
32
34
39
XO
30
24
36
24
24
24
24
20
24
24
24
32
33
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XO
130
130
130
130
130
130
130
130
130
130
130
130
130
130
130
18
18
18
18
XY
XY
131
131
22
30
22
22
18
22
XY
22
22
18
XY
XY
XY
132
133
133
132
134
18
22
18
22
18
18
XY
XY
22
XY
134
132
1
134
133
132
26
26
26
26
Xyp
Xyp
1
1
20
20
Xyp
125
18
16
18
18
27
18
12
18
18
18
16
18
Xyp
Xyp
Xyp
18
12
18
18
18
30
30
Xyp
NeoXY
Xyp
Xyp
135
135
135
135
135
135
135
1
1
1
1
1
18
33
22
30
30
173
XY
Xyp
Xyr
Taxa
Hydrous indicus
Hydrous piceus
Hydrous triangularis
Sternolophus rufipes
Tropisternus lateralis
Tropisternus nimbatus
Hygrobiidae
Hygrobia hermanni
Laemophloeidae
Cryptolestes ferrugineus
Cryptolestes pusilloides
Cryptolestes pusillus
Cryptolestes spartii
Cryptolestes turcicus
Cryptolestes ugandae
Lampyridae
Aspisoma aegrotum
Aspisoma hesperum
Aspisoma ignium
Aspisoma laterale
Aspisoma maculatum
Aspisoma sticticum
Bicellonycha lividipennis
Cratomorphus dorsalis
Diphotus vittatus
Ellychnia californica
Ellychnia corrusca
Lucidota diaphanura
Luciola cruciata
Luciola lateralis
Photinus australis
Photinus consanguineus
Photinus macdermotti
Photinus pyralis
Photinus sp. 1 (aff. pyralis)
Photinus sp. 2
Photuris congener
Photuris pennsylvanica
Pyractomena angulata
Pyractomena borealis
Pyractomena galeata
Pyractomena heterodoxa
Leiodidae
Cantabrogeus luquei
Cantabrogeus sp
Catops coracinus
Drimeotus kovacsi viehmanii
Espanoliella luquei
Fresnedaella lucius
Leiodes calcarata
Notidocharis whagoni
Pholeuon knirschi
Quaestus pasensis
Speonomus delarouzeei
Speonomus hydrophilus
Speonomus pyrenaeus
Troglocharinus ferreri pallaresi
Troglocharinus jacasi
Troglocharinus kiesenwetteri
Troglocharinus schibii
Troglocharinus variabilis
Lucanidae
Cerchus lignarius
Dorcus parallelipipedus
Figulus binodulus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
30
Xyp
30
18
18
18
30
30
30
18
18
18
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
1
36
36
XY
118
18
18
18
18
18
20
18
18
18
18
18
20
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
1
1
1
1
1
1
20
20
20
20
20
20
18
20
20
20
20
20
18
18
20
20
20
20
20
20
19
19
19
19
19
19
18
19
19
19
19
19
17
17
19
19
19
19
19
19
18
19
19
19
19
19
XO
XO
XO
XO
XO
XO
NeoXY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
136
136
136
136
136
136
136
136
136
136
136
136
136
136
137
137
138
137
136
136
138
136
136
136
136
136
22
33
22
24
22
22
22
22
24
22
24
24
24
24
24
24
24
24
22
Xyp
22
24
22
22
22
22
24
22
24
24
24
Xyp
24
24
24
24
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
141
Xyp
Xyp
Xyp
Xyp
20
18
20
20
18
20
Xyp
NeoXY
XY
20
20
20
20
20
parth
3
24
174
XO
XO
XO
XO
XO
139
139
139
140
139
139
139
139
140
139
141
142
142
141
141
141
141
1
1
143
Taxa
Sexual
System
Figulus boninensis
Figulus punctatus
Lamprima adolphinae
Lucanus cervus
Lucanus maculifermoratus
Macrodorcas binervis
Macrodorcas rectus
Nipponodorcus rubrofemoratus
10
Prosopocoilus inclinatus hachijoensis
Prosopocoilus inclinatus inclinatus
Sinodendron rugosum
Lycidae
Calopteron bifasciatum
Calopteron corrugatum
-B chromosomes
Calopteron scapulare
-B chromosomes
Linoptes imbrex
Metriorrhynchus rhipidius
Thonalmus chevrolati
Melandryidae
Dircaea 4-maculata
Prothalpia undata
Serropalpus barbatus
Serropalpus substriatus
Meloidae
Cyaneolytta n. sp.
Epicauta atomaria
Epicauta cinerea
Epicauta grammica
Epicauta isthmica
Epicauta murina
Epicauta n. sp.
Epicauta pennsylvanica
Epicauta pluvialis
Epicauta rosilloi
Epicauta rufipedes
Lytta picta
Meloe sp.
Mylabris balteata
Mylabris macilenta
Mylabris phalerata
Mylabris pustulata
Mylabris thunbergi
Paniculolytta sanguineoguttata
Pryrota decorata
Psalydolytta sp. nr. Rouxi
Sybaris paraeustus
Sybaris tastaceus
Tetraonyx frontalis
Tetraonyx quadrimaculata
Zonitis tarasca
Melyridae
Astylus antis
Astylus variegatus
-B chromosomes
Collops sp.
Endeodes collaris
Hoppingiana hudsonica
Micromalthidae
Micromalthus deblis
-type of HD not specified
Ploidy
20
20
parth
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
26
18
22
26
14
18
10
10
20
20
18
20
26
18
22
26
14
18
Xyr
10
20
Xyp
18
XY
Xyp
XY
XY
Xyr
Xyp
Xyr
1
NeoXY
Xyp
145
Xyp
143
143
144
144
1
145
145
32
32
31
31
XO
XO
1
1
32
31
XO
1
32
34
33
31
33
XO
XO
1
1
146
20
18
18
20
20
18
18
20
Xyp
NeoXY
Xyp
Xyp
1
1
1
1
20
20
20
20
21
22
20
24
20
20
20
20
20
20
20
20
20
20
22
22
22
22
Xyp
20
20
20
20
20
20
20
Xyp
Xyp
Xyyp
Xyyyp
XY
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
45
45
45
1
1
1
1
1
1
45
45
1
13
1
13
1
1
1
1
18
18
18
Xyp
147
148
18
20
14
17
19
14
XO
XO
Xyp
1
1
1
20
10
20
24
20
20
20
20
20
20
20
20
20
20
22
22
22
22
20
20
20
20
20
20
20
20
175
1
145
1
1
1
1
13
1
1
1
1
Taxa
Nitidulidae
Nitidula bipunctata
Nitidula rufipes
Noteridae
Canthydrus diopthalmus
Neohydrocoptus jaechi
Noterus clavicornis
Noterus crassicornis
Noterus laevis
Synchortus imbricatus
Oedemeridae
Oedemera podagrariae
Oedemera virescens
Passalidae
Aulacocyclus edentulus
Chondrocephalus debilis
Chondrocephalus gemmae
Chondrocephalus granulum
Chondrocephalus purulensis
Coniger ridiculus
Heliscus tropicus
Odontotaenius disjunctus
Odontotaenius striatopunctatus
Odontotaenius zodiacus
Ogyges politus
Oileus bifidus
-B chromosomes
Oileus rimator
Oileus sargi
Passalus aculeatus
Passalus alius
Passalus binomiatus
Passalus coniferus
Passalus glaberrinus
Passalus interruptus
Passalus interstitialis
Passalus mancus
Passalus mirabilis
Passalus morio
Passalus occipitalis
Passalus perplexus
Passalus plicatus
Passalus punctatostriatus
Passalus punctiger
Passalus quadricollis
Passalus sp. 1
Passalus suturalis
Passalus unicornis
Paxillus leachi
Petrejoides mazatecus
Petrejoides nebulosum
-B chromosomes
Petrejoides orizabae
Popilius eclipticus
Proculejus brevis
Proculejus c. brevis 1
-B chromosomes
Proculejus c. brevis 2
-B chromosomes
Proculus beckeri
Pseudacanthus aztecus
Pseudacanthus mexicanus
Pseudacanthus violetae
Ptichopus angulatus
Sexual
System
Ploidy
24
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
20
20
20
Xyp
Xyp
1
1
20
14
22
22
22
28
20
14
21
21
21
27
XY
NeoXY
XXY
XXY
XXY
XXY
149
150
149
149
149
151
22
22
22
22
Xyp
Xyp
4
4
30
38
28
24
44
36
34
26
24
26
30
26
18
30
38
28
24
39
36
34
26
XY
26
30
26
18
XY
XY
XY
XY
XXXXXO
XY
XY
XY
152
XY
XY
XY
XY
152
152
152
152
152
152
152
152
18
18
30
26
26
26
26
26
26
32
26
32
26
26
26
26
26
26
30
26
26
26
38
38
18
18
29
25
25
25
25
25
25
31
25
31
25
25
25
25
25
25
29
25
25
25
38
38
XY
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XY
152
152
152
152
152
153
152
152
152
152
152
153
152
152
152
152
153
152
152
152
154
152
152
152
28
34
28
32
28
34
28
32
XY
XY
XY
XY
152
152
152
152
20
20
XY
152
38
34
40
38
26
38
34
40
38
25
XY
XY
XY
XY
XO
152
152
152
152
152
176
152
152
152
152
Taxa
Sexual
System
Publius agassizi
Spasalus robustus
Spurius bicornis
Spurius halffteri
Undulifer acapulcae
Verres corticicola
Verres hageni
-B chromosomes
Veturius assimilis
Veturius transversus
Vindex agnoscendens
Vindex c. sculptilis
Vindex sp. nov.
Phalacridae
Phalacrid sp.
Pleocomidae
Pleocoma crinita
Pleocoma dubitalis
Pleocoma minor
Pleocoma simi
Ptiliidae
Ptinella aptera
Ptinella errabunda
Ptinella taylorae
Scarabaeidae
Adoretus bombinator
Adoretus decanus
Adoretus duvauceli
Adoretus epipleuralis
Adoretus incurvatus
Adoretus lasiopygus
Adoretus limbatus
Adoretus sp.
Adoretus sp. (M-42)
Adoretus versutus
Adorrhinyptia dorsalis
Adorrhinyptia sp.
Aegiala arenaria
Aegiala blanchardi
Aegialia arenaria
Aeqialia arenaria
Aeqialia blanchardi
Allomyrina dichotoma
Amaurodes passerini
Amphimallon majale
Amphimallon solstitialis
Anomala bengalensis
Anomala corpulenta
Anomala cuprea
Anomala dorsalis
Anomala dubia
Anomala lucens
Anomala luciae
Anomala polita
Anomala ruficapilla
Anomala rufocuprea
Anomala sp.
Anomala superflua
Anomala varicolor
Anomala vestigator
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
26
30
32
38
44
28
28
28
25
30
32
38
44
28
28
XY
XO
XY
XY
XY
XY
XY
XY
152
152
152
152
152
152
152
152
28
28
28
18
18
28
28
28
18
18
XY
XY
XY
XY
XY
152
152
152
152
152
16
15
XO
1
20
20
20
20
20
20
20
20
Xyp
Xyp
Xyp
Xyp
125
125
125
125
155
155
155
parth
22
22
22
22
22
22
22
22
22
22
24
22
16
18
20
20
20
20
20
20
20
18
18
20
20
20
20
20
20
18
20
20
20
20
177
22
22
22
22
22
22
22
22
22
22
24
22
16
18
20
20
20
20
20
20
20
20
20
20
18
18
20
20
20
20
20
20
20
20
18
20
20
20
20
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
Xyr
Xyr
Xyp
XY
Xyp
Xyp
XY
XY
XY
XY
Xyp
XY
Xyr
XY
Xyp
Xyp
XY
Xyp
Xyp
Xyp
Xyp
156
125
125
156
125
125
125
125
125
125
125
125
125
125
125
1
1
127
125
125
125
144
144
125
125
125
125
125
144
125
144
125
125
125
125
125
125
125
125
Taxa
Anomiopsides heteroclyta
Aphodius abdominalis
Aphodius ater
Aphodius bonvouloiri
Aphodius borealis
Aphodius coniugatus
Aphodius conspurcatus
Aphodius consputus
Aphodius constans
Aphodius contaminatus
Aphodius depressus
Aphodius distinctus
Aphodius distintus
Aphodius elevatus
Aphodius erraticus
Aphodius fasciatus
Aphodius fimetarius
Aphodius foetens
Aphodius foetidus
-B chromosomes
Aphodius fossor
Aphodius haemorrhoidalis
Aphodius lapponum
Aphodius lineolatus
Aphodius lividus
-B chromosomes
Aphodius luridus
Aphodius moestus
-B chromosomes
Aphodius nemoralis
Aphodius niger
-B chromosomes
Aphodius obliteratus
Aphodius paganettii
Aphodius paykulli
Aphodius pedellus
-B chromosomes
Aphodius plagiatus
Aphodius prodromus
Aphodius pseudolividus
Aphodius pusillus
Aphodius rufipes
Aphodius rufus
Aphodius scrutator
Aphodius sphacelatus
Aphodius sticticus
Aphodius subterraneus
Aphodius vittatus mundus
Aphodius wilsonae
-B chromosomes
Apogonia carinata
Apogonia ferruginea
Apogonia nigricans
Apogonia proxima
Apogonia sp, nr. nigricans
Apogonia sp.
Apogonia unistraita
Archophileurus vervex
Aserica pilula
Aserica sp.
Ataenius fossor
Ataenius haemorrhoidalis
Ataenius merdarius
Sexual
System
Ploidy
Female
2n
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Male
2n
18
19
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Sex Chrom. Source
Karyotype
XO
XY
XY
Xyp
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
45
157
158
157
158
159
160
161
158
160
160
125
160
125
162
158
163
159
159
20
20
20
20
20
20
20
20
20
20
XY
XY
XY
XY
XY
162
162
158
160
164
20
22
20
22
XY
XY
160
125
20
20
20
XY
158
165
20
20
20
20
20
20
20
20
XY
XY
XY
XY
160
165
160
163
20
20
20
XY
XY
XY
20
20
20
20
20
20
20
20
20
20
20
20
20
20
XY
XY
165
161
164
124
160
158
162
161
160
125
158
165
20
20
20
20
20
20
22
20
20
16
20
20
20
20
20
20
19
19
20
20
19
21
20
20
16
20
19
20
20
20
Xyp
XO
XO
Xyp
Xyp
XO
XO
Xyp
XY
NeoXY
Xyp
XO
XY
XY
XY
124
125
125
125
124
125
125
125
125
45
125
125
125
125
125
20
20
20
20
20
178
XY
Xyp
XY
XY
XY
Taxa
Ataenius rufipus
Ataenius rufus
Ataenius scrutator
Ataenius spretulus
Atheuchus sp
Athyreus excavatus
Autoserica assamensis
Autoserica sp.
Bolbites onitoides
Bothynus striatellus
Bubas bison
Bubas bubaloides
Bubas bubalus
-B chromosomes
Caccobius schreberi
Canthidium breve
Canthochilum andyi
Canthochilum hispidum
Canthochilum histeroides
Canthochilum oakleyi
Canthon aff carbonarius
Canthon chalybaeus
Canthon indigaceus
Canthon muticus
Canthon septemmaculatus
Canthon staigi
Catharsius aff. Sagax
Catharsius molossus
Catharsius pithecius
Catharsius sagax
Catharsius sp1
Catharsius sp2
Catharsius sp3
Cetonia aurata
Cetonia aurataeformis
Cetonia roelofsi
Chalcosoma atlas
Cheironitis furcifer
Chiron digitatus
Chlorocala africana
Clinteria spilota
Coenochilus trabecula
Copris fricator
Copris frictator
Copris hispanus cavolinii
Copris hispanus hispanus
Copris incertus
Copris lugubris
Copris lunaris
Copris sinicus
Copris sp.
Copris tullius
Coprophanaeus cyanescens
Coprophanaeus dardanus
Coprophanaeus ensifer
Cotalpa lanigera
Cremastocheilus armatus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
20
20
20
20
16
20
30
18
20
20
20
20
20
20
XY
20
20
18
18
18
18
18
20
20
18
20
20
18
20
20
20
20
20
20
18
20
20
20
20
20
20
20
20
20
14
20
20
21
21
19
18
18
18
14
14
20
14
14
20
20
20
20
20
20
20
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
Xyp
Xyp
Xyp
Xyp
Xyr
Xyp
XY
20
16
20
30
18
20
20
20
20
20
20
20
20
18
18
18
18
18
20
20
18
20
20
18
20
20
20
20
20
20
18
20
20
20
20
20
20
20
14
20
22
22
18
18
18
14
14
20
14
14
20
20
20
20
20
20
179
XY
XY
Xyp
XY
Xyp
Xyp
Xyp
Xyp
XY
XY
XY
NeoXY
XY
XY
Xyp
XY
Xyp
XO
XO
XY
XY
XY
Xyp
Xyp
XY
Xyp
Xyp
Xyp
XY
XY
XY
Xyp
Xyp
125
125
125
125
166
125
125
125
125
45
45
167
167
167
168
125
45
1
1
1
125
169
169
1
45
45
166
1
1
125
125
125
125
1
1
1
125
170
125
144
1
125
1
144
125
125
125
1
1
171
171
167
125
1
167
167
125
125
172
169
172
125
125
125
Taxa
Sexual
System
Cyclocephala insulicola
Cyclocephala lutea
Cyclocephala maffafa grandis
Cyclocephala melanocephala rubiginosa
Cyclocephala putrida
Cyclocephala tridentata dominicensis
Cyclocephala tridentata tridentata
Cyprolais hornimani
Deltochilum aff amazonicum
Deltochilum aff morbillosum
Deltochilum calcaratum
Ploidy
20
14
Dichotomius aff mundus
Dichotomius aff sericeus
Dichotomius affinis
Dichotomius bos
Dichotomius bosqui
Dichotomius carolinus
Dichotomius crinicollis
Dichotomius depresicollis
Dichotomius geminatus
-B chromosomes
Dichotomius laevicollis
Dichotomius mormon
Dichotomius nisus
Dichotomius semianeus
Dichotomius semisquamosus
Dichotomius sericeus
Diloboderus abderus
Diplotaxis obscura
Diplotaxis sierrae
Diplotaxis sp.
Dynamopus athleta
Dynastes hercules hercules
Dynastes tityus
Dyscinetus bidentatus
Dyscinetus gagates
Dyscinetus rugifrons
Ectinohoplia rufipes
Enema pan
Eophileurus chinensis
Eophileurus platypterus
Epicometis hirta
-B chromosomes
Epicometis squalida
Male
2n
Sex Chrom. Source
Karyotype
20
20
20
20
20
20
20
20
20
14
14
20
20
Xyp
20
20
20
20
20
20
NeoXY
Xyp
Xyp
173
XY
Xyp
Xyp
XY
Xyp
XY
169
175
20
173
45
144
174
45
144
173
144
20
18
18
18
14
14
20
14
14
20
20
20
18
18
18
18
20
20
18
18
18
18
18
18
18
18
18
18
18
18
20
20
20
20
20
20
18
18
18
18
18
18
18
18
18
20
20
XY
20
20
20
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
Xyp
XY
XY
144
XY
Xyr
XY
178
178
178
180
178
180
178
180
178
144
144
20
20
20
20
20
20
22
18
18
20
20
20
20
18
Xyp
Xy/Xyp/Xyr
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
NeoXY
Xyp
Xyp
Xyp
Xyr
Xyp
20
20
20
20
20
20
20
20
20
22
18
18
20
20
20
20
18
20
20
20
20
Xyp
XY
Xyp
169
166
45
125
125
125
125
181
144
45
45
45
125
45
125
125
125
125
20
20
Xyp
125
14
20
14
14
20
20
20
18
18
18
18
Deltochilum elevatum
Deltochilum morbillosum
Deltochilum valgum
Deltochilum verruciferum
Diabroctis mimas
Dichotomius sp
Dichranocephalus wallichii
Dicronorhina derbyana
Dicronorhina derbyana oberthuri
Dicronorhina micans
Digitonthophagus bonasus
Digitonthophagus gazella
-B chromosomes
Female
2n
180
NeoXY
Xyp
NeoXY
NeoXY
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
175
166
166
166
45
176
169
177
178
178
178
45
45
1
178
178
179
144
1
168
Taxa
Sexual
System
Eucranium arachnoides
Eudicella aethiopica
Eudicella gralli
Eudicella smithi
Euetheola humilis
Euonticellus fulvus
Euonticellus pallipes
Euothonphagus amyntas
-B chromosomes
Euothonphagus atramentarius
Euphoria inda
Euphoria lurida
Eurysternus caribaeus
Exomala hirtella
Geniates borelli
Glycyphana fulvistemma
Glyphoderus sterquilinus
Gnorimus nobilis
Gnorimus variabilis
Goliathus goliathus
Gromphas lacordairei
Gymnetis pantherina
Gymnopleurus cyaneus
Gymnopleurus geoffroyi
Gymnopleurus koenigi
Gymnopleurus sturmi
Haplidia etrusca
Haplidia transversa
Heliocopris bucephalus
Heliocopris gigas
Holotrichia longipennis
Holotrichia problematica
Holotrichia serrata
Hoplia communis
Hoplia uniformis
Isocopris inhiatus
Jumnos ruckieri
Lachnosterna longipennis
Leucothyreus guadulpiensis
Leucothyreus nolleti
Ligyrodes relictus
Ligyrus cuniculus
Ligyrus gibbosus burmeisteri
Lygirus ebenus
Lyogenys fuscus
Macraspis dichroa cribata
Macraspis festiva
Macraspis moreo
Macraspis tristis
Maladera alcocki
Malagoniella aff astianax
Mecynorrhina polyphemus confluens
Mecynorrhina torquata
Megalorrhina harrisi
Megasoma actaeon
Melolontha hippocastani
Melolontha melolontha
Melolontha pectoralis
Microcopris doriae
Microcopris hidakai
Mimela glabra
Mimela sp.
Oniticellus fulvus
Oniticellus pallipes
Onitis belial
Ploidy
20
20
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
20
20
20
20
20
20
20
18
20
20
20
20
20
20
20
Xyp
XY
XY
XY
Xyp
20
20
20
8
20
20
20
18
20
XY
20
20
8
20
20
20
18
20
20
20
20
20
20
20
20
18
20
20
20
18
20
20
20
20
20
20
20
20
20
18
20
20
20
18
20
20
20
20
18
14
20
20
20
20
20
Xyp
20
20
18
18
20
18
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
168
Xyp
Xyp
NeoXY
XY
Xyp
XY
Xyp
XY
NeoXY
XY
Xyp
XY
Xyp
Xyp
Xyp
XY
NeoXY
XY
XY
Xyp
Xyp
Xyp
Xyp
20
18
14
20
20
20
20
20
20
20
18
18
18
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
181
XY
XY
XY
Xyp
NeoXY
Xyp
XY
XY
Xyp
Xyp
45
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
XY
Xyp
XY
XY
XY
Xyp
Xyp
Xyp
Xyp
Xyp
XY
45
144
144
144
45
168
168
168
125
45
182
144
183
125
45
144
49
144
45
144
1
167
125
184
125
144
1
167
125
156
125
125
144
177
185
156
144
144
125
173
183
186
45
177
187
188
156
169
144
144
144
173
125
189
189
167
167
156
125
125
125
167
Taxa
Sexual
System
Onitis crassus
Onitis ion
Onitis philemon
Ontherus appendiculatus
Ontherus sulcator
Onthophagus albicornis
Onthophagus amyntas
Onthophagus andalusicus italicus
Onthophagus bifasciatus
Onthophagus catta
Onthophagus coenobita
Onthophagus crassus
Onthophagus dama
Onthophagus fracticornis
Onthophagus furcatus
Onthophagus hecate
Onthophagus hirculus
Onthophagus hirtus
Onthophagus illyricus
Onthophagus joannae
Onthophagus lemur
Onthophagus lucidus
Onthophagus maki
Onthophagus marginicollis
Onthophagus massai
Onthophagus mopsus
Onthophagus mopsus gracillicornis
Onthophagus nuchicornis
Onthophagus opacicollis
Onthophagus ovatus
Onthophagus pacificus
Onthophagus pennsylvanicus
Onthophagus punctatus
Onthophagus quaestus
Onthophagus ramosellus
Onthophagus rufescens
Onthophagus ruficapillus
Onthophagus similis
-B chromosomes
Onthophagus sp. 1
Onthophagus sp. 2
Onthophagus sp. 3
Onthophagus sp. 4
Onthophagus stylocerus
Onthophagus taurus
Onthophagus vacca
-B chromosomes
Onthophagus verticicornis
Onthophagus verticornis
Ophthalmo serica karafutoensis
Orizabus cultripes
Orphnus impressus
Orphnus mysoriensis
Oruscatus davus
Oryctes nasicornis
Oryctes rhinoceros
Osmoderma eremita
-B chromosomes
Osmoderma lassallei
Osmoderma scabra
-B chromosomes
Oxysternon silenus
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Xyp
XY
Xyp
Xyp
Xyp
XY
Xyp
XY
Xyp
Xyp
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Xyp
Xyp
Xyp
XY
XY
1
167
125
166
45
168
125
125
156
125
1
168
125
125
190
168
125
45
168
168
168
125
168
168
1
171
125
1
1
190
168
156
125
125
125
125
1
168
190
20
20
18
18
20
20
20
20
20
20
18
18
20
20
20
20
Xyp
Xyp
Xyp
Xyp
XY
XY
Xyp
XY
1
1
1
1
168
168
125
168
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Xyp
Xyp
Xyr
XY
Xyp
Xyr
XY
XY
Xyp
Xyp
XY
XY
XY
XY
XY
XY
Xyp
Xyp
20
20
18
18
20
20
18
18
20
18
18
20
20
20
18
18
20
18
18
18
20
182
125
1
125
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
NeoXY
125
125
125
45
125
144
125
144
18
18
NeoXY
NeoXY
144
49
19
XO
169
Taxa
Sexual
System
Ploidy
Oxythyrea funesta
Paracopris ramosiceps
Paragymnopleurus sinuatus
Pelidnota pallidipennis
Pelidnota punctata
Pelidnota sumptuosa
Pentodon bidens punctatus
Pentodon bispinifrons
Pentodon idiota
Pentodon sp.
Phalops divisus
Phalops olivaceous
Phanaeus aff yucatanus
Phanaeus chalcomelas
Phanaeus daphnis
Phanaeus ensifer
Phanaeus igneus
Phanaeus mexicanus
Phanaeus splendidulus
Phanaeus vindex
Phileurus didymus
Phileurus sp.
Phileurus valgus guadulpiensis
Phyllognathus dionysius
Phyllognathus silensis
Phyllopertha campestris
Phyllophaga aff capillata
Phyllophaga anxia
Phyllophaga delata
Phyllophaga drakii
Phyllophaga fusca
Phyllophaga gracilis
Phyllophaga pleei
Phyllophaga sandersoniella
Phyllophaga sp. crenulata group
Phyllophaga tristis
Phyllophaga vestita
Plaesiorrhinella watkinsiana
Pocalta ursina
Popillia japonica
Propomacrus bimucronatus
Protaetia aeruginosa
Protaetia angustata
Protaetia aurichalcea
Protaetia cuprea bancoi
Protaetia cuprea cuprea
Protaetia cuprea metallica
Protaetia cuprea obscura
Protaetia fieberi
Protaetia lugubris
Protaetia mirifica
Protaetia morio
Protaetia oblonga
Protaetia opaca
Protaetia speciosa
Psammodius oregonesis
Psammoporus sabuleti
Rhamphorrhina bertolonii
Rhinyptia indica
Rhomborrhina polita
Rhomborrhina unicolor
Rutela striata
Scarabaeidae nr. Autoserica and Neoserica
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
20
18
18
20
20
20
20
20
20
20
20
20
14
12
12
19
20
18
18
20
20
20
19
20
20
19
20
20
14
12
12
20
12
12
20
12
12
16
14
XY
20
18
20
20
20
20
20
20
20
20
20
XO
XY
Xyp
Xy+
Xyp
XY
Xyr
XO
Xyp
XY
XO
Xyp
Xyp
Xyp
XY
NeoXY
12
12
20
12
12
16
20
20
20
18
20
20
20
20
20
20
20
20
20
20
18
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
183
20
20
20
20
18
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Xyp
XY
NeoXY
Xyp
XY
NeoXY
NeoXY
144
Xyp
NeoXY
XY
XY
XY
XY
XY
XY
125
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
Xyp
XY
XY
Xyp
XY
125
44
125
167
1
183
125
187
191
125
144
125
128
128
1
169
1
187
125
1
166
125
125
144
187
125
125
125
186
125
125
125
125
125
189
144
125
186
144
125
125
144
170
144
144
170
170
170
170
170
170
170
170
170
170
170
125
127
144
156
125
125
144
Taxa
Sexual
System
Ploidy
Scarabaeus cristatus
Scarabaeus laticollis
Scarabaeus sacer
Scarabaeus semipunctatus
Schizonycha fuscescens
Schizonycha ruficollis
Serica assamensis
Serica falli
Serica sericea
Serica tristis
Serica umbrinella
Sisyphus schaefferi
Stephanorrhina guttata
Stephanorrhina princeps
Strategus surinamensis hirtus
Strategus syphax
20
Strategus validus
Sulcophanaeus imperator
Sulcophanaeus menelas
Tiniocellus spinipes
Trichiotinus assimilis
Trichius fasciatus
Trichius rosaceus zonatus
Trichius sexualis
Trichius succinctus
Trichius zonatus
Tropinota hirta
-B chromosomes
Xylotrupes gideon
Scolytidae
Blastophagus minor
Male
2n
Sex Chrom. Source
Karyotype
20
20
20
20
20
20
20
22
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
22
20
20
20
20
20
20
20
20
Xyp
20
20
20
20
20
24
20
20
20
20
20
20
20
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
XY
XY
183
XY
Xyp
20
20
XY
144
26
30
26
30
24
26
30
26
30
23
Xyp
Xyp
Xyp
Xyp
XO
1
1
1
1
1
192
20
20
24
20
20
20
20
20
Blastophagus pinierda
Cactopinus desertus
Coccotrypes dactyliperda
-type of HD not specified
Coccotrypes declivis
-based on one brood
Coccotrypes graniceps
-type of HD not specified
Conopthorus coniperda
Conopthorus resinosae
Conopthorus sp. 1
Cryphalus abietis
Cryphalus piceae
Crypturgus pusillus
Dendroctonus adjunctus
Dendroctonus approximatus
Dendroctonus brevicomis
Dendroctonus frontalis
Dendroctonus jeffreyi
-B chromosomes
Dendroctonus mexicanus
Dendroctonus murrayanae
Dendroctonus parallelocollis
Dendroctonus ponderosae
Dendroctonus pseudotsugae
Dendroctonus punctatus
Dendroctonus rhizophagus
Dendroctonus rufipennis
Dendroctonus simplex
Female
2n
NeoXY
NeoXY
Xyp
Xyp
XY
XY
XY
XY
haplodiploid
193
haplodiploid
194
20
18
167
125
167
125
125
125
125
125
156
125
125
125
156
1
144
144
144
173
187
45
45
125
125
125
144
144
125
125
170
NeoXY
XY
26
26
26
14
12
12
16
24
20
18
18
26
26
26
14
12
12
16
24
Xyp
Xyp
Xyp
NeoXY
NeoXY
NeoXY
Xyp
NeoXY
1
1
1
1
1
1
1
1
1
1
195
12
30
28
24
30
32
28
30
30
12
30
28
24
30
32
28
30
30
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
196
1
196
195
1
1
196
1
1
184
Taxa
Sexual
System
Ploidy
Female
2n
26
28
28
26
28
Hylastes angustatus
26
Hylastes ater
26
Hylastes cunicularius
26
Hylastes varius
26
Hylurgops glabratus
26
Hylurgops palliatus
26
Hylurgops pinifex
30
Hypothenemus hamperi
PGE
14
-in males one set of chromosomes is always highly condensed
14
-Subsequent studies failed to replicate
Ips acuminatus
16
Ips avulsus
34
-B chromosomes
Ips bonanseai
32
Ips borealis
32
Ips calligraphus calligraphus
32
32
Ips calligraphus ponderosae
32
32
Ips concinnus
16
Ips confusus
32
Ips cribricollis
32
Ips emarginatus
32
Ips hoppingi
32
Ips hunteri
32
Ips integer
32
Ips interstitialis
32
Ips knausi
32
Ips latidens
20
Ips lecontei
32
Ips mexicanus
16
Ips montanus
32
Ips Ochagnoni Sw.O
32
Ips paraconfusus
32
Ips perroti
32
Ips pertubatus
parth
48
32
Ips pilifrons
32
Ips pini
30
32
Ips pini (=oregonis Eichh.)
32
Ips plastographus maritimus
32
32
Ips plastographus plastographus
32
Ips sexdentatus
26
Ips spinifer
20
Ips tridens
parth
48
32
Ips typographus
26
30
Ips woodi
32
Leperisinus pruinosis
26
Orthotomicus caelatus
22
Orthotomicus erosus
16
Orthotomicus laricis
16
Orthotomicus proximus
14
Orthotomicus sabinianae
Orthotomicus suturalis
16
Phloeotribus scarabaeoides
26
Pityogenes bidentatus
16
Pityogenes chalcographus
16
20
Dendroctonus terebrans
Dendroctonus valens
Dryocoetes affaber
Dryocoetes autographus
185
Male
2n
Sex Chrom. Source
Karyotype
26
28
28
26
28
26
26
26
26
26
26
30
14
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
1
1
1
1
1
1
1
197
15
XXY
1
16
32
Xyp
Xyp
1
1
32
32
Xyp
Xyp
16
32
32
32
32
32
32
32
32
20
32
16
32
32
32
32
Xyp
Xyp
1
1
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
32
32
30
32
32
Xyp
32
26
20
Xyp
Xyp
Xyp
Xyp
Xyp
1
Xyp
Xyp
Xyp
32
26
30
32
26
22
16
16
14
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
16
26
16
16
20
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Taxa
Sexual
System
Ploidy
Pityogenes crannulatus
Pityogenes fossifrons
- XpneoXneoYp
Pityogenes hopkinsi
Pityogenes knechteli
Pityogenes quadridens
Pityokteines ornatus
Pityphthorus lichtensteini
Pityphthorus sp. 1
Polygraphus grandiclava
Polygraphus jezoensis
Polygraphus poligraphus
Polygraphus rugipennis
Polygraphus subopacus
Scolytus dahuricus
Scolytus mali
Scolytus multistriatus
Scolytus ratzeburgi
Scolytus rugulosus
Trypodendron lineatum
Xyleborus dispar
Xylosandrus affinis
haplodiploid
-evidence: sex ratio and/or all male broods
Xylosandrus compactus
haplodiploid 20
-type of HD not specified
Xylosandrus dispar
haplodiploid
-evidence: sex ratio and/or all male broods
Xylosandrus ferrugineus
haplodiploid
-evidence: sex ratio and/or all male broods
Xylosandrus germanus
haplodiploid 16
-type of HD not specified
Silphidae
Nicrophorus sayi
Nicrophorus vespilloides
Phosphuga atrata
Silpha americana
Silpha noveboracensis
Silpha perforata
Silpha surinamensis
Thanatophilus sinuatus
Silvanidae
Ahasverus advena
Oryzaephilus mercator
Oryzaephilus surinamensis
Staphylinidae
Aleochara sp.
Batrisodes globosus
Creophilus maxillosis villosus
36
Creophilus maxillosus
Nudobius cephalus
Ontholestes cingulatus
Philonthus fuscipennis
Philonthus intermedius
Philonthus politus
Philonthus varius
Platyprosophus tamulus
Pselaphus fustifer
Quedius fuliginosus
Staphylinus sp.
Staphylinus violaceus
Tachinus lignorum
Stenotrachelidae
Cephaloon lepturides
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
20
18
19
Xyp
XXY
1
1
20
20
16
20
18
26
20
20
16
20
18
26
28
16
24
16
12
16
22
12
16
12
16
16
26
40
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
16
16
12
16
22
12
16
12
16
16
26
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
193
10
1
193
193
8
1
14
14
34
40
26
40
26
26
13
13
34
40
26
40
26
26
XO
XO
Xyp
XY
Xyp
XY
Xyp
Xyp
198
198
198
198
198
198
1
198
18
18
18
18
18
18
Xyp
Xyp
Xyp
1
1
1
18
28
36
40
28
26
24
56
40
32
20
30
34
40
44
26
18
28
XY
40
28
26
24
56
40
31
20
30
33
40
44
26
NeoXY
Xyp
198
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XO
XY
Xyp
XO
Xyp
XY
XY
198
1
18
18
Xyp
1
186
45
198
198
198
198
198
198
198
1
198
45
198
198
Taxa
Tenebrionidae
Adesmia biskrensis
Adesmia cancellata
Adesmia metallica
Adesmia montana
Aemymone cariosa
Agapanthia korostelevi
Agapanthia walteri
Akis acuminata
Akis bacarozzo
Akis bremeri
Akis discoidea
Alobates pennsylvanica
Alphasida depressa
Alphasida ibicensis
Alphasida sp.
Alphitobius diaperinus
Alphitobius pliceus
Alphitobius sp.
Ammobius rufus
Arthrodeis curtus
Arthrodeis inflatus
Arthrodeis punctulatus
Arthromacra aenea
Asida cardonae
Asida glacialis
Asida glacialis rustica
Asida jurinei
Asida maraguesi
Asida planipennis
Belopus elongatus
Blaps bedeli torres-salai
Blaps cribrosa
Blaps gibba
Blaps gigas
Blaps judaeorum
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
20
18
20
18
20
20
16
16
16
16
20
20
18
20
18
20
18
20
20
16
16
16
16
20
20
20
18
19
20
20
20
20
20
20
14
20
20
20
20
20
20
20
34
36
38
35
19
21
37
19
35
36
36
34
36
34
25
20
19
35
20
20
18
18
20
20
20
20
14
26
26
20
20
20
20
20
Xyp
Xyp
Xyp
XY
Xyp
Xyp
Xyp
NeoXY
NeoXY
NeoXY
NeoXY
Xyp
Xyp
18
20
20
20
20
20
20
20
14
20
20
20
20
20
20
44
38
Blaps lethifera
Blaps lusitanica
Blaps mortisaga
Blaps mucronata
Blaps sulcata
Blaps tenuicollis
Blaps waltli
Blaps wiedemanni
Blapstinus sp.
Bolitopherus cornutus
Caenoblaps nitida
Clitobius ovatus opacus
Coelometopus clypeatus
Cossyphus depressus
Cossyphus hoffmannseggi
Cryptycus gibbulus
Cynaeus angustus
Dailognatha pumila
Derosphaerus cribum
Diaperis boleti
Diastolinus fortipes
Elenophorus collaris
Eleodes armata
Eleodes cordata
Eleodes dentipes
Erodius emondi
Erodius emondi laevis
20
20
20
18
20
20
20
20
14
26
26
20
20
20
20
20
187
Xyp
XO
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
XXXY
XXXXXXXY
XXXXY
XXY
XXY
XXY
XXY
XXXXY
XXXY
XXXY
XXXY
XXXXYY
XXXY
XXXXY
Xyp
XO
XXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
NeoXY
NeoXY
NeoXY
Xyr
Xyp
199
199
199
200
45
107
107
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
1
201
201
201
199
201
199
199
201
199
202
202
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
201
199
199
199
199
107
199
199
45
199
199
199
199
199
199
Taxa
Sexual
System
Erodius exilipes
Erodius lefranci
Erodius nitidicollis
Erodius orientalis
Eulabris sp.
Glabrasida goudoti
Glabrasida melillense
Glabrasida zapateri
Gnaptor spinimanus
Gnathocerus cornutus
Gnathocerus maxillosus
Gonocephalum bilineatum
Gonocephalum depressum
Gonocephalum dorsogranosum
Gonocephalum elongatum
Gonocephalum hoffmannseggi
Gonocephalum oblongum
Gonocephalum parallelum
Gonocephalum patruele
Gonocephalum rusticum
Gonocephalum sp.
Gonocephalum vagum
Hegeter amaroides
Hegeter brevicollis
Hegeter costipennis
Hegeter fernandezi
Hegeter grancanariensis
Hegeter lateralis
Hegeter politus
Hegeter tenuipunctatus
Hegeter transversus
Hegeter tristis
Himatismus fasciculatus
Hoplobrachium asperipenne
Hoplobrachium dentipes
Hylocrinua sp.
Isocerus balearicus
Isomira quadrastriata
Isomira sp. 1
Isomira variabilis
Laena reitteri
-B chromosomes
Latheticus oryzae
Leptonychus curvicornis
Melanochrus blairi
Melanochrus lacordairei
Mesomorphus villiger
Mesostema angustata praesahariana
Mesostema lineatopunctata
Misolampus goudoti erichsoni
Misolampus subglaber
Morica hybrida
Morica planata
Navicularis latihumeralis
Nesotes conformis grancanariensis
Nesotes gomerensis
Nesotes picescens
Nesotes porrectus
Nesotes viridicollis
Nyctelia rugosa
Nyctelia sp.
Nyctobates gigas
Opatrinus aciculatus
Opatrinus validus
Opatroides vicinus
Ploidy
20
20
20
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
20
20
20
20
20
18
20
38
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
22
22
20
20
20
20
14
18
20
20
20
20
20
20
18
20
36
20
20
20
20
Xyp
20
Xyp
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
22
22
20
20
20
20
14
18
Xyp
Xyp
Xyr
Xyp
Xyp
Xyp
Xyp
Xyp
XXXY
Xyp
Xyp
Xyp
Xyp
199
Xyr
199
Xyp
XY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
199
199
199
199
199
201
199
201
49
199
199
199
199
20
20
20
20
20
20
20
20
20
16
16
20
20
20
20
20
20
18
18
18
20
20
20
20
20
20
20
19
20
20
Xyp
20
16
16
20
20
20
20
20
20
18
18
18
20
20
20
Xyp
Xyp
Xyp
Xyp
XO
Xyr
Xyp
199
Xyp
NeoXY
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
Xyp
Xyp
XY
199
199
199
199
199
199
199
188
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
1
1
1
203
201
199
199
199
199
199
199
199
199
45
45
204
199
199
199
Taxa
Oxycara mellyi
Oxycarops fuscipes
Pachycera buprestroides
Pachychila dejeani
Pachychila frioli
Pachychila germari haroldi
Pachychila sublunata
Pachyscelis musiva
Paivaea hispida
Palembus dermestoides
Palorus subdepressus
Phaleria acuminata
Phaleria variabilis
Phylan abbreviatus
Phylan gibbulus
Phylan mediterraneus
Phylan nitidicollis
Phylan obesus
Phylan semicostatus
Pimelia angulata latesti
Pimelia aranacea
Pimelia ascendens
Pimelia atlantis atlantis
Pimelia atlantis frigioides
Pimelia baetica
Pimelia bipunctata
Pimelia boyeri
Pimelia canariensis
Pimelia capito
Pimelia cephalenica
Pimelia costata
Pimelia cribra
Pimelia echidna
Pimelia elevata
Pimelia estevezi
Pimelia fernandez-lopezi
Pimelia fornicata
Pimelia grandis echidniformis
Pimelia grandis latesti
Pimelia granulicollis
Pimelia grossa
Pimelia inexpectata
Pimelia integra
Pimelia interjecta
Pimelia interstitialis
Pimelia laevigata costipennis
Pimelia laevigata laevigata
Pimelia laevigata validipes
Pimelia lutaria
Pimelia maura
Pimelia mauritanica
Pimelia modesta
Pimelia monticola
Pimelia radula ascendens
Pimelia radula oromii
Pimelia radula radula
Pimelia rugosa
Pimelia scabricollis
Pimelia scabrosa
Pimelia sericella
Pimelia servilei
Pimelia sparsa albohumeralis
Pimelia sparsa serrimargo
Sexual
System
Ploidy
20
18
18
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
20
21
20
22
20
20
20
20
20
18
20
20
20
20
20
20
20
20
26
20
26
18
18
18
18
18
18
18
18
18
18
18
18
20
18
20
18
18
18
Xyp
18
18
18
18
18
20
18
Xyp
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
Xyp
18
Xyp
Xyy
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
Xyr
Xyp
Xyp
Xyp
Xyp
Xyr
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
199
Xyr
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
207
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyr
207
Xyp
20
22
20
20
20
20
20
18
20
20
20
20
20
20
20
20
26
20
26
18
18
18
18
18
18
18
18
18
18
18
18
20
18
20
18
18
18
20
18
18
18
18
18
20
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
18
189
199
205
199
199
199
199
199
199
199
107
199
206
199
199
199
199
199
199
199
199
199
199
199
207
207
207
207
199
207
207
107
199
207
207
207
207
207
207
207
199
207
199
199
207
207
199
207
207
207
207
199
199
207
199
207
207
207
199
207
199
199
207
Taxa
Pimelia sparsa sparsa
Pimelia subquadrata valdani
Pimelia variolosa
Platydema impressifrons
Platynotus excavatus
Platynotus punctatipennis
Praocis compacta
Probaticus ebeninus
Pseudoblaps meteeyi
Pseudolamus seriatoporus
Rhytinota sp.
Scaurus punctatus
Scaurus striatus
Scaurus vicinus
Scleron asperulum
Scleron ganutipenne
Scleron reitteri
Scleron sp.
Sexual
System
Ploidy
18
Scotobates calcaratus
Scotobius miliaris
Scotobius muricatus
Scotobius tristis
Sitophagus holoeptoides
Sphaenariopsis impolita
Sphaenariopsis tristis
Spyrathus sp.
Stenosis intricata
Tenebrio molitor
Tenebrio obscurus
Tenebrio picipes
Tentyria grossa
Tentyria laevis
Tentyria latreillei
Tentyria mucronata
Tentyria ophiusae
Tentyria rotunda
Tentyria schaumi
Tentyria subcosta
Trachycelis aphodiodes
Tribolium anaphe
Tribolium audax
-B chromosomes
Tribolium brevicornis
Tribolium castaneum
Tribolium confusum
Tribolium destructor
Tribolium freemani
Tribolium madens
-B chromosomes
Uloma impressicollis
Upis ceramboides
Uyttenboogaartia punctipennis
Zophobas aff. confusus
Zophosis bicarinata
Tetratomidae
Penthe obliquata
Torridincolidae
Ytu zeus
-B chromosomes
Trachypachidae
Trachypachus holmbergi
20
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
18
18
20
20
20
20
20
20
20
20
20
24
24
20
22
22
20
22
18
14
18
18
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
22
Xyp
199
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
NeoXY
NeoXY
NeoXY
NeoXY
Xyp
Xyp
XY
Xyr
NeoXY
Xyp
Xyp
NeoXY
Xyp
Xyp
Xyp
Xyp
Xyp
XY
Xyp
Xyr
Xyr
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
20
18
Xyr
18
20
20
20
20
20
20
20
20
20
24
24
20
22
22
20
22
18
14
18
18
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
22
18
20
20
18
18
20
20
207
Xyp
207
199
199
199
199
201
199
199
199
199
199
199
199
199
199
199
199
199
199
45
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
199
208
199
18
20
18
18
20
20
Xyp
NeoXY
NeoXY
Xyp
XY
208
199
199
199
199
199
20
20
20
20
20
20
20
Xyp
20
20
Xyp
Xyp
199
Xyp
Xyp
45
199
16
16
16
16
XY
NeoXY
1
1
20
20
Xyp
209
38
37
XO
210
190
204
199
Taxa
Trachypachus slevini
Trogidae
Trox borrei
Trox costatus
Trox foveicollis
Trox granulatus
Trox monachus
Trox omacanthus
Trox oricensis
Trox punctatus
Trox scaber
Trox scutellaris
Trox spinulosus dentibius
Trogossitidae
Lophoceters pusillus
Tenebroides mauritanicus
Zopheridae
Zopherus haldemani
Dermaptera
Arixenidae
Arixenia esau
Forficulidae
Anechura bipunctata
Apterygida albipennis
Forficula auricularia
Forficula auricularia
Forficula auricularia
Forficula auricularia
Forficula auricularia
Forficula auricularia
Forficula auricularia
Forficula scudderi
Forficula smyrnensis
Forficula sp
Forficula auricularia
Forficula auricularia
Forficula auricularia
Forficula scudderi
Forficula smyrnensis
Pseudochelidura sinuata
Hemimeridae
Hemimerus bouvieri
Labiduridae
Anisolabis annuipes
Anisolabis marginalis
Anisolabis maritima
Anisolabis sp
Euborellia annulipes
Euborellia moesta
Euborellia stali
Gonolabis brunneri
Labidura bengalensis
Labidura riparia
Labidura riparia
Labidura truncata
Labidura bidens
Labidura riparia
Labidura truncata
Nala lividipes
Nala lividipes
Nala lividipes
Notolabis occidentalis
Parisopsalis spryi
Labiidae
Chaetospania brunner
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
38
37
XO
210
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
20
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
Xyp
45
156
125
125
125
125
125
125
125
125
125
18
24
18
24
Xyp
Xyp
1
1
16
16
XY
199
60
60
XY
252
24
24
24
24
24
25
24
24
24
XY
XXXY
XXY
24
24
21
27
24
24
25
24
21
25
XY
XXY
XXY
XXY
XXY
XY
XXY
XXY
XXY
XXY
XXY
Xyy
XXY
XXY
252
252
253
253
253
253
253
253
253
254
254
255
252
252
252
252
252
252
8
7
XXY
252
26
26
26
44
26
26
26
24
14
12
12
10
12
14
10
34
40
38
24
34
25
25
25
42
25
25
25
24
14
12
12
10
12
14
10
34
40
37
24
32
complex XY
XXY
XXY
XXXY
XXY
XXY
XXY
XXY
XY
XY
XXY
XXY
XY
XY
XY
XY
XXY
XXY
XY
XXXY
253
252
252
252
252
252
252
252
253
256
257
258
252
252
252
252
252
252
252
252
34
31
XXXXY
252
26
26
24
29
28
24
24
24
24
22
26
191
25
25
Taxa
Chaetospania sp.1
Chaetospania sp.2
Labia minor
Labia sp.
Nesogaster erichsoni
Nesogaster halli
Prolabia arachidis
unknown Sp.
Pygicranidae
Dacnodes shortridgei
Diplatys gladiator
unknown sp.
Diptera
Agromyzidae
Amauromyza flavifrons
Amauromyza verbasci
Cerodontha angulata
Cerodontha eucaricis
Cerodontha ireos
Cerodontha pseuderrans
Cerodontha scirpi
Cerodontha spinata
Cerodontha vignae
Phytomyza abdominalis
Phytomyza crassiseta
Phytomyza crassiseta
Phytomyza ilicis
Phytomyza plantaginis
Phytomyza primulae
Anisopodidae
Mycetobia pallipes
Anthomyiidae
Achaetella varipes
Cordilura ciliata
Cordilura ontario
Fucellia marina
Fucellia merina
Homalomya sp
Hydrophoria conica
Hylemya antiqua
Hylemya antiqua
Hylemya brassicae
Hylemya brassicae
Hylemya cana
Hylemya cana
Hylemya cilicrura
Hylemya cilicrura
Hylemya crucifera
Hylemya crucifera
Hylemya echinata
Hylemya floralis
Hylemya floralis
Hylemya florilegea
Hylemya fugax
Hylemya fugax
Hylemya planipalpus
Hylemya planipalpus
Hylemya trichodactula
Hylemya trichodactyla
Ophyra leucostoma
Orthochaeta hirtipes
Pegomya betae
Pegomya bicolor
Pegomya geniculata
Pegomyia geniculata
Sexual
System
parth
Ploidy
3
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
22
14
18
22
22
40
38
21
21
14
18
21
19
38
37
XXY
XXY
XXY
XY
XXY
XXY
XXXY
XXY
252
252
252
252
252
252
252
252
12
18
20
11
18
20
XXY
XY
XXY
252
255
252
6
6
6
12
12
12
12
12
12
10
12
18
12
6
6
6
12
12
12
12
12
12
10
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
12
XY
12
12
XY
259
259
259
259
259
259
259
259
259
259
259
259
259
260
259
6
6
XY
261
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
14
14
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
13
13
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XXY
XXY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
262
262
262
263
262
263
262
264
262
264
262
264
262
264
262
264
262
262
264
262
262
265
264
264
262
262
264
263
262
262
262
262
264
XY
parth
192
Taxa
Phorbia brassica
Phorbia brassica
Scatophaga pallida
Scatophaga stercoraria
Asilidae
Asilus lecythus
Asilus notatus
Asilus sericeus
homomorphic
Dasyllis grossa
Dasyllis thoracica
Deromyia winthemi
Erax rufibarbis
Leptogaster badius
Septogaster badius
Bibionidae
Bibio bortulanus
Bibio hortulanus
homomorphic
Penthetria holosericea
Bombyliidae
Acrophthalrnyda paulseni
homomorphic
Anthrax anthrax
Anthrax cordillerensis
Anthrax lateralis
homomorphic
Anthrax mystaceus
Anthrax oedipus
Anthrax sinuosa
Anthrax tigrinus
Anthrax yamashiroensis
Bombylius fulvescens
Bombylius pygmaeus
Exoprosopa atrinasis
Exoprosopa sp.
Ligyra cerberus
Ligyra proserpina
Neodiplocampta roederi
Spogostylum simson
Systoechus vulgaris
Thyridanthrax abrupta
Thyridanthrax niveifrons
Thyridanthrax sp.
Thyridanthrax sp.
Toxophora javana
Usia florea
Villa cingulum
Villa gayi
Villa gorgon
Villa lateralis
Villa morio
Villa scylla
Villa sinuosa
Villa sp
Villa sp
Villa sp.1
Villa sp.1
Villa sp.2
Villa sp.2
Villa sp.3
Villa sp.3
Villa sp.lateralis grp.
Villa sp.nr.blanchardiana
Villa spB
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
XY
XY
XY
XY
262
266
262
262
14
14
10
14
14
10
XY
XY
XY
263
263
263
10
10
12
10
10
10
9
10
12
10
10
10
XO
XY
XY
XY
XY
XY
263
263
263
263
263
266
10
10
10
10
10
10
12
12
16
12
12
16
12
12
14
18
18
12
10
12
12
18
14
18
18
18
12
14
16
18
18
18
8
10
16
18
14
12
18
10
18
18
18
14
14
14
14
18
18
14
12
12
14
18
18
12
10
12
12
18
14
18
18
18
12
14
16
18
18
18
8
10
16
18
14
12
18
10
18
18
18
14
14
14
14
18
18
14
12
12
193
266
261
XY
261
267
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
267
267
263
267
267
263
267
267
267
267
267
267
267
267
267
263
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
267
Taxa
Villa spC
Villa velutina
Villa velutina
Villa velutina I
Villa velutina II
Calliphoridae
Albuquerquea latifrons
Amenia chrysame
Anthracomyza atratula
Bengalia inermis
Bengalia lyneborgi
Blepharicnema splendens
Boreellus atriceps
Bufolucilia silvarum
Calliphora viridescens
Calliphora vomitoria
Calliphora accepta
Calliphora bomitoria
Calliphora croceipalpis I
Calliphora croceipalpis II
Calliphora erythrocephala
Calliphora erythrocephala
Calliphora fallax
Calliphora fulviceps
Calliphora fulvicoxa
Calliphora hilli
Calliphora lilaea
Calliphora peruviana
Calliphora plebeia
Calliphora sternalis
Calliphora stygia
Calliphora tibialis
Calliphora varifrons
Calliphora vicina
Calliphora vicina
Calliphora vicina I
Calliphora vicina II
Calliphora viridenscens
Calliphora viridescens
Calliphora vomitoria
Calliphora vomitoria
Calliphora vomitoria I
Calliphora vomitoria II
Calliphora vomitoria III
Callitroga hominivorax
Callitroga hominivorax
Callitroga macellaria
Callitroga mocellaria
Chrysomia albiceps
Chrysomia cholorpyga
Chrysomia putoria
Chrysomya albiceps
homomoprhic; monogeny
Chrysomya marginalis
Chrysomya megacephala
Chrysomya phaonis
Chrysomya pinguis
Chrysomya putoria
Chrysomya rufifacies
homomoprhic; monogeny
Chrysomya saffraneta
Chrysomya varipes
Chrysomyia albiceps
Chrysomyia chloropyga I
Chrysomyia chloropyga II
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
14
18
18
18
18
14
18
18
18
18
XY
XY
XY
XY
XY
267
267
267
267
267
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
268
268
268
268
268
268
268
268
262
262
268
266
268
268
269
262
268
268
268
268
268
268
268
268
268
268
268
269
262
268
268
270
262
269
262
268
268
268
269
262
269
262
262
262
262
271
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
271
271
271
271
271
271
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
271
271
268
268
268
194
Taxa
Chrysomyia megacephala
Chrysomyia megacephala
Chrysomyia pachymera
Chrysomyia putoria I
Chrysomyia putoria II
Chrysomyia rufifacies
Chrysomyia rufifacies
Cochliomyia hominivorax
Cochliomyia macellaria I
Cochliomyia macellaria II
Cosmina testaceipes
Cynomya cadaverina
Cynomya mortuorum I
Cynomya mortuorum II
Cynomya mortuorum III
Cynomyia mortyorum
Cynomyopsis cadaverina
Cynomyopsis cadaverina
Discritomyia fasciata
Discritomyia fulgens
Discritomyia new species
Discritomyia terryi
Eucalliphora lilaea
Eucalliphora lilaea
Euphumosia setigera
Eurhyncomyia anterotes
Hemilucilia sp
homomorphic
Hemilucilia sp
homomorphic
Hemilucilia sp
homomorphic
Hemipyrellia brunneipes
Hemipyrellia fernandica
Hemipyrellia ligurriens
Hemipyrellia taeniops
Hemipyrellia tagaliana
Huascaromusca nigrifrons
Isomyia connivens
Isomyia prasina
Lucilia caesar
Lucilia caesar
Lucilia cuprina
Lucilia illustria
Lucilia sericata
Lucilia ampullacea
Lucilia caesar
Lucilia cuprina
Lucilia illustris
Lucilia porphyrina
Lucilia sericata
Melinda biseta
Melinda sp
Mesembrinella peregrina I
Mesembrinella peregrina II
Metallea cuprea
Metallea puncticeps
Microcalliphora varipes
Paralucilia fulvicrura
Paralucilia wheeleri
Paratricyclea surcoufi
Phaenicia caeruleiviridis
Phaenicia eximia
Phaenicia sericata
Phaenicia caeruleiviridis
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
268
262
268
268
268
268
262
268
268
268
268
268
268
268
268
262
269
262
268
268
268
268
270
262
268
268
268
12
12
XY
268
12
12
XY
268
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XXYY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
268
268
268
268
268
268
268
268
269
262
262
262
262
268
268
271
269
268
271
268
268
268
268
268
268
268
268
268
268
262
262
262
268
195
Taxa
Sexual
System
Phaenicia coernleiviridis
Phaenicia cuprina
Phaenicia eximia
Phaenicia eximia
Phaenicia sericata
Phormia regina
Phormia terrae-novae
Phormia regina
Phormia regina
Phormia terraenovae
Phumosia promittens
Pollenia rudis
Pollenia nigripes
Pollenia rudis
Pollenia rudis I
Pollenia rudis II
Pollenia rudis III
Polleniopsis hokurikuensis
Protocalliphora hirundo
Protocalliphora sialia
Protocalliphora sp
Protocalliphora aenea
Protocalliphora aenea
Protocalliphora avium
Protocalliphora avium
Protocalliphora hirundo
Protocalliphora hirundo
Protocalliphora metallica
Protocalliphora metallica
Protocalliphora sialia
Protocalliphora sialia
Protophormia terraenovae
Protophormia terraenovae
Protophormia terraenovae
Protophormia terraenovae I
Protophormia terraenovae II
Rhinia apicalis
Rhinia discolor
Rhinia subapicalis
Rhyncomyia italica
Rhyncomyia stannocuprea
Sarconesiopsis chilensis
Sphenometrops tergata
Stomorhina armatipes
Stomorhina lunata
Stomorhina obsoleta
Cecidomyiidae
Asphondylia monacha
Asteromyia rubra
Caryomyia sp
Cecidomyia serotinae
Cecidomyia sp.
Cecidomyia vicola
Dasyneura affinis
Lasioptera asterspinosae
Lestodiplosis sp
Miastor americana
octoploid in germ line, paedogenetic
Miastor metraloas
Miastor sp
Monarthropalpus buxi
Oligarces paradoxus
Oligotrphus pattersoni
Ontarinia canadensis
Phytophaga celtiphyllia
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
270
268
269
268
269
262
262
269
268
269
268
262
268
269
268
268
268
268
268
268
268
269
262
269
262
270
262
269
262
270
262
269
262
271
268
268
268
268
268
268
268
268
262
268
268
268
8
8
8
8
8
8
8
8
8
12
7
7
7
7
7
7
7
7
7
11
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
266
266
266
266
266
266
266
266
266
266
XO
XO
XO
XO
XO
XO
XO
266
266
266
266
266
266
266
8
7
8
8
12
7
7
196
Taxa
Phytophaga destructor
Rhopalomyia sabinae
Taxomyia taxi
Trishormomyia helianthi
Walshomyia texana
Chamaemyiidae
Cremifania nigrocellulata
homomorphic
Cremifania nigrocellulata
Leucopis obscura
Leucopmyia obscura
Chironomidae
Chironimus plumosus
homomorphic
Chironomus alluaudi
homomorphic
Chironomus alpestris
Chironomus bathophilus
Chironomus confinis
Chironomus dorsalis
Chironomus formosipennis
homomorphic
Chironomus plumosus
Chironomus pulcher
homomorphic
Chironomus riparius
homomorphic
Chironomus riperiua
Chironomus sp
homomorphic
Chironomus sp.
Chironomus sp.
homomorphic
Chironomus sp.
homomorphic
Chironomus thummi
Chironomus transvaalensis
homomorphic
Corynoneura sp nr scutellata
Cryptochironomus defectus
Endochironomus sp
Glyptotendipes barbipes
homomorphic
Glyptotendipes polytomus
Glyptotendipes sp
Glyptotendipes sp
Limnophyes vestitus
Lundstroemia parthenogenetica
Micropsectra praecox
Microtendipes pedellus
Sergentia profundorum
Stictochironomus histrio
Culicidae
Aedes albopictus
homomorphic
Aedes canadensis
homomorphic
Aedes japonicus
homomorphic
Aedes koreicus
homomorphic
Aedes togoi
homomorphic
Aedes triseriatus
homomorphic
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
9
8
8
8
8
8
7
7
7
7
XO
XO
XO
XO
XO
266
266
266
266
266
6
6
XY
272
6
10
10
6
10
10
XY
262
272
262
8
8
261
8
8
273
8
8
8
8
8
8
8
8
8
8
266
266
266
266
273
8
8
8
8
266
273
8
8
273
8
8
8
8
266
273
8
6
8
6
266
273
8
8
273
8
8
8
8
266
273
6
8
8
6
8
8
274
266
266
261
8
6
8
8
6
8
8
8
6
8
8
8
6
8
266
266
266
274
274
266
266
266
266
6
6
266
6
6
266
6
6
266
6
6
266
6
6
266
6
6
266
parth
parth
parth
197
Taxa
Anopheles agrgyritarsus
Anopheles albimanus
Anopheles aquasalis
Anopheles aztecus
Anopheles claviger
Anopheles darlingi
Anopheles freeborni
Anopheles gambiae
Anopheles maculipennis
Anopheles noroestensis
Anopheles occidentalis
Anopheles pseudopunctipennis
Anopheles punctipennis
Anopheles punctipennis
Anopheles quadrimaculatus
Anopheles sp
Anopheles stephensi
Anopheles strodei
Anopheles strodei
Armigeres obturbans
Chaoborus plumicornis
Corethra plumicornis
Culex hayashii
homomorphic
Culex pipiens
homomorphic
Culex pipiens
Culex spicalis
homomorphic
Culex tarsalis
homomorphic
Culex territans
homomorphic
Culex tritaeniorhynchus
homomorphic
Lutzia fuscana
homomorphic
Mochlonyx sp
Theobaldia incidens
homomorphic
Cuterebridae
Cuterebra emasculator
Cuterebra emasculator
Cylindrotomidae
Phalacrocera replicata
Diopsidae
Sphyracephala brevicornis
Dryomyzidae
Dryomyza anilis
Neuroctena analis
Helemyzidae
Suillia nemorum
Suillia nemorum
Suillia sp
Hippoboscidae
Melophagus ovinus
Olfersia bisulcata
Hypodermatidae
Hypoderma bovis
Hypoderma lineatum
Lauxaniidae
Calliopum aeneum
Depressa striatipennis
Homoneura armata
Homoneura biroi
Sexual
System
Ploidy
6
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
8
6
6
6
6
6
6
6
6
6
6
6
6
6
XY
6
6
6
6
6
6
6
6
8
6
6
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
275
XY
XY
XY
XY
XY
XY
XY
6
6
266
12
6
12
6
263
266
6
6
266
6
6
266
6
6
266
6
6
266
8
6
8
6
XY
266
266
12
12
12
12
XY
XY
262
276
10
10
XY
261
10
10
XY
262
12
12
12
12
XY
XY
262
262
12
12
12
12
12
12
XY
XY
XY
262
262
262
18
8
18
8
XY
XY
266
266
12
12
12
12
XY
XY
276
276
12
12
12
12
12
12
12
12
XY
XY
XY
XY
272
272
272
272
198
275
275
275
275
275
275
275
275
275
275
275
275
266
275
266
275
275
275
266
266
266
266
Taxa
Homoneura horvathi
Homoneura laticosta
Homoneura sp1
Homoneura sp2
Homoneura spA
Homoneura spB
Lauxania cylindricornis
Lyciella bicoloripes
Lyciella hirtiventris
Lyciella illota
Lyciella rorida
Lyciella sp1
Lyciella sp2
Lyciella spinigera
Minettia flaveola
Minettia flaveola
Minettia lupulina
Minettia lupulina
Paralauxania elevata
Paranomina sp
Physegenua vittata
Physegenua vittata
Physegenua vittata
Sapromyza avicola
homomorphic
Sapromyza sp
Sapromyza stigmatica
Sp sp
Sp sp
Sp sp
Sp sp
Sp. sp.
Leptoceratidae
Leptocera fontinalis
Limoniidae
Dicranomyia trinotata
homomorphic
Thaumastoptera calceata
homomorphic
ltonididae
Phytophaga destructor
homomorphic
Muscidae
Calliphora erythrocephala
Fannia canicularis
Fannia canicularis
Fannia glaucescens
Fannia glaucescens
Fannia sp
Fannia sp.
Fucellia marina
Haematobia irritans
Haematobia irritans
homomorphic
Homalomyia sp
Hydrophoria conica
homomorphic
Hydrotaea houghi
Hydrotaea scambus
homomorphic
Hydrotaea scambus
Macrorchus ausoba
Musca autymnalis
Musca domestica
Musca sorbens
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
10
10
12
12
12
10
10
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
10
10
12
12
12
10
10
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
272
272
272
272
272
272
272
272
272
272
272
272
272
272
272
262
272
272
272
272
272
272
262
272
8
12
12
12
12
12
12
8
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
272
272
272
272
272
272
262
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
7
266
6
6
261
6
6
261
8
8
261
12
12
12
12
12
12
12
12
10
10
12
12
12
12
12
12
12
10
10
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
263
270
262
270
262
270
270
270
262
270
12
12
12
12
XY
XY
262
270
10
10
XY
XY
270
270
10
12
12
12
12
10
12
12
12
12
XY
XY
XY
262
270
262
262
262
199
Taxa
Musca vestustissima
Musca autumnalis
Musca autumnalis
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica
Musca domestica curviforceps
Musca domestica curviforceps
Musca domestica domestica
Musca domestica domestica
Musca domestica domestica
Musca domestica domestica
Musca domesticacalleva
Musca domesticacalleva
Musca domesticacurviforceps
Musca sorbens
Musca sorbens
Musca veltustissima
heteromorphic
Musca velustissima
Muscina stabulans
Muscina stabulans
homomorphic
Muscina stabulans
Muscina stabulans
homomorphic
Mydaea neglecta
Ophyra leucostoma
homomorphic
Ophyra leucostoma
Ophyra leucostoma
Ophyra leucostoma
Orthellia nudissima
Orthellia nudissima
homomorphic
Orthellia nudissima
homomorphic
Pegomya betae
Pegomya bicolor
homomorphic
Pegomya geniculata
Peregle radicum
homomorphic
Phaonia basalis
Phaonia variegata
Phaonia basalis
Phaonia basalis
Phaonia variegala
homomorphic
Phaonia variegata
homomorphic
Phormia regina
Pseudopyrellia cornicina
Stomoxya calicitrans
Stomoxys calcitrans
homomorphic
Sexual
System
Ploidy
12
11
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
11
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
270
270
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
12
10
10
12
10
10
XY
XY
XY
270
262
270
12
10
12
10
XY
270
277
12
12
12
XY
XY
270
270
12
12
12
10
10
12
12
12
10
10
XY
XY
XY
XY
XY
270
277
262
262
270
10
10
12
12
12
12
XY
XY
270
270
12
12
12
12
XY
XY
270
270
12
10
12
12
10
12
10
12
12
10
XY
XY
XY
XY
XY
262
262
270
277
270
10
10
12
12
10
10
12
12
10
10
200
262
270
277
270
270
270
270
270
270
270
277
277
277
266
263
270
270
270
270
270
277
277
270
277
277
277
277
XY
XY
XY
263
270
262
270
Taxa
Mycetophilidae
Brachypeza radiata
Exechia indecisa
Exechia indecisa
Exechia speciosa
Fungivora blanda
Fungivora blanda
Fungivora funorum
Fungivora guttata
Fungivora guttata
Fungivora lineola
Fungivora unipunctata
Mycetophila punctata
Mycetophila punctata
Rhymosia domestica
Rhymosia fenestralis
Oestridae
Cephenemyia phobifer
Cephenomyia phobifer
Hypoderma bovis
Hypoderma lineatum
Oestrus ovis
Oestrus ovis
Ortalidae
Camptoneura picta
Camtpneura picta
Chaetopsis fulvifrons
Otitidae
Camptonoura picta
Ceroxys latiusculus
Ceroxys latiusculus
Ceroxys urticae
Chaetopsis fulvifrons
Chaetopsis fulvifrons
Chaetopsis ulvi
Chaetopsis ulvi
Delphinia picta
Delphinia picta
Delphinia picta
Euxesta notata
Euxesta notata
Euxesta notata sp.1
Euxesta notata sp.2
Melieria crassipennis
Melieria crassipennis I
Melieria crassipennis II
Myrmecothea myrmecoides
Myrmecotheca myrmecoides
Physiphora aenea
Pterocalla sp.
Seioptera vibrans
Seioptera vibrans
Phoridae
Aphiocaeta sp
Phora sp.
Phryneidae
Phryne fenestralis
Piophilidae
Piophila casei
Platystomatidae
Amphicnephes sp.
Laglasia sp.
Lenophila caerulea
Plagiostenopterina sp. 1
Plagiostenopterina sp. 2
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
14
14
12
8
8
14
8
8
14
14
14
14
14
8
12
14
14
12
8
8
14
8
8
14
14
14
14
14
8
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
266
266
261
266
266
261
266
266
261
266
266
266
261
266
266
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
262
276
262
262
262
276
12
12
8
12
12
8
12
12
12
12
8
8
8
8
12
12
12
12
12
12
12
8
8
8
10
12
10
10
6
6
12
12
12
12
8
8
8
8
12
12
12
12
12
12
12
8
8
8
10
12
10
10
6
6
6
8
6
8
8
8
12
12
6
12
12
6
6
6
12
12
6
6
201
263
266
263
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
262
262
278
278
278
262
263
263
278
263
263
262
278
278
278
262
278
278
262
278
278
278
262
278
266
266
XY
261
266
XY
XY
XY
XY
XY
278
278
278
278
278
Taxa
Platystoma lugubre
Pogonortalis doclea
Rivellia viridulans
Rivellia sp.
Rivellia sp. Probably viridulans
Psilidae
Chamaepsila rosae
Psychodidae
Lutzomyia carmelinoi
homomorphic
Lutzomyia columbiana
homomorphic
Lutzomyia erwindonaldoi
homomorphic
Lutzomyia gomezi
homomorphic
Lutzomyia trapidoi
homomorphic
Lutzomyia walkeri
homomorphic
Ptychopteridae
Liriope cinerasceus
Liriope sp
Rhagionidae
Chrysophilus sp
Leptis sp
Leptis sp.
Richardiidae
Setellia pernix
possibly XO
Sapromyzidae
Physegenua vittata
Physegenua vlittata
Sarcophagidae
Acridiophaga aculeata
heteromorphic
Acridiophaga aculeata
Arachnidiomya aldrichi
Blaesoxipha hunteri
Blaesoxipha hunteri
Blaesoxipha hunteri
Blaesoxipha opifera
Blaesoxipha opifera
Boettcheria cimbicis
Boettcheria cimbicis
Boettcheria cimbicis
Euboettcheria sp
Euboettcheria sp
Euboettcheria sp.
Helicobia rapax
Helicobia rapax
Helicobia rapax
Helicobia sp
Helicobia sp
Helicobia sp.
Hystricocnema plinthopyga
Hystricocnema plinthopyga
Hystricocnoma plinthopyga
Kellymyia kellyi
heteromorphic
Kellymyia kellyi
Kellymyia kellyi
Neobellaria bullata
Neobellieria bullata
Neobellieria cooleyi
Sexual
System
Ploidy
6
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
10
6
6
6
12
10
6
6
XY
XY
XY
XY
XY
278
8
8
262
8
8
279
8
8
279
8
8
279
8
8
279
6
6
279
8
8
279
10
10
10
10
XY
XY
266
266
10
10
10
10
10
10
XY
XY
XY
280
266
280
10
10
XY
278
12
12
12
12
XY
266
263
12
12
XY
281
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
262
270
270
270
262
270
262
282
270
262
270
262
282
282
270
262
270
262
282
282
270
262
281
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
270
262
262
282
270
202
12
278
278
262
278
Taxa
Paraphrissopoda chrysostoma
Paraphrissopoda chrysostoma
Paraphrisspdda chrysostoma
Parasarcophaga argyrostoma
Protodexia australis
Protodexia hunteri
Protodexis australis
Protodexis hunteri
Pseudosarcophaga affinis
Pseudosarcophaga affinis
Pseudosarcophaga affinis
homomorphic
Ravinia communis
Ravinia peniculata
Ravinia duplicata
Ravinia l'herminieri
Ravinia peniculata
Ravinia pusiola
Ravinia querula
Ravinia querula
Rawinia peniculata
Sacrophaga carnria
Sacrophaga sgrracinae
Sacrophaga sp
Sacrophaga tuberosa
Sarcophaga carnaria
Sarcophaga sp
Sarcophaga tuberosa
Sarcophaga aldrichi
Sarcophaga aldrichi
Sarcophaga argyrostoma
Sarcophaga argyrostoma
Sarcophaga argyrostoma
Sarcophaga argyrostoma
Sarcophaga bullata
Sarcophaga carnaria
Sarcophaga cooley
Sarcophaga cooleyi
Sarcophaga crassipalpus
Sarcophaga crassipalpus
Sarcophaga dalmatina
Sarcophaga exuberans
Sarcophaga exuberans
Sarcophaga exuberans
Sarcophaga falculata
Sarcophaga falculata
Sarcophaga falculata
Sarcophaga H
Sarcophaga H
Sarcophaga lalisterna
Sarcophaga latisterna
Sarcophaga occipitalis
Sarcophaga occipitalis
Sarcophaga occipitalis
Sarcophaga raversa
Sarcophaga reversa
Sarcophaga reversa
Sarcophaga securifera
Sarcophaga securifera
Sarcophaga serracaniae
Sarcophaga serraceniae
Sarcophaga serraceniae
Sarcophaga sp
Sarcophaga sp.
Sarcophage latisterna
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
12
18
20
18
XY
XY
12
12
12
12
12
12
18
20
18
282
262
XY
XY
XY
XY
XY
XY
XY
270
282
281
281
262
262
262
281
270
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
262
262
270
270
266
270
270
262
263
266
266
266
266
262
263
262
281
262
282
270
270
262
270
270
262
281
270
262
270
282
270
262
282
270
262
281
270
270
262
282
270
262
262
281
262
282
262
262
270
270
262
270
262
203
12
12
12
12
12
12
12
12
12
12
12
12
Taxa
Servasia aculeata
Servasia australis
Servasia reversa
Sphenometopa tergata
Sphenometopa tergata
Wohlfahrlia meigeni
Wohlfahrlia opaca
Wohlfarhtia meigeni
Wohlfarhtia opaca
Wolfahrtia meigeni
Wolfahrtia opaca
Scathophagidae
Cordilura ontario
Cordilura praensia
Cordilura varipes
Neochirosia nuda
Orthochaeta hirtipes
Scaptophaga furcala
Scaptophaga nigrolimbata
Scaptophaga pallida
Scaptophaga stercoraria
Scathophaga pallida
Scathophaga stercoraria
Scatopsidae
Scatopse notata
Scatopse sp
Sciaridae
Sciara agarica
Sciara coprphila
Sciara fenestralis
Sciara nacta
Sciara ocellaris
Sciara pauciseta
Sciara prolifica
Sciara reynoldsi
Sciara similans
Sciara sp
Sciomyzidae
Antichaeta melanosoma
Antichaeta melanosoma
Atrichomelina pubera
Atrichomelina pubera
Colobaea americana
Dichetophora boyesi
Dichrochirosa dissimilis
Dichrochirosa sp
Dictya sabroski
Dictya texensis
Dictya txensis
Dictya atlantica
Dictya atlantica
Dictya brimleyi
Dictya brimleyiE
Dictya floridensis
Dictya sabroskyi
Dictya stricta
Dictya texensis
Dictya umbrarum
Dictya umbrarum series II
Dictyodes dictyodes
Elgiva cucularia
Elgiva divisa
Elgiva rufa
Elgiva rufa
Euthycera chaerophylli
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
270
270
270
270
262
282
282
270
270
262
262
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
270
270
270
270
270
270
270
270
270
266
266
12
12
12
12
XY
XY
261
261
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
266
266
266
266
266
266
266
266
266
266
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
12
XY
262
283
262
283
283
284
283
283
262
262
262
283
262
283
262
283
283
283
283
284
284
283
284
284
284
283
283
8
8
8
8
8
8
8
8
8
8
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
204
7
7
7
Taxa
Sexual
System
Helosciomyza ferruginea
Hoplodictya spinicornis
Hydromya dorsalis
Hydromya dorsalis series II
Knutsonia lineata
Limnia boscii
Limnia paludicola
Limnia unguicornis
Neuroctena analis
Neuroctena analis
Perilimnia albifacies
Pherbellia dubia
Pherbellia grisescens
Pherbellia nana
Pherbellia sp.
Pherbellia albocostata
Pherbellia brunnipes
Pherbellia chiloensis
Pherbellia grisescens
Pherbellia humilis
Pherbellia javana
Pherbellia lapponica
Pherbellia nana
Pherbellia obscura
Pherbellia obtusa
Pherbellia patagonensis
Pherbellia scutellaris
Pherbellia sp. 1 (n.sp.near patagonensis)
Pherbellia sp.4 (n.sp.near chiloensis)
Pherbellia trabeculata
Pherbina corlleti
Pherbina coryleti
Protodictya chilensis
Protodictya guttularis
Psacadina zerni
Psacadina zernyi
Pteromicra glabricula
Pteromicra pectorosa
Pteromicra similis
Renocera fuscinervis
Salticella fasciata
Sepedon fuscipennis
Sepedon armipes
homomorphic
Sepedon armipes
Sepedon fuscipennis
Sepedon macropus
Sepedon plumbella
Sepedon saegeri
Sepedon sphegea
Sepedon spinipes americana
Tetanocera loewiE
Tetanocera sp.
Tetanocera sparsa
Tetanocera arrogans
Tetanocera elata
Tetanocera ferruginea
Tetanocera ferruginea
Tetanocera hyalipennis
Tetanocera loewi
Tetanocera montana
Tetanocera plebeia
Tetanocera plumosa
Tetanocera silvatica
Tetanocera unicolor
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
10
10
10
12
12
12
12
12
11
12
12
12
12
12
XY
XO
XY
XY
XY
XY
XY
XY
12
12
12
12
12
12
12
12
12
12
10
12
12
12
12
12
12
12
12
12
12
205
12
12
12
12
11
12
12
12
12
12
12
11
12
12
12
12
12
11
12
12
12
12
12
12
12
12
12
12
10
9
XY
XY
XO
XY
XY
XY
XY
XY
XY
XO
XY
XY
XY
XY
XY
XO
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XO
12
12
12
11
XY
XY
XY
XO
11
12
12
12
12
12
11
12
12
12
10
12
12
12
12
12
12
12
12
XO
XY
XY
XY
XY
XY
XXY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
12
XY
284
283
284
284
284
283
284
283
266
263
283
284
262
262
262
284
284
283
283
283
284
284
283
284
283
283
284
283
283
283
283
284
283
283
262
283
284
283
283
284
284
262
262
283
283
283
284
284
283
283
262
262
262
284
283
284
283
284
283
283
283
283
283
284
Taxa
Tetanocera vicina
Tetanoceroides mesopleuralis
Tetanoceroides occidentalis
Trypetoptera punctulata
Sepsidae
Piophila casei
Simuliidae
Eusimulium vernum
Simulium sp
Stratiomyidae
Actina incisurali
Adoxomyia rustica
Beris chalybeata
Chloromyia formosa
Hermetia illucens
Himantoloba flavopilosa
Lasionemopoda hirsuta
Merosargus melanothorax
Neoexaireta spinigera
Odontomyia regisgeorgei
Ptecticus australis
Ptecticus longipennis
Ptecticus matsumurae
Ptecticus n. sp 5
Ptecticus n.sp 1
Ptecticus n.sp 2
Ptecticus n.sp 3
Ptecticus n.sp 4
Ptecticus sackenii
Ptecticus tenebrifer
Ptecticus trivittatus
Ptecticus trivittatus
Ptecticus trivittatus
Ptecticus trivittatus
Ptecticus wulpii
Sargus cuprarius
Sargus fasciatus
Straliomys barbata
Stratiomys sp
Syrphidae
Blera badia
Blera fallax
Blera scitula
Allograpta exotica
Allograpta hortensis
Allograpta neotropica
Allograpta obliqua
Allograpta pfeifferi
Allograpta piurana
Allograpta piurana
Allograpta pulchra (hybrid?)
Allograpta pulchra (I)
Asarcina rostrata
Baccha alicia
Baccha asthenia
Baccha clavata (I)
Baccha clavata (II)
Baccha debasa
Baccha flavipennis
Baccha funebris
Baccha gastrostacta
Baccha livida
Baccha melanorrhina
Baccha norina
Baccha nymphaea
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
XY
XY
12
XY
283
283
283
284
12
12
XY
263
6
6
6
6
XY
285
266
10
8
10
14
14
16
16
16
12
12
16
16
16
16
16
10
8
10
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
280
280
280
280
280
280
280
280
280
280
280
280
280
280
280
280
280
280
263
280
280
266
263
263
280
280
280
280
280
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
286
286
286
287
287
287
288
288
287
287
287
287
288
287
287
287
287
287
287
287
287
287
287
287
287
16
16
16
16
16
16
16
16
16
16
8
14
12
14
16
16
16
12
12
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
16
8
14
12
12
12
12
10
8
10
10
12
10
10
8
10
12
10
10
10
10
10
10
10
10
8
10
8
10
12
12
12
10
8
10
10
12
10
10
8
10
12
10
10
10
10
10
10
10
10
8
10
8
10
206
XY
XY
XY
XY
XY
XY
XY
Taxa
Baccha polista
Baccha scutellata
Baccha virginio
Baccha vittiger
Baccha zilla
Baccha fascipennis
Baccha obscuripennis
Blera analis
Blera badia
Blera confusa
Blera confusa
Blera nigra
Blera nigra
Caliprobola crawfordi
Calliprobola crawfordi
Calliprobola crawfordi
Carposcalis obscura
Carposcalis sp.
Carposcalis sp.(3)
Carposcalis sp.(4)
Carposcalis sp.(5)
Carposcalis sp.(6)
Carposcalis sp.(8)
Cheilosia pagana
Cheilosia plutonia
Cheilosia tristis
Cheilosia vernalis
Chrysogaster metallina
Chrysogaster nitida
Chrysogaster solstitialis
Chrysogaster splendita
Chrysogaster viduata
Chrysotoxum arcuatum
Chrysotoxum arcuatum
Chrysotoxum bicinctum
Chrysotoxum bicinctum
Chrysotoxum cautum
Chrysotoxum derivatum
Chrysotoxum derivatum
Chrysotoxum elegans
Chrysotoxum fasciolatum
Chrysotoxum festivum
Chrysotoxum festivum
Chrysotoxum intermedium
Chrysotoxum latifasciatum
Chrysotoxum lessonae
Chrysotoxum octomaculatum
Chrysotoxum parmense
Chrysotoxum ventricosum
Chrysotoxum verralli
Cnemodon latitarsis
Criorhina berberina
Criorhina berberina
Criorhina berberina
Criorhina caudata
Criorhina coquilIetti
Criorhina luna
Criorhina pachymera
Criorhina tricolor
Criorhina tricolor
Didea laxa
Doros conopseus
Epistrophe bifasciata
Epistrophe bifasciata
Episyrphus balteatus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
10
10
10
10
6
6
12
12
12
12
12
12
10
10
10
8
8
8
8
8
8
8
12
12
10
10
12
12
12
12
12
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
8
8
8
8
12
12
12
8
12
12
10
8
10
10
10
10
9
10
10
10
6
6
12
12
12
12
12
12
10
10
10
8
8
8
8
8
8
8
12
12
10
10
12
12
12
12
12
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
8
8
8
8
12
12
12
8
12
12
10
8
10
10
10
XY
XXY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
207
287
287
287
287
287
288
288
286
288
288
286
288
286
288
286
286
288
287
287
287
287
287
287
288
288
288
288
288
288
288
288
288
288
289
288
289
289
288
289
289
289
288
289
289
289
289
289
289
289
289
290
286
286
286
286
286
286
286
286
286
288
288
290
288
288
Taxa
Eristalinus sepulcralis
Eristalis abusivus
Eristalis aeneus
Eristalis anthophorinus
Eristalis arbustorum
Eristalis bastardi
Eristalis bastardi
Eristalis bastardi
Eristalis dimidiatus
Eristalis horticola
Eristalis intricarius
Eristalis lenax
Eristalis meigeni
Eristalis nemorum
Eristalis obscurus
Eristalis pertinax
Eristalis pratorum
Eristalis tenax
Eristalis tenax
Eumerus obliquus
Eumerus strigatus
Eumerus tuberculatus
Hadromyia grandis
Helophilus fasciatus
Helophilus hybridus
Helophilus pendulus
Helophilus trivittatus
Ischiodon aegyptium
Ischiodon aegyptium
Ischyrosyrphus velutinus
Lejops lineatus
Lejops lunulatus
Lejops obsoletus
Lejops relictus
Lejops rex
Lejops stipatus
Lejops transfugus
Lejops versicolor
Lejota cyanea
Lejota cyanea
Leucozona lucorum
Mallota posticata
Melangyna cincta
Melangyna lasiophthalma
Melangyna pullulus
Melanostoma mellinum
Melanostoma pictipes
Melanostoma scalare
Meliscaeva cinctella
Meliscaeva sp
Merodon equestris
Mesogramma marginata
Mesogramma marginata
Mesograpta marginata
Metasyrphus americanus
Metasyrphus astutus
Metasyrphus canadensis
Metasyrphus corollae
Metasyrphus luniger
Metasyrphus perplexus
Metasyrphus snowi
Microdon piperi
Mitesia virginiensis
Myathropa florea
Myolepta luteola
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
10
12
12
12
12
8
10
8
12
10
14
10
12
12
12
12
12
12
10
12
10
10
10
8
8
8
10
10
12
12
12
12
8
8
8
8
8
8
8
12
12
10
10
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
10
12
12
12
12
8
10
8
12
10
14
10
12
12
12
12
12
12
10
12
10
10
10
8
8
8
10
10
12
12
12
12
8
8
8
8
8
8
8
12
12
10
10
XY
XY
XY
XY
XY
XY
XY
208
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
288
288
288
288
288
290
288
263
288
288
288
290
288
288
288
288
288
288
263
288
288
288
286
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
286
288
288
288
288
288
288
288
288
288
288
288
290
263
288
288
288
288
288
288
288
288
288
286
288
288
Taxa
Myolepta luteola
Myolepta luteola
Myolepta pretiosa
Myolepta strigilata
Neoascia globosa
Neoascia podagrica
Neocnemodon latitarsus
Neocnemodon vitripennis
Ornidia obesa
Orphnabaccha sp.
Paragopsis strigalus
Paragus bicolor
Paragus borbonicus
Paragus longiventris
Paragus minutus
Paragus tibialis
Phalacrodira rectoides
Phalacrodira vittiger
Pipiza femoralis
Pipiza nigripilosa
Pipiza sp.
Pipizella varipes
Pipizella virens
Platycheirus albimanus
Platycheirus angustatus
Platycheirus clypeatus
Platycheirus fulviventris
Platycheirus peltatus
Platycheirus quadratus
Platycheirus scambus
Platycheirus scutatus
Pseudoscaeva sericea
Rhingia campestris
Rhingia nasica
Rhysops currani
Rhysops neotropica
Rhysops nigrans
Rhysops sp. (13)
Salpingogaster nigra
Scaeva melanostoma
Scaeva occidentalis
Scaeva pyrastri
Scaeva selenitica
Sericomyia chalcopyga
Sericomyia chrysotoxoides
Sericomyia lata
Sericomyia militaris
Sericomyia species 1
Somula decora
Somula decora
Sphaerophoia scripta
Sphaerophoria menthastri
Sphaerophoria robusta
Sphaerophoria rueppellii
Sphaerophoria scripta
Sphaerophoria Species 6
Sphaerophoria Species 7
Sphaerophoria sylindrica
Sphecomyia brevicornis
Sphecomyia occidentalis
Sphecomyia pattonii
Sphegina armatipes
Sphegina californiaca
Sphegina flavimana
Sphegina infuscata
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
10
10
10
8
8
8
8
12
10
12
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
10
10
10
10
14
8
8
8
10
10
8
8
12
10
12
12
10
12
10
8
8
8
8
8
8
8
8
14
10
12
12
12
12
12
10
10
10
10
8
8
8
8
12
10
12
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
8
10
10
10
10
14
8
8
8
10
10
8
8
12
10
12
12
10
12
10
8
8
8
8
8
8
8
8
14
10
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
209
286
286
286
286
288
288
288
288
288
287
290
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
288
287
288
288
287
287
287
287
287
287
287
288
288
291
291
291
291
291
288
286
290
288
288
288
288
288
288
288
286
286
286
288
288
288
288
Taxa
Spilomyia digitata
Spilomyia longicornis
Syritta pipiens
Syritta pipiens
Syrphus octomaculatus
Syrphus balteatus
Syrphus corollae
Syrphus lasiophthalmus
Syrphus luniger
Syrphus melanostomoides
Syrphus melanostomoides
Syrphus phaeostigma
Syrphus rectus
Syrphus reedi
Syrphus ribesii
Syrphus ribesii
Syrphus sp.
Syrphus torvus
Syrphus vitripennis
Syrphus vittafrons
Tcmnostoma alternans
homomorphic
Temnostoma acra
homomorphic
Temnostoma acra
homomorphic
Temnostoma alternans
homomorphic
Temnostoma balyras
Temnostoma balyras
homomorphic
Temnostoma vespiforme
Toxomerus anthrax
Toxomerus calceolatus
Toxomerus croesus
Toxomerus geminatus
Toxomerus lachrymosus
Toxomerus laciniosus
Toxomerus laenas
Toxomerus occidentalis
Toxomerus philippii
Toxomerus politus
Toxomerus portius
Toxomerus sp. (10)
Toxomerus sp. (11)
Toxomerus sp. (12)
Toxomerus sp. (15)
Toxomerus sp. (16)
Toxomerus sp. (19)
Toxomerus sp. (4)
Toxomerus sp. (6)
Toxomerus sp.(3)
Toxomerus steatogaster
Toxomerus vitreus
Tropidia quadrata
Tropidia quadrata
Tropidia scita
Tropidia scita
Tubifera sp
Tubifera sp.
Volucella bombylans
Volucella obesa
Volucella obesa
Volucella pellucens
Xanthogramma ornatum
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
10
10
10
8
10
8
10
8
10
10
8
8
8
8
10
8
8
8
8
8
10
10
10
10
8
10
8
10
8
10
10
8
8
8
8
10
8
8
8
8
8
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
286
286
288
286
287
290
290
290
290
290
288
287
288
287
288
288
288
288
288
288
286
12
12
XY
286
12
12
XY
286
8
8
XY
286
8
8
8
8
XY
288
286
8
10
10
10
10
12
10
10
10
10
10
10
10
10
10
10
8
10
10
12
10
10
10
10
10
10
10
14
14
12
12
12
12
10
8
10
10
10
10
12
10
10
10
10
10
11
10
10
10
10
8
10
10
12
10
10
10
10
10
10
10
14
14
12
12
12
12
10
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XYY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
286
287
287
287
288
287
287
287
288
287
287
287
287
287
287
287
287
287
287
287
287
287
287
288
286
288
286
290
288
288
290
263
288
288
210
Taxa
Xylota _subfasciata
Xylota angustiventris
Xylota annulifera
Xylota atlantica
Xylota atlantica
Xylota barbara
Xylota barbata
Xylota barbata_
Xylota bigelowi
Xylota bigelowi
Xylota chalybea
Xylota chalybea
Xylota chalybea
Xylota coeruleiventris
Xylota coeruleiventris
Xylota curvaria
Xylota curvaria
Xylota ejuncida
Xylota flavitibia
Xylota flavitibia
Xylota flexa
Xylota florum
Xylota hinei
Xylota ignava
Xylota inarmata
Xylota lenta
Xylota libo
homomorphic
Xylota nemorum
Xylota nemorum
Xylota nemorum
Xylota pigra
Xylota pigra
Xylota pigra
Xylota quadrimaculata
Xylota quadrimaculata
Xylota segnis
Xylota segnis
Xylota segnis
Xylota sp2
Xylota subfasciata
Xylota sylvarum
Xylota sylvarum
Xylota sylvarum
Xylota tuberculata
Xylota tuberculata
Xylota tuberculata
Xylota vecors
Tabanidae
Atylotus bicolor
Atylotus bicolor
Atylotus fulvus
Atylotus fulvus
Atylotus horvathi
Atylotus loewianus
Atylotus obioensis
Atylotus ohioensis
Atylotus pulchellus karybenthinus
Chrysops aberrans
Chrysops aberrans
Chrysops aestuans
Chrysops caecutiens
Chrysops flavipes
Chrysops frigidus
Chrysops frigidus
Sexual
System
Ploidy
18
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
10
10
10
10
10
10
10
8
8
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
8
8
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
286
286
286
288
286
286
288
286
288
286
288
286
286
286
286
286
286
286
288
286
286
286
286
286
286
286
286
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
10
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
288
286
286
288
286
286
288
286
288
286
286
286
288
288
286
286
288
286
286
286
18
18
18
18
12
18
18
18
18
12
12
10
10
10
10
10
18
18
18
18
12
18
18
18
XY
12
12
10
10
10
10
10
XY
XY
XY
XY
XY
XY
XY
XY
292
XY
XY
XY
XY
XY
XY
XY
292
293
292
293
292
292
292
293
211
292
293
293
292
292
293
292
Taxa
Sexual
System
Chrysops indus
Chrysops Indus
Chrysops ludens
Chrysops mlokosiewiczi
Chrysops pictus
Chrysops relictus
Chrysops shermani
Chrysops shermani
Chrysops suavis
Chrysops vanderwulpi
Chrysops vittatus
Chrysops vittatus
homomorphic
Chrysops vittatus
Dasyrhamphis umbrinus
Haematopota crassicornis
Haematopota italica
Haematopota pallens
Haematopota pellucens
Haematopota pluvialis
Haematopota pluvialis
Haematopota scutellata rossica
Haematopota subcylindrica
Haematopota tamerlani
Hybomitra arpadi
Hybomitra bimaculata
Hybomitra brevis
Hybomitra ciureai
Hybomitra distinguenda distinguenda
Hybomitra erberi
Hybomitra lasiophthalma
Hybomitra lasiophthalma
Hybomitra lundbecki
Hybomitra m. acrocentrica
Hybomitra montana montana
Hybomitra muhlfeldi
Hybomitra nigella
Hybomitra peculiaris
Hybomitra stenopselapha
Hybomitra tarandina
Hybomitra tarandinoides
Hybomitra ussuriensis
Tabanus autumnalis
Tabanus bifarius
Tabanus bifarius
Tabanus bovines
Tabanus bromius
Tabanus buddha
Tabanus colchidicus
Tabanus cordiger
Tabanus cordiger
Tabanus dolini
Tabanus flavofemoratus
Tabanus geminus
Tabanus hauseri
Tabanus indrae
Tabanus infestus
Tabanus maculicornis
Tabanus marginalis
Tabanus marginalis
Tabanus miki
Tabanus pleskei
Tabanus quatuornotatus
Tabanus sabuletorum
Tabanus shelkovnikovi
Ploidy
18
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
10
10
10
10
10
12
8
10
10
12
8
10
10
10
10
10
10
12
8
10
10
12
8
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
293
292
292
292
292
292
293
292
292
292
292
293
12
12
18
14
18
14
18
18
18
26
18
12
14
18
12
16
10
14
14
18
16
16
10
18
14
18
18
10
18
14
16
16
14
10
12
16
12
12
12
10
14
14
10
10
10
10
10
10
12
16
12
12
12
12
18
14
18
14
18
18
XY
26
18
12
14
18
12
16
10
14
14
18
16
16
10
18
14
18
18
10
18
14
16
16
14
10
12
16
12
12
12
10
14
14
10
10
10
10
10
10
12
16
12
12
XY
XY
XY
XY
XY
XY
XY
XY
292
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
293
292
292
292
292
292
292
293
212
XY
XY
XY
XY
292
292
292
292
292
292
292
292
293
292
292
292
292
292
292
292
292
292
292
292
292
292
293
292
292
292
292
292
293
292
292
292
292
292
292
292
292
293
292
292
292
292
292
Taxa
Tabanus spectabilis
Tabanus sudeticus
Tabanus unifasciatus
Tabanus zimini
Tachinidae
Aplomya caesar
Aplomya caesar
Aplomya caesar
Aplomya mitis
Aplomya mitis
AplomyaE mitis
Archylas apicifer
Archytas apicifera
Bessa selecta
Bessa selecta
Bessa selecta
Bessa selecta
Ceracia dentata
Ceracia dentata
Ceromasia auricaudata
Ceromasia auricaudata
Ceromasia auridaudata
Drino behemica
Drino bohemica
Drino bohemica
Eumea westermanni
Eumea westermanni
Eumea westermanni
Lydella grisescens
Lydella grisescens
Lydella grisescens
Madremyia saundersii
Madremyia saundersii
Madremyia saundersii
Mericia ampelus
Mericia ampelus
Namorilla pyste
Nemorella pyste
Nemorilla pyste
Neopharocera hamata
Neopharocera hamata
Neophorocera hamata
Neophorocera hemata
Omoloma fumiferanae
Omotoma fumiferanae
Omotoma fumiferonae
Peleteria iterans
Peletieria iterans
Phorocera hamata
Phorocera hamata
Phorocera hemata
Phryxe pecosensis
Phryxe pecosensis
Phyrxe pecosensis
Spathimeigenia sp
Spathimeigenia sp.
Winthemia dantanae
Winthemia datanae
Winthemia datanae
Winthemia occidentis
Winthemia occidentis
Winthemia occidentis
Winthemia rufopicla
Winthemia rufopicta
Tephritidae
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
14
12
12
10
14
12
12
XY
XY
XY
XY
292
292
292
292
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
294
270
262
270
262
294
270
262
262
262
270
270
294
270
294
270
262
262
294
270
294
270
262
294
270
262
294
270
262
294
270
262
270
294
270
270
294
262
270
294
262
262
270
262
266
262
294
262
270
270
294
262
294
270
294
270
262
270
262
213
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
Taxa
Acinia fucata
Acinia mallochi
Afrodacus jarvisi
Anastepha ludens
Anastrepha amita
Anastrepha aphelocentema
Anastrepha aphelocentema
Anastrepha aphelocentoma
Anastrepha barnesi
Anastrepha barnesi
Anastrepha bistrigata
Anastrepha bistrigata
Anastrepha distincta
homomorphic
Anastrepha distincta
Anastrepha distincta
Anastrepha fraterculus
homomorphic
Anastrepha fraterculus
Anastrepha fraterculus
Anastrepha fraterculus
Anastrepha grandis
Anastrepha leptozona
Anastrepha ludens
Anastrepha ludens
Anastrepha ludens
homomorphic
Anastrepha mombinpraeoptans
Anastrepha montei
Anastrepha oblicua
Anastrepha obliqua
Anastrepha pickeli
Anastrepha pickeli
Anastrepha pseudoparalella
Anastrepha pseudoparallela
Anastrepha serpentina
Anastrepha serpentina
Anastrepha serpentina
Anastrepha sororcula
Anastrepha spatula
Anastrepha spatulata
Anastrepha spatulata
Anastrepha striata
Anastrepha striata
Anastrepha striata
Anastrepha turpiniae
Anastrepha zenildae
homomorphic
Anastrepha zuelaniae
Anastrepha zuelaniae
Anastrepha zuelaniae
homomorphic
Austrodacus cucumis
Bactrocera cucurbitae
Bactrocera diversus
Bactrocera oleae
Bactrocera zonatus
Cecidocharella borrichia
Ceratitis capitata
Ceratitis capitata
Chrysotrypanea sp
Chrysotrypanea trifasciata
Dacus olea
Dioxina chilensis
Diplodacus sienatiles
Sexual
System
Ploidy
12
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
14
10
12
12
12
12
12
12
11
12
12
12
12
14
10
12
12
12
12
12
12
12
11
12
ZW
ZW
XY
12
12
12
12
12
12
XY
XY
262
296
295
12
12
12
12
10
10
10
12
12
12
12
12
10
10
10
12
XY
XY
XY
XY
XY
XY
XY
295
262
296
296
296
295
262
296
12
8
12
12
8
8
12
12
11
10
12
12
12
12
12
12
12
12
12
12
XY
8
12
12
8
8
12
12
12
10
11
12
12
12
12
12
12
12
12
12
262
XY
XY
XY
XY
XY
XY
XY
XXY
XY
XXY
XY
XY
XY
XY
XY
XY
XY
XY
XY
296
295
296
295
296
296
295
295
262
296
296
295
262
296
295
262
296
296
296
12
12
12
12
12
12
XY
XY
XY
295
262
296
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
XY
XY
XY
XY
XY
ZW
XY
XY
ZW
ZW
XY
262
295
295
295
295
295
295
262
295
295
262
295
262
214
XY
XY
XY
XY
XY
XY
XXY
XXY
XY
295
295
262
266
296
295
296
262
295
296
295
296
295
Taxa
Dyseuaresta impluvita
homomorphic
Dyseuaresta mexicana
Epochra canadensis
Euaresta melanogaster
Euaresta melanogsaster
Procecidochares utilis
Rhachiptera limbata
Rhagoletis basiola
Rhagoletis berberedis
Rhagoletis berberis
Rhagoletis boycei
homomorphic
Rhagoletis cerasi
Rhagoletis cingulata
homomorphic
Rhagoletis completa
homomorphic
Rhagoletis conivora
Rhagoletis conversa
Rhagoletis fausta
homomorphic
Rhagoletis indifferens
homomorphic
Rhagoletis jungladis
homomorphic
Rhagoletis juniperina
homomorphic
Rhagoletis meigeni
Rhagoletis mendax
homomorphic
Rhagoletis nova
Rhagoletis pomella
homomorphic
Rhagoletis ribicola
homomorphic
Rhagoletis striatelia
Rhagoletis suavis
homomorphic
Rhagoletis tabelaria
homomorphic
Rhagoletis tomatis
Rhagoletis zephyria
homomorphic
Rhagoletis zoqui
homomorphic
Strumata bryoniae
Strumata cacuminata
Strumata humeralis
Strumata tryoni
Tehpritis arnicae
Tephritis arnicae
Tephritis arnicae
Tephritis sp.
Trupanea chrysanthemifolii
homomorphic
Trupanea foliosi
homomorphic
Trupanea footei
homomorphic
Trupanea thuriferae
homomorphic
Trypanaresta marisolae
homomorphic
Zonosemata electa
Sexual
System
Ploidy
Female
2n
Male
2n
12
12
12
10
12
12
12
12
10
12
14
12
12
10
12
12
12
12
10
12
14
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
295
12
12
295
12
12
295
10
12
10
12
XO
295
295
12
12
12
12
XY
295
295
12
12
11
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
11
12
12
12
12
12
12
12
11
12
12
12
12
12
295
12
12
295
12
12
295
12
12
295
12
12
215
Sex Chrom. Source
Karyotype
295
XY
XY
ZW
XY
XY
XY
XY
295
262
263
266
295
295
295
295
295
295
295
295
295
XY
XY
295
295
295
295
XXY
295
295
295
XY
295
295
295
XY
XY
XY
XY
XY
XO
XO
ZW
XY
262
262
262
262
262
266
295
295
295
295
Taxa
Zonosemata vittigera
Tetanoceratidae
Tetanocera sparea
Thaumaleidae
Thaumalea testacea
Tipulidae
Dictenidia bimaculala
Tipula paludosa
Tipula paludosa
Tipula sp
Tipula sp.
Trichoceridae
Melusine sp.
homomorphic
Ulididae
Chaetopsis fulvifrons
Xylomyidae
Solva pallipes
Embiidina
Embiidae
Cleomia guareschii
Embia tyrrhenica
Embia nuragica
Embia ramburi
Oligotomidae
Haploembia solieri
Haploembia palaui
Oligotoma japonica
Oligotoma saundersi
Ephemeroptera
Baetidae
Baetis rhodani
Baetis vernus
Cloeon dipterum
Cloeon triangulifer
Caenidae
Caenis horaria
Ephemerellidae
Ephemerella ignita
Ephemeridae
Ephemera danica
Heptageniidae
Ecdyonurus dispar
Siphlonuridae
Ameletus costalis
Grylloblata
Grylloblatidae
Galloisiana nipponensis
heteromorphic
Grylloblatta campodeioformis
Hemiptera
Aclerdidae
Aclerda berlesii
Aclerda sp.
Aleyrodidae
Aleurotulus nephrolepidis
holocentric HD arrhenotoky
Aleyrodes proletella
holocentric HD arrhenotoky
Bemisia tabaci
holocentric HD arrhenotoky
Trialeurodes vaporarium
holocentric HD arrhenotoky
Asterolecaniidae
Asterodiaspis quercicola
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
12
12
XY
295
12
12
XY
266
6
6
XY
261
8
8
8
8
8
8
8
8
8
8
XY
261
266
261
266
261
6
6
261
8
8
266
10
10
XY
280
22
22
24
22
21
21
23
21
XO
XO
XO
XO
297
297
297
297
20
22
20
22
19
21
19
21
XO
XO
XO
XO
297
297
297
297
10
10
10
10
10
10
XY
XY
XY
298
298
298
299
9
8
XO
300
16
16
XY
300
12
11
XO
300
20
20
XY
300
18
18
XY
298
30
30
XY
243
XY
243
XY
XY
parth
301
302
18
parth
26
13
303
28
14
303
22
11
303
22
11
303
24
216
302
Taxa
Russellaspis pustulans
Astrolecaniidae
Abditicoccus acaciae
Amorphococcus mesuae
Anomalococcus indicus
Endernia sp.
Mycetococcus ehrhorni
PGE
Callipappidae
Callipappus rubiginosus
Platycoelostoma tasmanicum
Cerococcidae
Cerococcus indicus
Cerococcus quercus
PGE
Coccidae
Acanthopulvinaria orientalis
PGE
Ceroplastes _oridensis
Ceroplastes actiniformes
Ceroplastes ceriferus
Ceroplastes cirripediformis
Ceroplastes cirripediformis
Ceroplastes japonicus
PGE
Ceroplastes pseudoceriferus
Ceroplastes rubens
Cissococcus fulleri
Coccus capparidis
Coccus hesperidum
Coccus longulus
Coccus pseudomagnoliarum
Coccus viridis
Drepanococcus cajani
Ericerus pela
Eriopeltis lichtensteini
PGE
Eriopeltis stammeri
PGE
Eucalymnatus euterones
Eucalymnatus tessellatus
Eulecanium ciliatum
Eulecanium douglasi
Eulecanium tiliae
Lichtensia viburni
PGE
Luzulaspis dactylis
PGE
Neolecanium cornuparvum
PGE
Neopulvinaria innumerabilis
Parthenolecanium cerasifex
arrhenotoky
Parthenolecanium cerasifex
Parthenolecanium corni
PGE
Parthenolecanium pomeranicum
Parthenolecanium putmani
Physokermes hemicryphus
Protopulvinaria pyriformis
Pulvinaria _occifera
Pulvinaria aurantii
PGE
Pulvinaria hydrangeae
Pulvinaria peregrina
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
parth
304
parth
parth
305
305
302
305
302
14
6
14
parth
302
302
18
302
36
32
36
302
302
302
306
306
302
36
36
36
14
302
307
308
309
302
302
309
309
302
310
302
14
302
16
16
18
18
18
18
302
311
302
302
302
302
18
302
34
302
16
302
302
16
16
302
302
16
23
16
16
26
312
302
302
302
302
302
16
16
302
302
14
14
parth
parth
18
parth
parth
parth
parth
parth
parth
parth
parth
parth
2 or 3
302
305
14
18
parth
parth
parth
XO
217
Taxa
Pulvinaria polygonata
Pulvinaria psidii
Pulvinaria ribesiae
PGE
Pulvinaria vitis
Pulvinariella mesembryanthemi
Rhizopulvinaria variabilis
Saissetia coffeae
Sphaerolecanium prunastri
PGE
Vinsonia stellifera
Conchaspididae
Conchaspis lepagei
PGE
Dactylopiidae
Dactylopius ceylonicus
PGE
Dactylopius coccus
PGE
Dactylopius confusus
PGE
Dactylopius opuntiae
Dactylopius opuntiae
PGE
Diaspididae
Abgrallaspis cyanophylli
PGE
Abgrallaspis flavida
PGE
Acutaspis paulista
PGE
Acutaspis perseae
PGE
Africaspis chionaspiformis
PGE
Africaspis fici
PGE
Ancepaspis edentata
PGE
Ancepaspis tridentata
PGE
Aonidia lauri
PGE
Aonidia shastae
PGE
Aonidiella aurantii
PGE
Aonidiella citrina
PGE
Aonidiella orientalis
PGE
Aonidiella simplex
PGE
Aonidomytilus concolor
PGE
Aonidomytilus variabilis
PGE
Aspidaspis arctostaphyli
PGE
Aspidaspis densiflorae
PGE
Aspidiella hartii
PGE
Aspidiella sacchari
PGE
Sexual
System
parth
parth
parth
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
14
18
302
302
302
16
28
16
18
302
302
302
302
302
10
302
12
302
10
302
16
302
10
302
10
313
302
8
302
8
302
8
302
8
302
8
302
12
302
8
302
6
302
8
302
8
302
8
302
8
302
8
302
8
302
12
302
10
302
8
302
8
302
8
302
8
302
parth
218
Taxa
Aspidiotus cryptomeriae
PGE
Aspidiotus destructor
PGE
Aspidiotus hedericola
PGE
Aspidiotus nerii
PGE
Aspidiotus nerii
Aspidiotus simulans
PGE
Aulacaspis rosae
PGE
Aulacaspis spinosa
PGE
Aulacaspis tubercularis
PGE
Berlesaspis spinifera
Carulaspis minima
PGE
Chionaspis ortholobis
PGE
Chionaspis pinifoliae
PGE
Chionaspis pinifoliae
Chionaspis platani
PGE
Chionaspis salicis
PGE
Chrysomphalus aonidum
PGE
Chrysomphalus bifasciculatus
PGE
Chrysomphalus dictyospermi
Chrysomphalus pinnulifer
Clavaspis coursetiae
PGE
Clavaspis texana
PGE
Comstockiella sabalis
PGE
Cooleyaspis praelonga
PGE
Costalimaspis eugeniae
PGE
Crassaspis multipora
PGE
Diaspidiotus aesculi
PGE
Diaspidiotus ancylus
PGE
Diaspidiotus gigas
PGE
Diaspidiotus lenticularis
PGE
Diaspidiotus ostreaeformis
PGE
Diaspidiotus perniciosus
PGE
Diaspidiotus zonatus
PGE
Diaspis boisduvalii
PGE
Diaspis bromeliae
PGE
Sexual
System
parth
parth
parth
parth
Ploidy
2
2
2
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
8
302
8
302
8
302
8
302
8
8
302
302
8
302
8
302
8
302
8
305
302
8
302
8
302
8
8
302
302
8
302
8
302
8
302
8
8
8
302
302
302
8
302
10
302
18
302
12
302
8
302
8
302
8
302
8
302
8
302
8
302
8
302
8
302
8
302
8
302
219
Taxa
Diaspis echinocacti
PGE
Duplachionaspis sicula
PGE
Dynaspidiotus apacheca
PGE
Dynaspidiotus californicus
PGE
Dynaspidiotus pseudomeyeri
PGE
Epidiaspis leperii
Epidiaspis persimilis
PGE
Fiorinia fioriniae
Fiorinia japonica
PGE
Froggattiella penicillata
PGE
Furcaspis capensis
PGE
Furcaspis biformis
PGE
Greenaspis decurvata
PGE
Gymnaspis aechmeae
PGE
Helaspis mexicana
PGE
Hemiberlesia cupressi
Hemiberlesia lataniae
PGE
Hemiberlesia lataniae
Hemiberlesia palmae
Hemiberlesia quercicola
PGE
Hemiberlesia rapax
Howardia biclavis
Ischnaspis longirostris
Kuwanaspis bambusicola
Kuwanaspis pseudoleucaspis
Ledaspis reticulata
PGE
Ledaspis tenuiloba
PGE
Lepidosaphes beckii
PGE
Lepidosaphes conchiformis
PGE
Lepidosaphes tokionis
PGE
Lepidosaphes ulmi
PGE
Leucaspis lowi/loewi
Leucaspis pusilla
PGE
Lindingaspis ferrisi
PGE
Lindingaspis opima
PGE
Lindingaspis rossi
PGE
Melanaspis glomerata
PGE
Melanaspis inopinata
Melanaspis lilacina
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
16
302
8
302
8
302
314
parth
2
8
315
8
8
302
302
8
8
302
302
8
302
8
316
8
302
8
302
10
302
12
302
parth
2
8
8
302
302
parth
parth
2
2
8
8
8
302
302
302
parth
parth
parth
parth
parth
2
2
2
2
2
8
10
8
10
8
8
302
302
302
302
302
302
18
302
8
315
8
302
12
302
16
302
11
8
302
302
8
302
8
302
8
302
8
302
8
8
302
302
parth
2
220
Taxa
PGE
Morganella longispina
PGE
Mycetaspis juventinae
PGE
Mycetaspis personata
PGE
Neomorgania eucalypti
PGE
Neoselenaspidus kenyae
Neoselenaspidus silvaticus
Nicholiella bumeliae
PGE
Nikkoaspis shiranensis
PGE
Odonaspis ruthae
Odonaspis saccharicaulis
PGE
Opuntiaspis philococcus
PGE
Parlatoria crotonis
PGE
Parlatoria oleae
PGE
Parlatoria proteus
PGE
Parlatoria proteus
Parlatoria ziziphi
PGE
Pinnaspis aspidistrae
PGE
Pinnaspis buxi
Pinnaspis strachani
PGE
Poliaspoides formosana
Protodiaspis agrifoliae
PGE
Protodiaspis chichi
PGE
Protodiaspis didymus
PGE
Protodiaspis infidelis
PGE
Protodiaspis signata
PGE
Pseudaonidia baikeae
PGE
Pseudaonidia trilobitiformis
PGE
Pseudaulacaspis pentagona
PGE
Pseudischnaspis bowreyi
PGE
Pseudoparlatoria browni
PGE
Pseudoparlatoria ostreata
PGE
Pseudoparlatoria parlatorioides
PGE
Radionaspis indica
PGE
Rhizaspidiotus dearnessi
PGE
Rolaspis anacantha
PGE
Sexual
System
parth
parth
parth
parth
Ploidy
2
2
2
2
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
8
302
8
302
8
302
8
302
8
8
8
302
302
302
8
302
8
8
302
302
8
302
8
302
8
302
8
302
8
8
302
302
8
302
parth
2
8
8
302
302
parth
2
8
8
302
302
8
302
8
302
6
302
8
302
8
302
8
302
16
302
8
302
8
302
10
302
10
302
8
302
8
302
18
302
221
Taxa
Selenaspidus articulatus
PGE
Selenaspidus incisus
Situlaspis yuccae
PGE
Spinaspidiotus fissidens
PGE
Targionia bigeloviae
Targionia nigra
PGE
Targionia vitis
PGE
Targionia yuccarum
PGE
Trischnaspis bipindensis
Unachionaspis bambusae
PGE
Unaspis citri
PGE
Unaspis euonymi
PGE
Xanthophthalma concinnum
Xerophilaspis prosopidis
PGE
Eriococcidae
Acanthococcus agropyri
PGE
Acanthococcus insignis
PGE
Aculeococcus morrisoni
Aculeococcus yongpingensis
Apiomorpha attenuata
PGE
Apiomorpha baeuerleni
PGE
Apiomorpha calycina
PGE
Apiomorpha conica
PGE
Apiomorpha densispinosa
PGE
Apiomorpha dipsaciformis
PGE
Apiomorpha duplex
PGE
Apiomorpha excupula
PGE
Apiomorpha frenchi
PGE
Apiomorpha gullanae
Apiomorpha helmsii
PGE
Apiomorpha hilli
PGE
Apiomorpha intermedia
PGE
Apiomorpha karschi
PGE
Apiomorpha longiloba
PGE
Apiomorpha macqueeni
PGE
Apiomorpha maliformis
PGE
Apiomorpha malleeacola
Sexual
System
Ploidy
parth
parth
parth
parth
2
2
2
Female
2n
Sex Chrom. Source
Karyotype
8
302
8
10
302
302
8
302
8
8
302
302
8
302
8
302
8
8
302
302
8
302
8
302
16
8
302
302
16
302
16
302
56
305
305
317
5
317
92
317
19
317
4
317
18
317
26
317
18
317
17
317
73
98
318
302
42
302
12
302
93
302
178
302
192
302
18
302
32
302
parth
parth
parth
Male
2n
222
Taxa
PGE
Apiomorpha minor
PGE
Apiomorpha munita
PGE
Apiomorpha munita
PGE
Apiomorpha ovicola
PGE
Apiomorpha ovicoloides
PGE
Apiomorpha pedunculata
PGE
Apiomorpha pharetrata
PGE
Apiomorpha pileata
PGE
Apiomorpha regularis
PGE
Apiomorpha rosaeforma
PGE
Apiomorpha sessilis
PGE
Apiomorpha sloanei
PGE
Apiomorpha spinifer
PGE
Apiomorpha strombylosa
PGE
Apiomorpha subconica
PGE
Apiomorpha tepperi
PGE
Apiomorpha urnalis
PGE
Apiomorpha variabilis
PGE
Apiomorpha withersi
PGE
Ascelis praemollis
Ascelis schraderi
PGE
Calycicoccus merwei
PGE
Capulinia crateraformis
Capulinia jaboticabae
PGE
Capulinia orbiculata
PGE
Carpochloroides mexicanus
Carpochloroides viridis
Casuarinaloma leaii
PGE
Cryptococcus fagisuga
Cylindrococcus casuarinae
Cylindrococcus spiniferus
PGE
Cystococcus echiniformis
Cystococcus pomiformis
Eremococcus pirogallis
Eremococcus rugosus
Eremococcus rugosus
Eremococcus turbinata
Eriococcus abditus
PGE
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
47
302
154
302
15
302
62
302
57
302
38
302
29
302
19
302
108
302
14
302
4
302
128
302
148
302
41
302
67
302
64
302
121
302
80
302
40
302
16
18
302
302
319
parth
18
305
302
18
302
12
319
319
302
6
18
305
302
302
16
305
305
305
305
305
305
302
parth
parth
parth
parth
parth
223
Taxa
Eriococcus acutispinatus
Eriococcus araucariae
PGE
Eriococcus arcanus
Eriococcus cavellii
PGE
Eriococcus coriaceus
PGE
Eriococcus detectus
PGE
Eriococcus elytranthae
Eriococcus ericae
PGE
Eriococcus fossor
Eriococcus hakeae
Eriococcus lecanioides
PGE
Eriococcus leptospermi
PGE
Eriococcus minus
PGE
Eriococcus montanus
Eriococcus rata
PGE
Eriococcus rhodomyrti
PGE
Eriococcus williamsi
PGE
Floracoccus elevans
Gossyparia spuria
PGE
Kuwanina obscurata
Lachnodius eucalypti
homomorphic
Lachnodius hirsutus
Lachnodius lectularius
Madarococcus totarae
PGE
Madarococcus viridulus
PGE
Opisthoscelis convexa
Opisthoscelis fibularis
Opisthoscelis maculata
Opisthoscelis mammularis
Opisthoscelis maskelli
Opisthoscelis subrotunda
Ourococcus cobbi
PGE
Phacelococcus subcorticalis
Phloeococcus loriceus
PGE
Scutare lanuginosa
PGE
Sisyrococcus intermedius
PGE
Sphaerococcus morrisoni
Sphaerococcus pustulans
Sphaerococcus socialis
Stegococcus oleariae
Tectococcus ovatus
Halimococcidae
Colobopyga browni
PGE
Colobopyga pritchardiae
PGE
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
parth
18
305
302
18
305
302
18
302
18
302
14
305
302
18
18
305
302
302
18
302
16
302
14
305
302
16
302
14
302
28
305
302
parth
parth
parth
parth
parth
18
parth
XO
305
302
18
14
305
302
302
18
302
18
18
14
302
305
302
305
305
302
302
18
18
302
302
24
302
18
302
18
305
305
302
305
305
10
302
10
315
18
parth
parth
parth
224
Taxa
Halimococcus borassi
PGE
Platycoccus tylocephalus
PGE
Thysanococcus calami
Thysanococcus pandani
PGE
Kermesidae
Kermes quercus
PGE
Olliffiella cristicola
Reynvaania gallicola
Kerriidae
Kerria lacca
PGE
Tachardiella sp.
PGE
Kuwaniidae
Kuwania oligostigma
Lecanodiaspididae
Lecanodiaspis africana
PGE
Marchalinidae
Marchalina hellenica
Margarodidae
Margarodes vitis
Porphyrophora hamelii
Porphyrophora polonica
Matsucoccidae
Matsucoccus gallicolus
Matsucoccus pini
Monophlebidae
Aspidoproctus maximus
Auloicerya acaciae
haplodiploid type unspecified
Conifericoccus agathidis
Crypticerya rosae
haplodiploid type unspecified
Crypticerya sp.
haplodiploid type unspecified
Drosicha sp.
Echinicerya anomala
haplodiploid type unspecified
Gigantococcus maximus
haplodiploid type unspecified
Gueriniella serratulae
Icerya aegyptiaca
haplodiploid type unspecified
Icerya aegyptiaca
Icerya bimaculata
Icerya brasiliensis
haplodiploid type unspecified
Icerya littoralis
haplodiploid type unspecified
Icerya montserratensis
haplodiploid type unspecified
Icerya purchasi
Icerya similis
haplodiploid type unspecified
Icerya tremae
haplodiploid type unspecified
Llaveia axin
Llaveia oaxacoensis
Llaveiella taenechina
Nautococcus schraderae
Sexual
System
parth
Ploidy
2
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
302
10
302
10
10
302
302
26
302
parth
parth
305
305
18
302
20
302
16
XO
302
302
parth
18
302
parth
302
302
302
14
14
XO
XO
40
XXXXXXO
302
320
6
4
XO
302
321
parth
4
305
302
4
302
8
4
302
302
4
302
2
6
4
302
302
parth
hermaphrodite
4
4
4
302
302
302
4
302
4
302
4
4
302
302
4
302
parth
hermaphrodite
6
6
6
6
225
XO
XO
XO
XO
302
302
302
302
Taxa
Protortonia navesi
Protortonia primitiva
Steatococcus tuberculatus
haplodiploid type unspecified
Ortheziidae
Insignorthezia insignis
Newstedia sp.
Orthezia urticae
Praelongorthezia praelonga
homomorphic
Phenacoleachiidae
Phenacoleachia zealandica
Phoenicococcidae
Colobopyga browni
PGE
Colobopyga pritchardiae
PGE
Halimococcus borassi
PGE
Phoenicococcus marlatti
PGE
Platycoccus tylocephalus
PGE
Thysanococcus calami
Thysanococcus pandani
PGE
Pseudococcidae
Antonina crawi
Antonina evelynae
PGE
Antonina graminis
Antonina pretiosa
Atrococcus paludinus
PGE
Balanococcus singularis
PGE
Cataenococcus olivaceus
Chaetococcus bambusae
Chlorococcus straussiae
Chorizococcus pusillus
Chorizococcus rostellum
Clavicoccus tribulus
Coccidohystrix insolita
PGE
Coccura suwakoensis
Dysmicoccus boninsis
Dysmicoccus brevipes
PGE
Dysmicoccus brevipes
Dysmicoccus multivorus
PGE
Dysmicoccus neobrevipes
Dysmicoccus wistariae
Erium pygmaeum
Ferrisia virgata
PGE
Ferrisia virgata
Fonscolombia butorinae
PGE
Fonscolombia tomlinii
PGE
Formicococcus ireneus
Formicococcus robustus
PGE
Heterococcus nudus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
parth
XO
14
18
16
XO
XO
323
324
325
326
8
XO
327
parth
parth
parth
parth
parth
parth
parth
322
302
302
6
4
10
315
10
315
10
315
18
315
10
315
10
10
315
302
12
12
302
302
16
24
10
302
302
302
10
302
38
10
38
10
10
14
12
302
302
302
302
302
302
302
10
12
10
302
302
302
10
10
302
302
10
10
10
10
302
302
302
302
10
10
302
302
10
302
10
10
302
302
10
302
226
Taxa
PGE
Laminicoccus pandani
Laminicoccus vitiensis
Maconellicoccus hirsutus
PGE
Maconellicoccus ugandae
Melanococcus viridis
Mirococcopsis subterranea
Mirococcus inermis
PGE
Nairobia bifrons
Neochavesia caldasiae
Nesococcus pipturi
Nesopedronia acanthocauda
Nesopedronia cibotii
Nesopedronia dura
Nesopedronia hawaiiensis
Nipaecoccus aurilanatus
Nipaecoccus graminis
Nipaecoccus nipae
PGE
Nipaecoccus viridis
PGE
Palmicultor browni
Palmicultor palmarum
Paracoccus bruguierae
Paracoccus burnerae
Paracoccus diversus
Paraputo leveri
Pedronia strobilanthis
Phenacoccus acericola
PGE
Phenacoccus aceris
PGE
Phenacoccus defectus
Phenacoccus dicoriae
Phenacoccus gossypii
PGE
Phenacoccus graminicola
Phenacoccus helianthi
PGE
Phenacoccus infernalis
Phenacoccus manihoti
Phenacoccus phenacoccoides
PGE
Phenacoccus pumilus
PGE
Phenacoccus solani
Phenacoccus solenopsis
Planococcoides crassus
Planococcus citri
PGE
Planococcus lilacinus
Planococcus minor
Planococcus vovae
PGE
Porococcus tinctorius
Pseudococcus swezeyi
Pseudococcus _origer
Pseudococcus antricolens
Pseudococcus calceolariae
PGE
Pseudococcus comstocki
Pseudococcus kikuyuensis
Pseudococcus longispinus
Sexual
System
parth
Ploidy
Female
2n
Sex Chrom. Source
Karyotype
10
16
10
302
302
302
10
16
10
10
302
302
302
302
14
14
12
18
14
10
10
10
10
10
302
328
302
302
302
302
302
302
302
302
10
302
8
10
10
10
10
10
8
12
302
302
302
302
302
302
302
302
10
302
10
10
12
302
302
302
10
10
302
302
10
10
302
329
302
10
302
10
10
10
10
302
302
302
302
10
16
10
302
302
302
18
10
10
10
10
302
302
302
302
302
10
10
8
302
302
302
parth
parth
Male
2n
227
Taxa
Sexual
System
Pseudococcus lycopodii
Pseudococcus maritimus
Pseudococcus montanus
Pseudococcus nudus
Pseudococcus occidius
Pseudococcus pipturicolus
Pseudococcus viburni
PGE
Pseudotrionymus multiductus
parth
Rastrococcus iceryoides
PGE
Rhizoecus dianthi
Rhizoecus falcifer
Rhizoecus mayanus
Rhizoecus mexicanus
PGE
Saccharicoccus sacchari
Spilococcus mamillariae
PGE
Spilococcus mamillariae
parth
Spilococcus sequoiae
Trionymus aberrans
PGE
Trionymus caricis
Trionymus insularis
parth
Trionymus longipilosus
Trionymus perrisii
PGE
Vryburgia amaryllidis
Vryburgia transvaalensis
parth
Xenococcus annandalei
Putoidae
Puto albicans
Puto antennatus
Puto arctostaphyli
Puto paci_cus
Puto sp.
Puto superbus
Puto yuccae
Steingeliidae
Araucaricoccus queenslandicus parth
Steingelia gorodetskia
Steingelia gorodetskia
Stictococcidae
Stictococcus sp.
homomorphic
Hymenoptera
Andrenidae
Andrena (Hesperandrena) duboisi
arrhenotoky
Andrena (Hesperandrena) sp.
arrhenotoky
Andrena togashii
arrhenotoky
Aphelinidae
Aphelinus mali
arrhenotoky
Aphytis mytilaspidus
arrhenotoky
Archenomus orientalis
arrhenotoky
Encarsia berlesei
arrhenotoky
Encarsia pergandiella
arrhenotoky
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
10
10
18
10
10
10
10
302
302
302
302
302
302
302
18
10
302
302
12
12
10
8
302
302
302
302
10
20
302
302
20
24
16
302
302
302
8
10
10
16
302
302
302
302
10
10
302
302
305
20
XO
20
16
XO
XO
XO
20
XO
302
305
302
302
302
305
302
10
10
319
XO
XO
302
302
330
6
3
331
20
10
331
6
3
332
10
5
333
10
5
333
22
11
333
10
5
333
12
6
333
228
Taxa
Sexual
System
Apidae
Camargoia nordestina
arrhenotoky
Cephalotrigona capitata
arrhenotoky
Frieseomelitta doederleini
arrhenotoky
Frieseomelitta languida
arrhenotoky
Frieseomelitta varia
arrhenotoky
Geotrigona mombuca
arrhenotoky
Lestrimelitta limao
arrhenotoky
Mourella caerulea
arrhenotoky
Nannotrigona sp.1
arrhenotoky
Nannotrigona testaceicornis
arrhenotoky
Paratrigona subnuda
arrhenotoky
Ptilotrigona lurida
arrhenotoky
Scaptotrigona acantha
arrhenotoky
Scaptotrigona angustula
arrhenotoky
Scaptotrigona angustula
arrhenotoky
Scaptotrigona appendiculata
arrhenotoky
Scaptotrigona babai
arrhenotoky
Scaptotrigona barrocoloralensis
arrhenotoky
Scaptotrigona chanchamayoensis
arrhenotoky
Scaptotrigona clavipes
arrhenotoky
Scaptotrigona dentipes
arrhenotoky
Scaptotrigona depilis
arrhenotoky
Scaptotrigona esakii
arrhenotoky
Scaptotrigona fenestrata
arrhenotoky
Scaptotrigona fenestrata
arrhenotoky
Scaptotrigona fiebrigi
arrhenotoky
Scaptotrigona flavipes
arrhenotoky
Scaptotrigona fulgidus longiplumosis
arrhenotoky
Scaptotrigona fulviventris
arrhenotoky
Scaptotrigona fuscipennis
arrhenotoky
Scaptotrigona japonica
arrhenotoky
Scaptotrigona latitarsis
arrhenotoky
Ploidy
34
Female
2n
Male
2n
34
17
334
34
17
334
30
15
334
30
15
334
30
15
334
30
15
334
28
14
334
34
17
334
34
17
334
34
17
334
34
17
334
22
11
334
34
17
331
34
17
332
34
17
334
32
16
332
32
16
332
17
Sex Chrom. Source
Karyotype
332
34
17
335
34
17
334
34
17
332
34
17
334
32
16
332
32
16
336
32
16
331
34
17
337
34
17
332
26
13
331
32
16
338
34
17
334
34
17
332
34
17
334
229
Taxa
Sexual
System
Ploidy
Scaptotrigona megastigmata
arrhenotoky
Scaptotrigona minangkabou
arrhenotoky
Scaptotrigona muelleri
arrhenotoky
Scaptotrigona okinawana
arrhenotoky
Scaptotrigona pectoralis barrocoloradensis
34
arrhenotoky
Scaptotrigona postica
arrhenotoky
Scaptotrigona quadripunctata
34
arrhenotoky
Scaptotrigona recursa
arrhenotoky
Scaptotrigona smaragdula
arrhenotoky
Scaptotrigona sp.1
arrhenotoky
Scaptotrigona spinipes
arrhenotoky
Scaptotrigona spinipes
arrhenotoky
Scaptotrigona subterranea
arrhenotoky
Scaptotrigona violacea
arrhenotoky
Scaptotrigona violacea
arrhenotoky
Scaptotrigona xanthotricha
arrhenotoky
Anthophora acervorum villosula
18
arrhenotoky
Anthophora bomboides
arrhenotoky
Anthophora californica
arrhenotoky
Anthophora plumipes
arrhenotoky
Apis cerana
arrhenotoky
Apis cerana indica
arrhenotoky
Apis cerana japonica
arrhenotoky
Apis dorsata
arrhenotoky
Apis florea
arrhenotoky
Apis mellifera
arrhenotoky
Apis mellifera adansonii
arrhenotoky
Apis mellifera ligustica
arrhenotoky
Apis mellifera mellifera
arrhenotoky
Axestotrigona ferruginea
arrhenotoky
Bombus (Bombias) nevadensis
36
arrhenotoky
Bombus (megabombus) consobrinus wittenburghi
arrhenotoky
Bombus affinis
Female
2n
Male
2n
34
17
332
40
20
332
16
8
339
34
17
332
17
34
Sex Chrom. Source
Karyotype
331
17
17
334
334
34
17
334
28
14
332
34
17
334
34
17
331
34
17
334
34
17
331
32
16
331
32
16
340
34
17
334
9
332
36
18
331
38
19
331
18
9
331
32
16
341
32
16
342
32
16
332
32
16
343
32
16
344
32
16
331
32
16
345
32
16
332
32
16
346
36
18
347
18
348
38
19
332
36
18
348
230
Taxa
Sexual
System
arrhenotoky
Bombus atratus
arrhenotoky
Bombus deuteronymus maruhanabachi
arrhenotoky
Bombus diversus
arrhenotoky
Bombus honshuensis
arrhenotoky
Bombus hypocrita
arrhenotoky
Bombus ignitus
arrhenotoky
Bombus lucorum lucorum
arrhenotoky
Bombus moderatus
arrhenotoky
Bombus morio
arrhenotoky
Bombus pennsylvanicus
arrhenotoky
Bombus pseudobaicalensis
arrhenotoky
Bombus schrencki
arrhenotoky
Bombus terrestris
arrhenotoky
Bombus terricola
arrhenotoky
Bombus terricola occidentalis
arrhenotoky
Bombus ussurensis
arrhenotoky
Celetrigona longicornis
arrhenotoky
Cleptotrigona cubiceps
arrhenotoky
Cullumanobombus rufocincus
arrhenotoky
Dactylurina staudingeri
arrhenotoky
Diadasia enavata
arrhenotoky
Eufriesia violacea
arrhenotoky
Euglossa cyanaspis
arrhenotoky
Euglossa hyacinthina
arrhenotoky
Euglossa sp.
arrhenotoky
Exomalopsis aureopilosa
arrhenotoky
Exomalopsis sp.
arrhenotoky
Fervidobombus atratus
arrhenotoky
Fervidobombus californicus
arrhenotoky
Fervidobombus fervidus
arrhenotoky
Fervidobombus morio
arrhenotoky
Fervidobombus pennsylvanicus
arrhenotoky
Ploidy
36
Female
2n
Male
2n
40
20
339
46
23
332
36
18
332
34
17
332
36
18
332
36
18
332
36
18
331
36
18
348
40
20
339
36
18
331
34
17
332
34
17
331
36
18
349
36
18
348
36
18
348
36
18
332
30
15
350
36
18
347
38
19
348
34
17
331
30
15
331
30
15
331
42
21
331
40
20
331
42
21
331
18
9
331
16
8
331
40
20
348
38
19
348
36
18
348
40
20
348
18
231
Sex Chrom. Source
Karyotype
348
Taxa
Sexual
System
Frieseomelitta (Duckeola) ghiliani
arrhenotoky
Frieseomelitta doederleini
arrhenotoky
Frieseomelitta languida
arrhenotoky
Frieseomelitta varia
arrhenotoky
Hypotrigona braunsi
arrhenotoky
Hypotrigona gribodoi
arrhenotoky
Isepeolus viperinus
arrhenotoky
Leurotrigona muelleri
arrhenotoky
Leurotrigona pusilla
arrhenotoky
Meliplebeia beccari
arrhenotoky
Melipona asilvae
arrhenotoky
Melipona bicolor bicolor
arrhenotoky
Melipona capixaba
arrhenotoky
Melipona compressipes
arrhenotoky
Melipona compressipes manaosensis
arrhenotoky
Melipona crinita
arrhenotoky
Melipona favosa
arrhenotoky
Melipona interrupta fasciculata
arrhenotoky
Melipona mandacaia
arrhenotoky
Melipona marginata
arrhenotoky
Melipona marginata marginata
arrhenotoky
Melipona mondury
arrhenotoky
Melipona nigra
arrhenotoky
Melipona quadrifasciata
arrhenotoky
Melipona quinquefasciata
arrhenotoky
Melipona rufiventris
arrhenotoky
Melipona scutellaris
arrhenotoky
Melipona subnitida
arrhenotoky
Meliponula (Meliplebeia) becarii
arrhenotoky
Meliponula bocandei
arrhenotoky
Mourella caerulea
arrhenotoky
Nannotrigona sp.
arrhenotoky
Nannotrigona testaceicornis
Ploidy
Female
2n
30
15
18
18
34
Male
2n
Sex Chrom. Source
Karyotype
351
30
15
340
30
15
352
30
15
340
28
14
351
28
14
351
32
16
351
16
8
331
30
15
331
34
17
331
18
9
353
18
9
353
18
9
353
18
9
353
18
9
351
18
9
353
18
9
332
9
354
18
9
353
18
9
331
9
355
18
9
356
18
9
354
18
9
355
28
14
351
18
9
354
18
9
353
18
9
340
17
351
36
18
347
34
17
352
34
17
352
34
17
340
232
Taxa
arrhenotoky
Oxytrigona sp. cf. flaveola
arrhenotoky
Oxytrigona tataira
arrhenotoky
Partamona ailyae
arrhenotoky
Partamona cupira
arrhenotoky
Partamona helleri
arrhenotoky
Partamona mulata
arrhenotoky
Partamona pearsoni
arrhenotoky
Partamona peckolti
arrhenotoky
Partamona peckolti musarum
arrhenotoky
Partamona seridoensis
arrhenotoky
Partamona sp.n.
arrhenotoky
Partamona vicina
arrhenotoky
Plebeia (Friesella) schrottkyi
arrhenotoky
Plebeia denoiti
arrhenotoky
Plebeia droryana
arrhenotoky
Plebeia emerina
arrhenotoky
Plebeia remota
arrhenotoky
Plebeia sp.1
arrhenotoky
Plebeia subnuda
arrhenotoky
Plebeina denoiti
arrhenotoky
Psithyrus ashtoni
arrhenotoky
Psithyrus citrinus
arrhenotoky
Ptilothrix bombiformis
arrhenotoky
Pyrobombus ardens
arrhenotoky
Pyrobombus bimaculatus
arrhenotoky
Pyrobombus edwardsii
arrhenotoky
Pyrobombus ephippiatus
arrhenotoky
Pyrobombus huntii
arrhenotoky
Pyrobombus impatiens
arrhenotoky
Pyrobombus melanopygus
arrhenotoky
Pyrobombus mixtus
arrhenotoky
Pyrobombus perplexus
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
34
17
357
34
17
351
34
17
331
34
17
340
34
17
331
34
17
331
34
17
331
34
17
358
34
17
331
34
17
359
34
17
331
34
17
331
36
18
340
36
18
354
36
18
360
36
18
351
36
18
351
34
17
331
36
18
351
36
18
331
50
25
348
52
26
348
12
6
331
36
18
332
36
18
348
36
18
331
36
18
348
36
18
348
36
18
348
36
18
348
36
18
348
24
12
348
233
Sex Chrom. Source
Karyotype
Taxa
Pyrobombus sitkensis
arrhenotoky
Pyrobombus ternarius
arrhenotoky
Pyrobombus vagans
arrhenotoky
Pyrobombus vosnesenskii
arrhenotoky
Scaptotrigona depilis
arrhenotoky
Scaptotrigona postica
arrhenotoky
Separatobombus griseocollis
arrhenotoky
Subterraneobombus appositus
arrhenotoky
Subterraneobombus borealis
arrhenotoky
Svastra obliqua expurgata
arrhenotoky
Tetragona angustula
arrhenotoky
Tetragona clavipes
arrhenotoky
Trigona fuscipennis
arrhenotoky
Trigona spinipes
arrhenotoky
Trigona staudingeri
arrhenotoky
Trigona subterranea
arrhenotoky
Argidae
Arge cyanocrocea
arrhenotoky
Arge gracilicornis
arrhenotoky
Arge melanochroa
arrhenotoky
Arge nigripes
arrhenotoky
Arge pagana
arrhenotoky
Arge pectoralis
arrhenotoky
Arge ustulata
arrhenotoky
Bethylidae
Epyris nigir
arrhenotoky
Laelius utilis
arrhenotoky
Braconidae
Apanteles sp.1
arrhenotoky
Aphidius rhopalosiphi
arrhenotoky
Biosteres carbonarius
arrhenotoky
Charmon cruentatus
arrhenotoky
Cotesia glomeratus
arrhenotoky
Dacnusa sp.1
arrhenotoky
Sexual
System
Ploidy
32
Female
2n
Male
2n
36
18
348
36
18
348
36
18
348
36
18
348
34
17
340
34
17
340
38
19
348
16
Sex Chrom. Source
Karyotype
348
32
16
348
42
21
331
34
17
340
34
17
340
34
17
340
34
17
340
34
17
351
34
17
340
22
11
331
16
8
331
20
10
331
26
13
331
16
8
331
16
8
331
16
8
331
28
14
333
20
10
333
22
11
332
14
7
333
28
14
333
10
5
333
24
12
333
34
17
333
234
Taxa
Diaeretiella rapae
arrhenotoky
Ephedrus sp.1
arrhenotoky
Habrobracon hebetor
arrhenotoky
Habrobracon juglandis
arrhenotoky
Habrobracon pectinophorae
arrhenotoky
Habrobracon serinopae
arrhenotoky
Heterospilus prosopidis
arrhenotoky
Macrocentrus thoracicus
arrhenotoky
Meteorus gyrator
arrhenotoky
Meteorus pallipes
arrhenotoky
Meteorus versicolor
arrhenotoky
Mirax sp.1
arrhenotoky
Phaenocarpa persimilis
arrhenotoky
Praon abjectum
arrhenotoky
Rhysipolis decorator
arrhenotoky
Cephidae
Calameuta filiformis
arrhenotoky
Cephus cinctus
arrhenotoky
Janus integer
arrhenotoky
Chalcididae
Brachymeria intermedia
arrhenotoky
Brachymeria lasus
arrhenotoky
Brachymeria ovata
arrhenotoky
Dirhinus himalayanus
arrhenotoky
Chrysididae
Omalus djozanus hondonis
arrhenotoky
Cimbicidae
Abia candens
arrhenotoky
Cimbex femorata
arrhenotoky
Trichiosoma lucorum
arrhenotoky
Colletidae
Hylaeus ellipticus
arrhenotoky
Hylaeus nippon
arrhenotoky
Hylaeus sp.1
arrhenotoky
Hylaeus sp.2
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
12
6
333
34
17
333
20
10
333
20
10
333
20
10
333
20
10
333
34
17
333
26
13
333
20
10
333
20
10
333
16
8
333
20
10
333
34
17
333
8
4
333
12
6
333
40
20
331
18
9
331
48
24
331
6
3
333
10
5
333
10
5
333
10
5
333
38
19
332
32
16
331
16
8
331
16
8
331
22
11
331
16
8
332
36
18
332
56
28
332
235
Sex Chrom. Source
Karyotype
Taxa
Hylaeus stevensi
arrhenotoky
Hylaeus transversali
arrhenotoky
Cynipidae
Andricus curvator
arrhenotoky
Andricus fecundator
arrhenotoky
Andricus kollari
arrhenotoky
Andricus quercuscalicis
arrhenotoky
Aulacidea hieracii
arrhenotoky
Biorrhiza pallida
arrhenotoky
Cynips divisa
arrhenotoky
Cynips erinacea
arrhenotoky
Diastrophus nebulosus
arrhenotoky
Diplolepis elganteria
arrhenotoky
Diplolepis nervosum
arrhenotoky
Diplolepis rosae
arrhenotoky
Diplolepis spinosissimae
arrhenotoky
Dryocosmus kuriphilus
arrhenotoky
Neuroterus laeviusculus
arrhenotoky
Neuroterus numismalis
arrhenotoky
Neuroterus quescusbaccarum
arrhenotoky
Trigonaspis megaptera
arrhenotoky
Xestophanes potentillae
arrhenotoky
Diapriidae
Belyta depressa
arrhenotoky
Cinetus lanceolatus
arrhenotoky
Diprionidae
Diprion nipponicus
arrhenotoky
Diprion pini
arrhenotoky
Diprion similis
arrhenotoky
Gilpinia abieticola
arrhenotoky
Gilpinia frutetorum
arrhenotoky
Gilpinia hercyniae
arrhenotoky
Gilpinia pallida
arrhenotoky
Gilpinia polytoma
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
28
14
331
16
8
331
20
10
333
20
10
333
20
10
333
20
10
333
20
10
333
20
10
333
20
10
333
24
12
333
20
10
333
18
9
333
18
9
333
22
11
333
18
9
333
20
10
333
20
10
333
20
10
333
20
10
333
20
10
333
20
10
333
16
8
333
20
10
333
28
14
331
22
11
331
22
11
331
14
7
331
14
7
331
14
7
331
14
7
331
12
6
331
236
Sex Chrom. Source
Karyotype
Taxa
Macrodiprion nemoralis
arrhenotoky
Monoctenus nipponicus
arrhenotoky
Monoctenus suffusus
arrhenotoky
Neodiprion nanulus
arrhenotoky
Neodiprion sertifer
arrhenotoky
Neodiprion abietis
arrhenotoky
Neodiprion compar
arrhenotoky
Neodiprion dubiosus
arrhenotoky
Neodiprion lecontei
arrhenotoky
Neodiprion maurus
arrhenotoky
Neodiprion nigroscutum
arrhenotoky
Neodiprion pinetum
arrhenotoky
Neodiprion pratti banksianae
arrhenotoky
Neodiprion sp.1
arrhenotoky
Neodiprion sp.2
arrhenotoky
Neodiprion swainei
arrhenotoky
Neodiprion taedae taedae
arrhenotoky
Neodiprion tsugae
arrhenotoky
Neodiprion virginiana
arrhenotoky
Encryrtidae
Ageniaspis fuscicollis
arrhenotoky
Apoanagyrus lopezi
arrhenotoky
Copidosoma buyssoni
arrhenotoky
Copidosoma floridanum
arrhenotoky
Copidosoma gelechiae
arrhenotoky
Copidosoma truncatellum
arrhenotoky
Eucoilidae
Pseudeucoila bochei
arrhenotoky
Eulophidae
Cirrospillls dialllus
arrhenotoky
Colpodypells florus
arrhenotoky
Melittobia chalybii
arrhenotoky
Tetrastichus gigas
arrhenotoky
Tetrastichus megachilidis
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
14
7
331
30
15
331
14
7
331
14
7
331
14
7
331
16
8
331
14
7
331
14
7
331
14
7
331
14
7
331
14
7
331
14
7
331
14
7
331
14
7
331
16
8
331
16
8
331
14
7
331
14
7
331
14
7
331
20
10
333
20
10
333
24
12
333
20
10
333
22
11
333
20
10
333
20
10
333
12
6
333
12
6
333
10
5
333
12
6
333
12
6
333
237
Sex Chrom. Source
Karyotype
Taxa
Sexual
System
Eumenidae
Ancistrocerus adiabatus cytainus
arrhenotoky
Ancistrocerus densepilloserus
arrhenotoky
Ancistrocerus simulator
arrhenotoky
Ancistrocerus spilogaster
arrhenotoky
Ancistrocerus tuberculiceps sutterianus
arrhenotoky
Discoelius japonicus
arrhenotoky
Euodynerus foraminatus scutellaris
arrhenotoky
Euodynerus quadrifasciatus
arrhenotoky
Stenodynerus frauenfeldi
arrhenotoky
Eupelmidae
Anastatus catalonicus
arrhenotoky
Macroneura vesicularis
arrhenotoky
Eurytomidae
Eudecatoma biguttata
arrhenotoky
Eurytoma californica
arrhenotoky
Formicidae
Acanthomyrmex sp.1
arrhenotoky
Acanthomyrmex sp.2
arrhenotoky
Acanthomyrmex sp.3
arrhenotoky
Acanthomyrmex sp.4
arrhenotoky
Acromyrmex ambiguus
arrhenotoky
Acromyrmex crassipinus
arrhenotoky
Acromyrmex echinatior
arrhenotoky
Acromyrmex heyeri
arrhenotoky
Acromyrmex hispidus
arrhenotoky
Acromyrmex subterraneus
arrhenotoky
Acropyga acutiventris
arrhenotoky
Acropyga sp.1
arrhenotoky
Acropyga sp.2
arrhenotoky
Aenictus brevicornis
arrhenotoky
Aenictus laeviceps
arrhenotoky
Aenictus sp.1
arrhenotoky
Amblyopone australis
arrhenotoky
Amblyopone fortis
Ploidy
Female
2n
12
6
Male
2n
Sex Chrom. Source
Karyotype
331
12
6
332
14
7
331
12
6
331
20
10
331
24
12
332
16
8
331
10
5
332
20
10
332
10
5
333
10
5
333
18
9
333
20
10
333
24
12
361
22
11
361
22
11
361
22
11
361
38
19
361
38
19
361
36
18
362
38
19
361
38
19
361
38
19
361
30
15
361
32
16
361
30
15
361
24
12
361
22
11
361
30
15
361
48
24
361
44
22
361
238
Taxa
arrhenotoky
Amblyopone reclinata
arrhenotoky
Anochetus altisquamis
arrhenotoky
Anochetus graeffei
arrhenotoky
Anochetus graeffei.2
arrhenotoky
Anochetus horridus
arrhenotoky
Anochetus madaraszi
arrhenotoky
Anochetus modicus
arrhenotoky
Anochetus sp.1
arrhenotoky
Anochetus sp.2
arrhenotoky
Anochetus sp.3
arrhenotoky
Anochetus sp.4
arrhenotoky
Anochetus sp.5
arrhenotoky
Anochetus sp.6
arrhenotoky
Anochetus yerburyi
arrhenotoky
Anonychomyrma itinerans
arrhenotoky
Anonychomyrma sp.1
arrhenotoky
Anoplolepis gracilipes
arrhenotoky
Aphaenogaster beccarii
arrhenotoky
Aphaenogaster beccarii.2
arrhenotoky
Aphaenogaster depilis
arrhenotoky
Aphaenogaster famelica
arrhenotoky
Aphaenogaster fulva
arrhenotoky
Aphaenogaster gibbosa
arrhenotoky
Aphaenogaster iberica
arrhenotoky
Aphaenogaster lamellidens
arrhenotoky
Aphaenogaster longiceps
arrhenotoky
Aphaenogaster miamiana
arrhenotoky
Aphaenogaster osimensis
arrhenotoky
Aphaenogaster rudis
arrhenotoky
Aphaenogaster sardoa
arrhenotoky
Aphaenogaster senilis
arrhenotoky
Aphaenogaster smythiesi
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
38
19
361
30
15
361
30
15
361
38
19
361
46
23
361
28
14
361
30
15
361
24
12
361
38
19
361
30
15
361
30
15
361
34
17
361
34
17
361
30
15
361
16
8
361
16
8
361
34
17
361
30
15
361
46
23
361
34
17
361
34
17
361
36
18
361
32
16
361
34
17
361
38
19
361
46
23
361
36
18
361
32
16
361
40
20
361
34
17
361
32
16
361
28
14
361
239
Sex Chrom. Source
Karyotype
Taxa
Aphaenogaster sp.1
arrhenotoky
Aphaenogaster subterranea
arrhenotoky
Aphaenogaster testaceopilosa
arrhenotoky
Aphaenogaster tipuna
arrhenotoky
Aphaenogaster treatae
arrhenotoky
Apterostigma mayri
arrhenotoky
Apterostigma sp.1
arrhenotoky
Apterostigma sp.2
arrhenotoky
Apterostigma sp.3
arrhenotoky
Atta bisphaerica
arrhenotoky
Atta cephalotes
arrhenotoky
Atta columbica
arrhenotoky
Atta laevigata
arrhenotoky
Atta sexdens
arrhenotoky
Bothriomyrmex gibbus
arrhenotoky
Bothriomyrmex pusillus
arrhenotoky
Bothriomyrmex sp.1
arrhenotoky
Brachymyrmex sp.1
arrhenotoky
Calomyrmex sp, ANIC-1
arrhenotoky
Camponotus (Myrmanblys) sp1
arrhenotoky
Camponotus aethiops
arrhenotoky
Camponotus alii
arrhenotoky
Camponotus atriceps
arrhenotoky
Camponotus balzani
arrhenotoky
Camponotus bonariensis
arrhenotoky
Camponotus caryae
arrhenotoky
Camponotus cingulatus
arrhenotoky
Camponotus compressus
arrhenotoky
Camponotus consobrinus
arrhenotoky
Camponotus crassisquamis
arrhenotoky
Camponotus crassus
arrhenotoky
Camponotus cruentatus
arrhenotoky
Camponotus dolendus
Sexual
System
Ploidy
34
18
Female
2n
Male
2n
30
15
361
22
11
361
17
Sex Chrom. Source
Karyotype
361
34
17
361
42
21
361
24
12
361
20
10
361
24
12
361
32
16
361
22
11
361
44
22
362
22
11
361
22
11
361
22
11
361
22
11
361
22
11
361
22
11
361
18
9
361
28
14
361
9
361
42
21
361
42
21
361
40
20
361
40
20
361
40
20
361
40
20
361
40
20
361
30
15
361
46
23
361
40
20
361
20
10
361
38
19
361
20
10
361
240
Taxa
Sexual
System
arrhenotoky
Camponotus femoratus
arrhenotoky
Camponotus festinus
arrhenotoky
Camponotus foreli
arrhenotoky
Camponotus japonicus
arrhenotoky
Camponotus kiusiuensis
arrhenotoky
Camponotus lateralis
arrhenotoky
Camponotus ligniperdus
arrhenotoky
Camponotus mitis
arrhenotoky
Camponotus mus
arrhenotoky
Camponotus obscuripes
arrhenotoky
Camponotus parius
arrhenotoky
Camponotus pilicornis
arrhenotoky
Camponotus punctulatus
arrhenotoky
Camponotus rufipes
arrhenotoky
Camponotus rufoglaucus
arrhenotoky
Camponotus sericeiventris
arrhenotoky
Camponotus sericeus
arrhenotoky
Camponotus sp, ANIC-1
arrhenotoky
Camponotus sp, ANIC-10
arrhenotoky
Camponotus sp, ANIC-11
arrhenotoky
Camponotus sp, ANIC-12
arrhenotoky
Camponotus sp, ANIC-13
arrhenotoky
Camponotus sp, ANIC-14
arrhenotoky
Camponotus sp, ANIC-2
arrhenotoky
Camponotus sp, ANIC-3
arrhenotoky
Camponotus sp, ANIC-5
arrhenotoky
Camponotus sp, ANIC-8
arrhenotoky
Camponotus sp, ANIC-9
arrhenotoky
Camponotus sp. (impressus group)
arrhenotoky
Camponotus sp. variegatus complex
arrhenotoky
Camponotus sp.1
arrhenotoky
Camponotus sp.2
arrhenotoky
Ploidy
Female
2n
Male
2n
44
22
361
38
19
361
34
17
361
26
13
361
28
14
361
28
14
361
28
14
361
20
10
361
26
13
361
28
14
361
40
20
361
50
25
361
40
20
361
40
20
361
36
18
361
40
20
361
44
22
361
46
23
361
46
23
361
32
16
361
38
19
361
20
10
361
38
19
361
46
23
361
48
24
361
32
16
361
32
16
361
36
18
361
52
26
361
20
10
361
18
9
361
38
19
361
241
Sex Chrom. Source
Karyotype
Taxa
Sexual
System
Camponotus sp.3
arrhenotoky
Camponotus sp.4
arrhenotoky
Camponotus sp.5
arrhenotoky
Camponotus sp.6
arrhenotoky
Camponotus sylvaticus
arrhenotoky
Camponotus Taylori
arrhenotoky
Camponotus thraso
arrhenotoky
Camponotus vagus
arrhenotoky
Camponotus variegatus
arrhenotoky
Camponotus vitiosus
arrhenotoky
Cardiocondyla nuda
arrhenotoky
Cardiocondyla sp. (Myrmobrachys)
arrhenotoky
Cardiocondyla sp.1
arrhenotoky
Cardiocondyla sp.10
arrhenotoky
Cardiocondyla sp.11
arrhenotoky
Cardiocondyla sp.12
arrhenotoky
Cardiocondyla sp.13
arrhenotoky
Cardiocondyla sp.14
arrhenotoky
Cardiocondyla sp.15
arrhenotoky
Cardiocondyla sp.16
arrhenotoky
Cardiocondyla sp.17
arrhenotoky
Cardiocondyla sp.18
arrhenotoky
Cardiocondyla sp.19
arrhenotoky
Cardiocondyla sp.20
arrhenotoky
Cardiocondyla sp.21
arrhenotoky
Cardiocondyla sp.22
arrhenotoky
Cardiocondyla sp.7
arrhenotoky
Cardiocondyla sp.8
arrhenotoky
Cardiocondyla sp.9
arrhenotoky
Carebara asina
arrhenotoky
Carebara sauteri
arrhenotoky
Carebara sp, ANIC-6
arrhenotoky
Carebara sp.1
Ploidy
Female
2n
Male
2n
40
20
361
38
19
361
36
18
361
40
20
361
40
20
361
24
12
361
40
20
361
28
14
361
26
13
361
18
9
361
28
14
361
40
20
361
40
20
361
40
20
361
26
13
361
52
26
361
18
9
361
18
9
361
32
16
361
34
17
361
20
10
361
34
17
361
38
19
361
40
20
361
20
10
361
38
19
361
44
22
361
40
20
361
38
19
361
44
22
361
36
18
361
38
19
361
36
18
361
242
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Carebara sp.2
arrhenotoky
Carebara sp.3
arrhenotoky
Carebara sp.4
arrhenotoky
Carebara sp.5
arrhenotoky
Carebara sp.6
arrhenotoky
Carebara sp.7
arrhenotoky
Carebara sp.8
arrhenotoky
Cataglyphis bicolor
arrhenotoky
Cataglyphis iberica
arrhenotoky
Cataglyphis setipes
arrhenotoky
Centromyrmex feae
arrhenotoky
Cerapachys biroi
arrhenotoky
Cerapachys brevis
arrhenotoky
Cerapachys sp.1
arrhenotoky
Cerapachys sp.2
arrhenotoky
Chalepoxenus kutteri
arrhenotoky
Chalepoxenus muellerianus
arrhenotoky
Colobostruma alinodis
arrhenotoky
Colobostruma sp, ANIC-1
arrhenotoky
Colobostruma sp.1
arrhenotoky
Crematogaster biroi
arrhenotoky
Crematogaster brunnea
arrhenotoky
Crematogaster laboriosa
arrhenotoky
Crematogaster rothneyi
arrhenotoky
Crematogaster sp, ANIC-1
arrhenotoky
Crematogaster sp, ANIC-2
arrhenotoky
Crematogaster sp.1
arrhenotoky
Crematogaster sp.10
arrhenotoky
Crematogaster sp.11
arrhenotoky
Crematogaster sp.12
arrhenotoky
Crematogaster sp.13
arrhenotoky
Crematogaster sp.4
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
44
22
361
32
16
361
26
13
361
34
17
361
42
21
361
36
18
361
44
22
361
52
26
361
52
26
361
54
27
361
44
22
361
28
14
361
46
23
361
50
25
361
50
25
361
24
12
361
24
12
361
22
11
361
22
11
361
20
10
361
24
12
361
36
18
361
26
13
361
50
25
361
24
12
361
28
14
361
40
20
361
26
13
361
24
12
361
58
29
361
26
13
361
26
13
361
243
Sex Chrom. Source
Karyotype
Taxa
Crematogaster sp.5
arrhenotoky
Crematogaster sp.6
arrhenotoky
Crematogaster sp.7
arrhenotoky
Crematogaster sp.8
arrhenotoky
Crematogaster sp.9
arrhenotoky
Crematogaster subnuda
arrhenotoky
Cryptopone rotundiceps
arrhenotoky
Cryptopone sauteri
arrhenotoky
Cryptopone testacea
arrhenotoky
Cylindromyrmex brasiliensis
arrhenotoky
Cyphomyrmex cornutus
arrhenotoky
Cyphomyrmex costatus
arrhenotoky
Cyphomyrmex rimosus
arrhenotoky
Dacetinops concinnus
arrhenotoky
Diacamma rugosum
arrhenotoky
Diacamma sp.1
arrhenotoky
Diacamma sp.2
arrhenotoky
Diacamma sp.3
arrhenotoky
Diacamma sp.4
arrhenotoky
Diacamma sp.5
arrhenotoky
Dinoponera gigantea
arrhenotoky
Dinoponera lucida
arrhenotoky
Dinoponera quadriceps
arrhenotoky
Discothyrea sp.1
arrhenotoky
Doleromyrma sp.1
arrhenotoky
Doleromyrma sp.2
arrhenotoky
Dolichoderus quadripunctatus
arrhenotoky
Dolichoderus scabridus
arrhenotoky
Dolichoderus sp.1
arrhenotoky
Dolichoderus thoracicus
arrhenotoky
Dorymyrmex bicolor
arrhenotoky
Dorymyrmex flavus
arrhenotoky
Dorymyrmex pulchellus
Sexual
System
Ploidy
28
Female
2n
Male
2n
26
13
361
36
18
361
36
18
361
24
12
361
26
13
361
36
18
361
12
6
361
28
14
361
18
9
361
34
17
361
22
11
361
20
10
361
32
16
361
16
8
361
14
7
361
36
18
361
44
22
361
30
15
361
66
33
361
58
29
361
82
41
363
114
57
361
92
46
363
30
15
361
14
7
361
12
6
361
14
Sex Chrom. Source
Karyotype
361
28
14
361
18
9
361
30
15
361
26
13
361
26
13
361
18
9
361
244
Taxa
arrhenotoky
Dorymyrmex pyramicus
arrhenotoky
Dorymyrmex thoracicus
arrhenotoky
Echinopla sp.1
arrhenotoky
Ectatomma brunneum
arrhenotoky
Ectatomma edentatum
arrhenotoky
Ectatomma muticum
arrhenotoky
Ectatomma permagnum
arrhenotoky
Ectatomma tuberculatum
arrhenotoky
Eurhopalothrix sp.1
arrhenotoky
Forelius foetida
arrhenotoky
Forelius mccooki
arrhenotoky
Formica 3 spp (fusca gr.)
arrhenotoky
Formica 4 spp
arrhenotoky
Formica aquilonia
arrhenotoky
Formica candida
arrhenotoky
Formica cinerea
arrhenotoky
Formica cunicularia
arrhenotoky
Formica dakotensis
arrhenotoky
Formica exsecta
arrhenotoky
Formica frontalis
arrhenotoky
Formica fusca
arrhenotoky
Formica gagates
arrhenotoky
Formica gerardi
arrhenotoky
Formica japonica
arrhenotoky
Formica lemani
arrhenotoky
Formica lugubris
arrhenotoky
Formica montana
arrhenotoky
Formica obscuripes
arrhenotoky
Formica pergandei
arrhenotoky
Formica polyctena
arrhenotoky
Formica pratensis
arrhenotoky
Formica pressilabris
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
18
9
361
18
9
361
24
12
361
44
22
361
46
23
361
40
20
361
46
23
361
36
18
361
18
9
361
32
16
361
32
16
361
54
27
361
52
26
361
52
26
361
52
26
361
54
27
361
54
27
361
52
26
361
52
26
361
52
26
361
54
27
361
54
27
361
54
27
361
54
27
361
54
27
361
52
26
361
54
27
361
52
26
361
52
26
361
52
26
361
52
26
361
52
26
361
245
Sex Chrom. Source
Karyotype
Taxa
Formica reflexa
arrhenotoky
Formica rufa
arrhenotoky
Formica rufibarbis
arrhenotoky
Formica sanguinea
arrhenotoky
Formica subintegra
arrhenotoky
Formica subrufa
arrhenotoky
Formica truncorum
arrhenotoky
Formica ulkei
arrhenotoky
Formica uralensis
arrhenotoky
Formica yessensis
arrhenotoky
Formicoxenus chamberlini
arrhenotoky
Formicoxenus hirticolis
arrhenotoky
Formicoxenus nitidulus
arrhenotoky
Formicoxenus provancheri
arrhenotoky
Formicoxenus quebecensis
arrhenotoky
Gigantiops destructor
arrhenotoky
Gnamptogenys annulata
arrhenotoky
Gnamptogenys binghamii
arrhenotoky
Gnamptogenys menadensis
arrhenotoky
Gnamptogenys sp.1
arrhenotoky
Gnamptogenys sp.2
arrhenotoky
Gnamptogenys striatula
arrhenotoky
Harpagoxenus canadensis
arrhenotoky
Harpagoxenus sublaevis
arrhenotoky
Heteroponera dolo
arrhenotoky
Heteroponera relicta
arrhenotoky
Hypoponera confinis
arrhenotoky
Hypoponera pruinosa
arrhenotoky
Hypoponera sp, ANIC-1
arrhenotoky
Hypoponera sp, ANIC-2
arrhenotoky
Hypoponera sp.1
arrhenotoky
Hypoponera sp.2
arrhenotoky
Hypoponera sp.3
Sexual
System
Ploidy
Female
2n
Male
2n
52
26
361
52
26
361
54
27
361
52
26
361
52
26
361
52
26
361
54
27
361
52
26
361
52
26
361
52
26
361
28
14
361
32
16
361
30
15
361
22
11
361
28
14
361
78
39
361
68
34
361
22
11
361
42
21
361
36
18
361
46
23
361
34
17
361
36
18
361
40
20
361
24
12
361
22
11
361
38
19
361
24
12
361
38
19
361
38
19
361
38
19
361
38
19
361
36
18
361
246
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Iridomyrmex anceps
arrhenotoky
Iridomyrmex anceps.2
arrhenotoky
Iridomyrmex gracilis
arrhenotoky
Iridomyrmex mattiroloi
arrhenotoky
Iridomyrmex purpureus
arrhenotoky
Iridomyrmex sp, ANIC-11
arrhenotoky
Iridomyrmex sp, ANIC-12
arrhenotoky
Iridomyrmex sp, ANIC-13
arrhenotoky
Iridomyrmex sp, ANIC-14
arrhenotoky
Iridomyrmex sp, ANIC-15
arrhenotoky
Iridomyrmex sp, ANIC-16
arrhenotoky
Iridomyrmex sp, ANIC-17
arrhenotoky
Iridomyrmex sp, ANIC-5
arrhenotoky
Iridomyrmex sp, ANIC-6
arrhenotoky
Iridomyrmex sp.4
arrhenotoky
Lasius alienus
arrhenotoky
Lasius brunneus
arrhenotoky
Lasius emarginatus
arrhenotoky
Lasius flavus
arrhenotoky
Lasius fuliginosus
arrhenotoky
Lasius nearcticus
arrhenotoky
Lasius niger
arrhenotoky
Lasius pallitarsus
arrhenotoky
Lasius sakagamii
arrhenotoky
Lasius talpa
arrhenotoky
Lasius umbratus
arrhenotoky
Lepisiota capensis
arrhenotoky
Lepisiota sp.1
arrhenotoky
Lepisiota sp.2
arrhenotoky
Leptogenys borneensis
arrhenotoky
Leptogenys diminuta
arrhenotoky
Leptogenys hysterica
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
36
18
361
48
24
361
18
9
361
18
9
361
18
9
361
18
9
361
18
9
361
18
9
361
18
9
361
18
9
361
18
9
361
18
9
361
14
7
361
18
9
361
18
9
361
28
14
361
30
15
361
30
15
361
30
15
361
28
14
361
30
15
361
30
15
361
28
14
361
30
15
361
30
15
361
30
15
361
18
9
361
18
9
361
18
9
361
46
23
361
36
18
361
26
13
361
247
Sex Chrom. Source
Karyotype
Taxa
Leptogenys iridescens
arrhenotoky
Leptogenys kraepelini
arrhenotoky
Leptogenys minchinii
arrhenotoky
Leptogenys myops
arrhenotoky
Leptogenys peugueti
arrhenotoky
Leptogenys processionalis
arrhenotoky
Leptogenys sp.1
arrhenotoky
Leptogenys sp.2
arrhenotoky
Leptogenys sp.3
arrhenotoky
Leptomyrmex erytrocephalus
arrhenotoky
Leptothorax acervorum
arrhenotoky
Leptothorax albipennis
arrhenotoky
Leptothorax crassipilis
arrhenotoky
Leptothorax faberi
arrhenotoky
Leptothorax goesswaldi
arrhenotoky
Leptothorax gredleri
arrhenotoky
Leptothorax kutteri
arrhenotoky
Leptothorax muscorum
arrhenotoky
Leptothorax pacis
arrhenotoky
Leptothorax pocahontas
arrhenotoky
Leptothorax retractus
arrhenotoky
Leptothorax sp.1
arrhenotoky
Leptothorax sp.10
arrhenotoky
Leptothorax sp.2
arrhenotoky
Leptothorax sp.3
arrhenotoky
Leptothorax sp.4
arrhenotoky
Leptothorax sp.5
arrhenotoky
Leptothorax sp.6
arrhenotoky
Leptothorax sp.7
arrhenotoky
Leptothorax sp.8
arrhenotoky
Leptothorax sp.9
arrhenotoky
Leptothorax sphagnicola
arrhenotoky
Linepithema humile
Sexual
System
Ploidy
Female
2n
Male
2n
46
23
361
26
13
361
52
26
361
48
24
361
54
27
361
46
23
361
48
24
361
30
15
361
54
27
361
24
12
361
26
13
361
16
8
361
36
18
361
32
16
361
56
28
361
22
11
361
48
24
361
36
18
361
52
26
361
36
18
361
36
18
361
32
16
361
34
17
361
42
21
361
34
17
361
24
12
361
24
12
361
26
13
361
28
14
361
44
22
361
32
16
361
26
13
361
16
8
361
248
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Linepithema pilifer
arrhenotoky
Linepithema sp.1
arrhenotoky
Lophomyrmex bedoti
arrhenotoky
Lophomyrmex sp.1
arrhenotoky
Lordomyrma sp.1
arrhenotoky
Manica rubida
arrhenotoky
Mayriella abstinens
arrhenotoky
Meranoplus bicolor
arrhenotoky
Meranoplus minor
arrhenotoky
Meranoplus sp.1
arrhenotoky
Meranoplus sp.2
arrhenotoky
Meranoplus sp.3
arrhenotoky
Meranoplus sp.4
arrhenotoky
Messor aciculatus
arrhenotoky
Messor andrei
arrhenotoky
Messor barbarus
arrhenotoky
Messor sp.1
arrhenotoky
Monomorium dichroum
arrhenotoky
Monomorium glabrum
arrhenotoky
Monomorium indicum
arrhenotoky
Monomorium latinode
arrhenotoky
Monomorium minimum
arrhenotoky
Monomorium orientale
arrhenotoky
Monomorium pharaonis
arrhenotoky
Monomorium rothsteini
arrhenotoky
Monomorium scabriceps
arrhenotoky
Monomorium sp.1
arrhenotoky
Monomorium sp.10
arrhenotoky
Monomorium sp.11
arrhenotoky
Monomorium sp.12
arrhenotoky
Monomorium sp.2
arrhenotoky
Monomorium sp.3
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
18
9
361
18
9
361
38
19
361
38
19
361
22
11
361
44
22
361
18
9
361
16
8
361
22
11
361
20
10
361
22
11
361
22
11
361
22
11
361
44
22
361
40
20
361
42
21
361
40
20
361
16
8
361
38
19
361
22
11
361
70
35
361
22
11
361
20
10
361
22
11
361
22
11
361
38
19
361
22
11
361
38
19
361
22
11
361
22
11
361
42
21
361
32
16
361
249
Sex Chrom. Source
Karyotype
Taxa
Monomorium sp.4
arrhenotoky
Monomorium sp.5
arrhenotoky
Monomorium sp.6
arrhenotoky
Monomorium sp.7
arrhenotoky
Monomorium sp.8
arrhenotoky
Monomorium sp.9
arrhenotoky
Monomorium subopacum
arrhenotoky
Monomorium viride
arrhenotoky
Monomorium whitei
arrhenotoky
Mycocepurus goeldii
arrhenotoky
Mycocepurus sp.1
arrhenotoky
Myrmecia banksi
arrhenotoky
Myrmecia brevinoda
arrhenotoky
Myrmecia cephalotes
arrhenotoky
Myrmecia chasei
arrhenotoky
Myrmecia croslandi
arrhenotoky
Myrmecia forficata
arrhenotoky
Myrmecia fulvipes
arrhenotoky
Myrmecia gulosa
arrhenotoky
Myrmecia haskinsorum
arrhenotoky
Myrmecia imaii
arrhenotoky
Myrmecia mandibularis
arrhenotoky
Myrmecia michaelseni
arrhenotoky
Myrmecia nigrocincta
arrhenotoky
Myrmecia occidentalis
arrhenotoky
Myrmecia pavida
arrhenotoky
Myrmecia piliventris
arrhenotoky
Myrmecia pilosula
arrhenotoky
Myrmecia pyriformis
arrhenotoky
Myrmecia simillima
arrhenotoky
Myrmecia sp. cf. arnoldi
arrhenotoky
Myrmecia sp. cf. fulvipes
arrhenotoky
Myrmecia tepperi
Sexual
System
Ploidy
Female
2n
Male
2n
22
11
361
22
11
361
22
11
361
22
11
361
22
11
361
34
17
361
34
17
361
22
11
361
24
12
361
16
8
361
16
8
361
40
20
361
84
42
361
66
33
361
46
23
361
2
1
361
52
26
361
42
21
361
38
19
361
18
9
361
8
4
361
58
29
361
54
27
361
22
11
361
64
32
361
44
22
361
34
17
361
20
10
361
82
41
361
70
35
361
58
29
361
12
6
361
70
35
361
250
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Myrmecia vindex
arrhenotoky
Myrmecina americana
arrhenotoky
Myrmecina graminicola
arrhenotoky
Myrmecina sp.1
arrhenotoky
Myrmecina sp.2
arrhenotoky
Myrmica lobicornis
arrhenotoky
Myrmica rubra
arrhenotoky
Myrmica ruginodis
arrhenotoky
Myrmica sabuleti
arrhenotoky
Myrmica scabrinodis
arrhenotoky
Myrmica schencki
arrhenotoky
Myrmica sulcinodis
arrhenotoky
Myrmicaria brunnea
arrhenotoky
Myrmicaria sp.1
arrhenotoky
Myrmicaria sp.2
arrhenotoky
Myrmicaria sp.3
arrhenotoky
Myrmicaria sp.4
arrhenotoky
Myrmicaria sp.5
arrhenotoky
Myrmicocrypta sp.1
arrhenotoky
Myrmoxenus adlerzi
arrhenotoky
Myrmoxenus algeriana
arrhenotoky
Myrmoxenus bernardi
arrhenotoky
Myrmoxenus corsica
arrhenotoky
Myrmoxenus gordiagini
arrhenotoky
Myrmoxenus kraussei
arrhenotoky
Myrmoxenus ravouxi
arrhenotoky
Myrmoxenus stumperi
arrhenotoky
Mystrium camillae
arrhenotoky
Neivamyrmex texanus
arrhenotoky
Nothomyrmecia macrops
arrhenotoky
Notoncus ectatommoides
arrhenotoky
Ochetellus glaber
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
76
38
361
28
14
361
28
14
361
66
33
361
68
34
361
24
12
361
48
24
361
48
24
361
46
23
361
44
22
361
46
23
361
52
26
361
44
22
361
44
22
361
44
22
361
44
22
361
44
22
361
46
23
361
30
15
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
32
16
361
36
18
361
94
47
361
44
22
361
28
14
361
251
Sex Chrom. Source
Karyotype
Taxa
Odontomachus chelifer
arrhenotoky
Odontomachus hastatus
arrhenotoky
Odontomachus latidens
arrhenotoky
Odontomachus meinerti
arrhenotoky
Odontomachus rixosus
arrhenotoky
Odontomachus scalptus
arrhenotoky
Odontomachus simillimus
arrhenotoky
Odontomachus sp, ANIC-1
arrhenotoky
Odontomachus sp.1
arrhenotoky
Odontoponera transversa
arrhenotoky
Oecophylla longinoda
arrhenotoky
Oecophylla smaragdina
arrhenotoky
Opisthopsis rufithorax
arrhenotoky
Orectognathus clarki
arrhenotoky
Orectognathus darlingtoni
arrhenotoky
Orectognathus versicolor
arrhenotoky
Pachycondyla apicalis
arrhenotoky
Pachycondyla arhuaca
arrhenotoky
Pachycondyla astuta
arrhenotoky
Pachycondyla carinulata
arrhenotoky
Pachycondyla chinensis
arrhenotoky
Pachycondyla concava
arrhenotoky
Pachycondyla constricta
arrhenotoky
Pachycondyla crassinoda
arrhenotoky
Pachycondyla crenata
arrhenotoky
Pachycondyla curvinodis
arrhenotoky
Pachycondyla gilberti
arrhenotoky
Pachycondyla goeldii
arrhenotoky
Pachycondyla harpax
arrhenotoky
Pachycondyla impressa
arrhenotoky
Pachycondyla inversa
arrhenotoky
Pachycondyla leeuwenhoeki
arrhenotoky
Pachycondyla lutea
Sexual
System
Ploidy
Female
2n
Male
2n
44
22
361
44
22
361
32
16
361
44
22
361
30
15
361
44
22
361
44
22
361
44
22
361
44
22
361
44
22
361
24
12
361
16
8
361
50
25
361
30
15
361
22
11
361
22
11
361
40
20
361
24
12
361
40
20
361
24
12
361
22
11
361
54
27
361
30
15
361
62
31
361
26
13
361
26
13
361
30
15
361
24
12
361
96
48
361
94
47
361
30
15
361
16
8
361
16
8
361
252
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Pachycondyla luteipes
arrhenotoky
Pachycondyla marginata
arrhenotoky
Pachycondyla mesonotalis
arrhenotoky
Pachycondyla metanotalis
arrhenotoky
Pachycondyla moesta
arrhenotoky
Pachycondyla rubiginosa
arrhenotoky
Pachycondyla rubra
arrhenotoky
Pachycondyla rufipes
arrhenotoky
Pachycondyla sp.1
arrhenotoky
Pachycondyla sp.10
arrhenotoky
Pachycondyla sp.11
arrhenotoky
Pachycondyla sp.2
arrhenotoky
Pachycondyla sp.3
arrhenotoky
Pachycondyla sp.4
arrhenotoky
Pachycondyla sp.5
arrhenotoky
Pachycondyla sp.6
arrhenotoky
Pachycondyla sp.7
arrhenotoky
Pachycondyla sp.8
arrhenotoky
Pachycondyla sp.9
arrhenotoky
Pachycondyla stigma
arrhenotoky
Pachycondyla striata
arrhenotoky
Pachycondyla subversa
arrhenotoky
Pachycondyla succedanea
arrhenotoky
Pachycondyla tridentata
arrhenotoky
Pachycondyla unidentata
arrhenotoky
Pachycondyla venusta
arrhenotoky
Pachycondyla verenae
arrhenotoky
Pachycondyla villosa
arrhenotoky
Papyrius nitidus
arrhenotoky
Paratrechina indica
arrhenotoky
Paratrechina longicornis
arrhenotoky
Paratrechina parvula
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
22
11
361
46
23
361
26
13
361
70
35
361
26
13
361
76
38
361
36
18
361
48
24
361
22
11
361
48
24
361
52
26
361
38
19
361
44
22
361
60
30
361
22
11
361
28
14
361
22
11
361
36
18
361
36
18
361
12
6
361
104
52
361
28
14
361
14
7
361
28
14
361
12
6
361
48
24
361
60
30
361
34
17
361
16
8
361
30
15
361
16
8
361
14
7
361
253
Sex Chrom. Source
Karyotype
Taxa
Paratrechina sp, ANIC-1
arrhenotoky
Paratrechina sp.1
arrhenotoky
Paratrechina sp.10
arrhenotoky
Paratrechina sp.2
arrhenotoky
Paratrechina sp.3
arrhenotoky
Paratrechina sp.4
arrhenotoky
Paratrechina sp.5
arrhenotoky
Paratrechina sp.6
arrhenotoky
Paratrechina sp.7
arrhenotoky
Paratrechina sp.8
arrhenotoky
Paratrechina sp.9
arrhenotoky
Pheidole binghamii
arrhenotoky
Pheidole capellinii
arrhenotoky
Pheidole dentata
arrhenotoky
Pheidole dentigula
arrhenotoky
Pheidole desertorum
arrhenotoky
Pheidole fallax
arrhenotoky
Pheidole fervida
arrhenotoky
Pheidole hortensis
arrhenotoky
Pheidole hyatti
arrhenotoky
Pheidole indica
arrhenotoky
Pheidole latinoda
arrhenotoky
Pheidole megacephala
arrhenotoky
Pheidole mus
arrhenotoky
Pheidole nitidula
arrhenotoky
Pheidole noda
arrhenotoky
Pheidole pallidula
arrhenotoky
Pheidole plagiaria
arrhenotoky
Pheidole porcula
arrhenotoky
Pheidole soritis
arrhenotoky
Pheidole sp.1
arrhenotoky
Pheidole sp.10
arrhenotoky
Pheidole sp.11
Sexual
System
Ploidy
Female
2n
Male
2n
30
15
361
16
8
361
30
15
361
26
13
361
28
14
361
16
8
361
28
14
361
30
15
361
16
8
361
30
15
361
30
15
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
42
21
361
20
10
361
12
6
361
20
10
361
38
19
361
20
10
361
20
10
361
20
10
361
18
9
361
18
9
361
20
10
361
20
10
361
254
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Pheidole sp.12
arrhenotoky
Pheidole sp.13
arrhenotoky
Pheidole sp.14
arrhenotoky
Pheidole sp.15
arrhenotoky
Pheidole sp.16
arrhenotoky
Pheidole sp.17
arrhenotoky
Pheidole sp.18
arrhenotoky
Pheidole sp.19
arrhenotoky
Pheidole sp.2
arrhenotoky
Pheidole sp.20
arrhenotoky
Pheidole sp.21
arrhenotoky
Pheidole sp.22
arrhenotoky
Pheidole sp.23
arrhenotoky
Pheidole sp.24
arrhenotoky
Pheidole sp.25
arrhenotoky
Pheidole sp.26
arrhenotoky
Pheidole sp.27
arrhenotoky
Pheidole sp.28
arrhenotoky
Pheidole sp.29
arrhenotoky
Pheidole sp.3
arrhenotoky
Pheidole sp.30
arrhenotoky
Pheidole sp.31
arrhenotoky
Pheidole sp.32
arrhenotoky
Pheidole sp.33
arrhenotoky
Pheidole sp.34
arrhenotoky
Pheidole sp.35
arrhenotoky
Pheidole sp.36
arrhenotoky
Pheidole sp.37
arrhenotoky
Pheidole sp.38
arrhenotoky
Pheidole sp.39
arrhenotoky
Pheidole sp.4
arrhenotoky
Pheidole sp.40
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
20
10
361
34
17
361
20
10
361
16
8
361
38
19
361
18
9
361
20
10
361
22
11
361
20
10
361
38
19
361
20
10
361
20
10
361
20
10
361
18
9
361
20
10
361
20
10
361
20
10
361
36
18
361
42
21
361
20
10
361
20
10
361
30
15
361
20
10
361
20
10
361
255
Sex Chrom. Source
Karyotype
Taxa
Pheidole sp.41
arrhenotoky
Pheidole sp.42
arrhenotoky
Pheidole sp.43
arrhenotoky
Pheidole sp.44
arrhenotoky
Pheidole sp.45
arrhenotoky
Pheidole sp.46
arrhenotoky
Pheidole sp.47
arrhenotoky
Pheidole sp.48
arrhenotoky
Pheidole sp.49
arrhenotoky
Pheidole sp.5
arrhenotoky
Pheidole sp.50
arrhenotoky
Pheidole sp.51
arrhenotoky
Pheidole sp.6
arrhenotoky
Pheidole sp.7
arrhenotoky
Pheidole sp.8
arrhenotoky
Pheidole sp.9
arrhenotoky
Pheidole spininodis
arrhenotoky
Pheidole subarmata
arrhenotoky
Pheidole tepicana
arrhenotoky
Pheidole woodmasoni
arrhenotoky
Pheidologeton diversus
arrhenotoky
Pheidologeton sp.1
arrhenotoky
Philidris cordata
arrhenotoky
Plagiolepis pygmaea
arrhenotoky
Plagiolepis schmitzii
arrhenotoky
Plagiolepis sp.1
arrhenotoky
Plagiolepis sp.2
arrhenotoky
Platythyrea quadridenta
arrhenotoky
Platythyrea tricuspidata
arrhenotoky
Podomyrma adelaidae
arrhenotoky
Pogonomyrmex apache
arrhenotoky
Pogonomyrmex badius
arrhenotoky
Pogonomyrmex barbatus
Sexual
System
Ploidy
Female
2n
Male
2n
20
10
361
20
10
361
28
14
361
18
9
361
18
9
361
18
9
361
32
16
361
20
10
361
18
9
361
20
10
361
18
9
361
18
9
361
20
10
361
18
9
361
20
10
361
20
10
361
20
10
361
20
10
361
18
9
361
18
9
361
42
21
361
42
21
361
16
8
361
18
9
361
18
9
361
18
9
361
18
9
361
18
9
361
94
47
361
52
26
361
32
16
361
32
16
361
32
16
361
256
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Pogonomyrmex brevispinosus
arrhenotoky
Pogonomyrmex californicus
arrhenotoky
Pogonomyrmex comanche
arrhenotoky
Pogonomyrmex desertorum
arrhenotoky
Pogonomyrmex huachucanus
arrhenotoky
Pogonomyrmex imberbiculus
arrhenotoky
Pogonomyrmex magnacanthus
arrhenotoky
Pogonomyrmex maricopa
arrhenotoky
Pogonomyrmex montanus
arrhenotoky
Pogonomyrmex occidentalis
arrhenotoky
Pogonomyrmex rugosus
arrhenotoky
Pogonomyrmex subnitidus
arrhenotoky
Polyergus samurai
arrhenotoky
Polyrhachis ammon
arrhenotoky
Polyrhachis dives
arrhenotoky
Polyrhachis gribodoi
arrhenotoky
Polyrhachis hector
arrhenotoky
Polyrhachis hippomanes
arrhenotoky
Polyrhachis illaudata
arrhenotoky
Polyrhachis lacteipennis
arrhenotoky
Polyrhachis lamellidens
arrhenotoky
Polyrhachis rastellata
arrhenotoky
Polyrhachis sp, ANIC-1
arrhenotoky
Polyrhachis sp.1
arrhenotoky
Polyrhachis sp.2
arrhenotoky
Polyrhachis sp.3
arrhenotoky
Ponera japonica
arrhenotoky
Ponera pennsylvanica
arrhenotoky
Ponera scabra
arrhenotoky
Ponera sp.1
arrhenotoky
Ponera sp.2
arrhenotoky
Prenolepis imparis
arrhenotoky
Sexual
System
Ploidy
32
Female
2n
Male
2n
32
16
361
32
16
361
32
16
361
32
16
361
36
18
361
30
15
361
16
Sex Chrom. Source
Karyotype
361
32
16
361
32
16
361
32
16
361
32
16
361
32
16
361
54
27
361
42
21
361
42
21
361
48
24
361
42
21
361
40
20
361
32
16
361
42
21
361
42
21
361
42
21
361
42
21
361
42
21
361
42
21
361
40
20
361
12
6
361
12
6
361
8
4
361
12
6
361
12
6
361
16
8
361
257
Taxa
Prenolepis jerdoni
arrhenotoky
Pristomyrmex punctatus
arrhenotoky
Pristomyrmex sp.1
arrhenotoky
Pristomyrmex sp.2
arrhenotoky
Proatta sp.1
arrhenotoky
Probolomyrmex sp.1
arrhenotoky
Proceratium silaceum
arrhenotoky
Proceratium sp.1
arrhenotoky
Prolasius sp, ANIC-1
arrhenotoky
Prolasius sp, ANIC-2
arrhenotoky
Protomognathus americanus
arrhenotoky
Pseudolasius sp.1
arrhenotoky
Pseudolasius sp.2
arrhenotoky
Pseudolasius sp.3
arrhenotoky
Pseudolasius sp.4
arrhenotoky
Pseudolasius sp.5
arrhenotoky
Pseudomyrmex gracilis
arrhenotoky
Pseudomyrmex holmgreni
arrhenotoky
Pseudomyrmex penetrator
arrhenotoky
Pseudomyrmex schuppi
arrhenotoky
Pseudomyrmex sp.1
arrhenotoky
Pseudomyrmex sp.2
arrhenotoky
Pseudomyrmex sp.3
arrhenotoky
Pyramica dohertyi
arrhenotoky
Pyramica mutica
arrhenotoky
Pyramica sp.1
arrhenotoky
Pyramica sp.2
arrhenotoky
Pyramica sp.3
arrhenotoky
Recurvidris sp.1
arrhenotoky
Rhytidoponera aciculata
arrhenotoky
Rhytidoponera chalybaea
arrhenotoky
Rhytidoponera impressa
arrhenotoky
Rhytidoponera lamellinodis
Sexual
System
Ploidy
Female
2n
Male
2n
36
18
361
24
12
361
22
11
361
28
14
361
32
16
361
28
14
361
36
18
361
48
24
361
18
9
361
18
9
361
22
11
361
28
14
361
30
15
361
30
15
361
16
8
361
34
17
361
70
35
361
50
25
361
24
12
361
24
12
361
42
21
361
50
25
361
44
22
361
24
12
361
36
18
361
38
19
361
26
13
361
16
8
361
24
12
361
52
26
361
42
21
361
42
21
361
52
26
361
258
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Rhytidoponera maniae
arrhenotoky
Rhytidoponera mayri
arrhenotoky
Rhytidoponera mayri
arrhenotoky
Rhytidoponera metallica
arrhenotoky
Rhytidoponera punctata
arrhenotoky
Rhytidoponera purpurea
arrhenotoky
Rhytidoponera sp ANIC-10
arrhenotoky
Rhytidoponera sp ANIC-11
arrhenotoky
Rhytidoponera sp ANIC-13
arrhenotoky
Rhytidoponera sp ANIC-14
arrhenotoky
Rhytidoponera sp ANIC-15
arrhenotoky
Rhytidoponera sp ANIC-16
arrhenotoky
Rhytidoponera sp ANIC-9
arrhenotoky
Rhytidoponera tasmaniensis-1
arrhenotoky
Rhytidoponera tasmaniensis-2
arrhenotoky
Rhytidoponera victoriae
arrhenotoky
Sericomyrmex amabilis
arrhenotoky
Solenopsis aurea
arrhenotoky
Solenopsis fugax
arrhenotoky
Solenopsis geminata
arrhenotoky
Solenopsis invicta
arrhenotoky
Solenopsis molesta
arrhenotoky
Solenopsis richteri
arrhenotoky
Solenopsis saevissima
arrhenotoky
Solenopsis sp.1
arrhenotoky
Solenopsis sp.2
arrhenotoky
Solenopsis xyloni
arrhenotoky
Sphinctomyrmex steinheili
arrhenotoky
Stenamma brevicorne
arrhenotoky
Stenamma westwoodii
arrhenotoky
Stigmacros sp, ANIC-1
arrhenotoky
Stigmacros sp, ANIC-3
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
44
22
361
50
25
361
50
25
361
38
19
361
100
50
361
38
19
361
48
24
361
50
25
361
52
26
361
22
11
361
50
25
361
52
26
361
48
24
361
30
15
361
46
23
361
42
21
361
50
25
361
32
16
361
22
11
361
32
16
361
32
16
361
22
11
361
32
16
361
32
16
361
38
19
361
22
11
361
32
16
361
46
23
361
8
4
361
40
20
361
38
19
361
20
10
361
259
Taxa
Strongylognathus huberi
arrhenotoky
Strumigenys doriae
arrhenotoky
Strumigenys friedae
arrhenotoky
Strumigenys godeffroyi
arrhenotoky
Tapinoma erraticum
arrhenotoky
Tapinoma indicum
arrhenotoky
Tapinoma melanocephalum
arrhenotoky
Tapinoma nigerrimum
arrhenotoky
Tapinoma sessile
arrhenotoky
Tapinoma simrothi
arrhenotoky
Tapinoma sp.1
arrhenotoky
Technomyrmex albipes
arrhenotoky
Technomyrmex sp.1
arrhenotoky
Technomyrmex sp.2
arrhenotoky
Technomyrmex sp.3
arrhenotoky
Technomyrmex sp2 bicolor
arrhenotoky
Temnothorax affinis
arrhenotoky
Temnothorax ambiguus
arrhenotoky
Temnothorax andrei
arrhenotoky
Temnothorax angustulus
arrhenotoky
Temnothorax c.f. interrupus
arrhenotoky
Temnothorax c.f. lichtensteini
arrhenotoky
Temnothorax congruus
arrhenotoky
Temnothorax corticalis
arrhenotoky
Temnothorax curvispinosus
arrhenotoky
Temnothorax duloticus
arrhenotoky
Temnothorax exilis
arrhenotoky
Temnothorax flavicornis
arrhenotoky
Temnothorax gredosi
arrhenotoky
Temnothorax interruptus
arrhenotoky
Temnothorax kraussei
arrhenotoky
Temnothorax lichtensteini
arrhenotoky
Temnothorax longispinosus
Sexual
System
Ploidy
28
Female
2n
Male
2n
28
14
361
22
11
361
24
12
361
42
21
361
16
8
361
12
6
361
10
5
361
18
9
361
16
8
361
18
9
361
10
5
361
18
9
361
30
15
361
28
14
361
30
15
361
28
14
361
18
9
361
22
11
361
20
10
361
46
23
361
24
12
361
14
Sex Chrom. Source
Karyotype
361
18
9
361
22
11
361
46
23
361
48
24
361
22
11
361
22
11
361
34
17
361
24
12
361
28
14
361
28
14
361
24
12
361
260
Taxa
arrhenotoky
Temnothorax melas
arrhenotoky
Temnothorax niger
arrhenotoky
Temnothorax nigriceps
arrhenotoky
Temnothorax nylanderi
arrhenotoky
Temnothorax parvulus
arrhenotoky
Temnothorax rabaudi
arrhenotoky
Temnothorax racovitzai
arrhenotoky
Temnothorax recedens
arrhenotoky
Temnothorax rottenbergii
arrhenotoky
Temnothorax rugatulus
arrhenotoky
Temnothorax schaumii
arrhenotoky
Temnothorax sordidulus
arrhenotoky
Temnothorax specularis
arrhenotoky
Temnothorax spinosior
arrhenotoky
Temnothorax spinosius
arrhenotoky
Temnothorax ssp tuberum group
arrhenotoky
Temnothorax tristis
arrhenotoky
Temnothorax tuberum
arrhenotoky
Temnothorax unifasciatus
arrhenotoky
Tetramorium adelphon
arrhenotoky
Tetramorium brevidentatum
arrhenotoky
Tetramorium caespitum
arrhenotoky
Tetramorium eleates
arrhenotoky
Tetramorium forte
arrhenotoky
Tetramorium guineense
arrhenotoky
Tetramorium kheperra
arrhenotoky
Tetramorium lanuginosum
arrhenotoky
Tetramorium pacificum
arrhenotoky
Tetramorium pnyxis
arrhenotoky
Tetramorium semilaeve
arrhenotoky
Tetramorium seneb
arrhenotoky
Tetramorium simillimum
arrhenotoky
Sexual
System
Ploidy
24
Female
2n
Male
2n
26
13
361
36
18
361
18
9
361
22
11
361
28
14
361
18
9
361
42
21
361
24
12
361
22
11
361
28
14
361
18
9
361
22
11
361
34
17
361
24
12
361
32
16
361
12
Sex Chrom. Source
Karyotype
361
42
21
361
18
9
361
18
9
361
22
11
361
20
10
361
28
14
361
28
14
361
28
14
361
22
11
361
14
7
361
14
7
361
22
11
361
20
10
361
14
7
361
20
10
361
14
7
361
261
Taxa
Tetramorium smithi
arrhenotoky
Tetramorium sp.1
arrhenotoky
Tetramorium sp.10
arrhenotoky
Tetramorium sp.11
arrhenotoky
Tetramorium sp.12
arrhenotoky
Tetramorium sp.2
arrhenotoky
Tetramorium sp.3
arrhenotoky
Tetramorium sp.4
arrhenotoky
Tetramorium sp.5
arrhenotoky
Tetramorium sp.6
arrhenotoky
Tetramorium sp.7
arrhenotoky
Tetramorium sp.8
arrhenotoky
Tetramorium sp.9
arrhenotoky
Tetramorium spinosum
arrhenotoky
Tetramorium walshi
arrhenotoky
Tetraponera allaborans
arrhenotoky
Tetraponera sp.1
arrhenotoky
Tetraponera sp.2
arrhenotoky
Trachymyrmex septentrionales
arrhenotoky
Trachymyrmex sp.1
arrhenotoky
Trachymyrmex sp.2
arrhenotoky
Typhlomyrmex meire
arrhenotoky
Typhlomyrmex rogenhoferi
arrhenotoky
Vollenhovia emeryii
arrhenotoky
Vollenhovia sp, ANIC-3
arrhenotoky
Vollenhovia sp.1
arrhenotoky
Vollenhovia sp.2
arrhenotoky
Vollenhovia sp.3
arrhenotoky
Vollenhovia sp.4
arrhenotoky
Halictidae
Agapostemon virescens
arrhenotoky
Augochlora pura
arrhenotoky
Augochlorella michaelis
arrhenotoky
Sexual
System
Ploidy
20
Female
2n
Male
2n
26
13
361
20
10
361
36
18
361
20
10
361
24
12
361
18
9
361
20
10
361
18
9
361
20
10
361
22
11
361
18
9
361
26
13
361
14
7
361
26
13
361
14
7
361
32
16
361
44
22
361
42
21
361
10
Sex Chrom. Source
Karyotype
361
12
6
361
18
9
361
20
10
361
36
18
361
36
18
361
40
20
361
22
11
361
50
25
361
34
17
361
36
18
361
34
17
331
22
11
331
32
16
351
262
Taxa
Augochloropsis sparsilis
arrhenotoky
Halictus aerarius
arrhenotoky
Lasioglossum allodalum
arrhenotoky
Lasioglossum cooleyi
arrhenotoky
Lasioglossum duplex
arrhenotoky
Lasioglossum lineatulus
arrhenotoky
Lasioglossum rhythidophorum
arrhenotoky
Lasioglossum rhytidophorum
arrhenotoky
Lasioglossum taeniolellum
arrhenotoky
Lasioglossum zephyrus
arrhenotoky
Nomia nevadensis angelesia
arrhenotoky
Pseudaugochloropsis graminea
arrhenotoky
Pseudoaugochloropsis graminea
arrhenotoky
Ichneumonidae
Aethercerus discolor
arrhenotoky
Aethercerus dispar
arrhenotoky
Aethercerus nitidus
arrhenotoky
Aethercerus ranini
arrhenotoky
Agrothereutes extrematus
arrhenotoky
Aoplus pulchricornis
arrhenotoky
Aptesis puncticollis
arrhenotoky
Baeosemus dentifer
arrhenotoky
Baranisobas ridibundus
arrhenotoky
Chasmias motatorius
arrhenotoky
Coelichneumon cyaniventris
arrhenotoky
Coelichneumon sugillatorius
arrhenotoky
Colpognathus celerator
arrhenotoky
Cratichneumon viator
arrhenotoky
Diadromus prosopius
arrhenotoky
Diadromus pulchellus
arrhenotoky
Diadromus subtilicornis
arrhenotoky
Diadromus troglodytes
arrhenotoky
Diadromus varicolor
arrhenotoky
Sexual
System
Ploidy
12
Female
2n
Male
2n
16
8
331
28
14
332
40
20
332
36
18
364
18
9
332
24
12
331
6
Sex Chrom. Source
Karyotype
351
12
6
331
32
16
332
26
13
331
42
21
331
16
8
331
16
8
351
22
11
333
24
12
333
22
11
333
22
11
333
20
10
333
26
13
333
16
8
333
22
11
333
22
11
333
34
17
333
26
13
333
26
13
333
22
11
333
28
14
333
22
11
333
22
11
333
22
11
333
22
11
333
22
11
333
263
Taxa
Dicaelotus pumilis
arrhenotoky
Dicaelotus sp. nr. Parvulus
arrhenotoky
Diphyus latebricola
arrhenotoky
Diphyus raptorius
arrhenotoky
Dirophanes callopus
arrhenotoky
Dirophanes fulvitarsis
arrhenotoky
Dirophanes invisor
arrhenotoky
Dyspetes arrogator
arrhenotoky
Ephialtes manifestator
arrhenotoky
Eurylabus torvus
arrhenotoky
Gelis sp.1
arrhenotoky
Gen. sp.1
arrhenotoky
Glypta lapponica
arrhenotoky
Heterischnus nigricollis
arrhenotoky
Heterischnus truncator
arrhenotoky
Homotherus locutor
arrhenotoky
Ichneumon albiger
arrhenotoky
Ichneumon amphibolus
arrhenotoky
Ichneumon bucculentus
arrhenotoky
Ichneumon confusor
arrhenotoky
Ichneumon crassifemur
arrhenotoky
Ichneumon croceipes
arrhenotoky
Ichneumon extensorius
arrhenotoky
Ichneumon formosus
arrhenotoky
Ichneumon gracilentus
arrhenotoky
Ichneumon gracilicornis
arrhenotoky
Ichneumon ingratus
arrhenotoky
Ichneumon inquinatus
arrhenotoky
Ichneumon insidiosus
arrhenotoky
Ichneumon lugens
arrhenotoky
Ichneumon melanotis
arrhenotoky
Ichneumon minutorius
arrhenotoky
Ichneumon molitorius
Sexual
System
Ploidy
Female
2n
Male
2n
22
11
333
22
11
333
24
12
333
24
12
333
18
9
333
20
10
333
20
10
333
20
10
333
30
15
333
20
10
333
26
13
333
30
15
332
18
9
333
22
11
333
22
11
333
22
11
333
24
12
333
24
12
333
24
12
333
24
12
333
24
12
333
24
12
333
24
12
333
22
11
333
24
12
333
22
11
333
24
12
333
26
13
333
24
12
333
24
12
333
24
12
333
24
12
333
24
12
333
264
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Ichneumon nereni
arrhenotoky
Ichneumon sarcitorius
arrhenotoky
Ichneumon stramentarius
arrhenotoky
Ichneumon submarginatus
arrhenotoky
Ichneumon suspiciosus
arrhenotoky
Ichneumon validicornis
arrhenotoky
Lissonota sp.
arrhenotoky
Mastrus smithii
arrhenotoky
Oronotus binotatus
arrhenotoky
Orthocentrus sp.1
arrhenotoky
Orthopelma mediator
arrhenotoky
Patrocloides chalybeatus
arrhenotoky
Phaeogenes melanogonos
arrhenotoky
Phaeogenes nigridens
arrhenotoky
Phaeogenes semivulpinus
arrhenotoky
Phaeogenes spiniger
arrhenotoky
Polysphincta tuberosa
arrhenotoky
Pseudoamblyteles homocerus
arrhenotoky
Stenichneumon culpator
arrhenotoky
Sycaonia sicaria
arrhenotoky
Syspasis alboguttata
arrhenotoky
Syspasis scutellator
arrhenotoky
Tycherus australogeminus
arrhenotoky
Tycherus bellicornis
arrhenotoky
Tycherus dilleri
arrhenotoky
Tycherus fuscicornis
arrhenotoky
Tycherus ischiontelinus
arrhenotoky
Tycherus ophthalmicus
arrhenotoky
Tycherus osculator
arrhenotoky
Tycherus suspicax
arrhenotoky
Venturia canescens
arrhenotoky
Virgichneumon digrammus
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
22
11
333
24
12
333
20
10
333
24
12
333
24
12
333
24
12
333
22
11
333
26
13
333
22
11
333
28
14
333
22
11
333
16
8
333
22
11
333
22
11
333
18
9
333
22
11
333
18
9
333
18
9
333
28
14
333
22
11
333
22
11
333
22
11
333
22
11
333
20
10
333
22
11
333
22
11
333
18
9
333
22
11
333
22
11
333
22
11
333
22
11
333
34
17
333
265
Sex Chrom. Source
Karyotype
Taxa
Virgichneumon faunus
arrhenotoky
Vulgichneumon saturatorius
arrhenotoky
Leucospidae
Leucospis affinis
arrhenotoky
Megachilidae
Anthidium mormonum
arrhenotoky
Ashmeadiella sp.
arrhenotoky
Chalicodoma sculpturalis
arrhenotoky
Chalicodoma spissula
arrhenotoky
Coelioxys sp.
arrhenotoky
Dianthidium heterulkei heterulkei
arrhenotoky
Hoplitis robusta
arrhenotoky
Megachile ainu
arrhenotoky
Megachile pseudomonticola
arrhenotoky
Megachile relativa
arrhenotoky
Megachile rotundata
arrhenotoky
Osmia cornifrons
arrhenotoky
Osmia cornuta
arrhenotoky
Osmia cornuta
arrhenotoky
Osmia glauca
arrhenotoky
Osmia lignaria propinque
arrhenotoky
Osmia nigrifrons
arrhenotoky
Osmia pentstemonis
arrhenotoky
Osmia taurus
arrhenotoky
Stelis chlorocyanea
arrhenotoky
Trachusa gummifera
arrhenotoky
Ormyridae
Ormyrus sp.1
arrhenotoky
Pergidae
Perga sp.
arrhenotoky
Philomastix sp.
arrhenotoky
Pterygophorus sp.
arrhenotoky
Pompilidae
Anoplius concinnus
arrhenotoky
Anoplius viaticus
arrhenotoky
Sexual
System
Ploidy
30
Female
2n
Male
2n
22
11
333
18
9
333
12
6
333
32
16
331
32
16
331
32
16
332
32
16
332
32
16
331
15
Sex Chrom. Source
Karyotype
331
32
16
331
32
16
332
32
16
332
32
16
331
32
16
331
32
16
332
32
16
365
32
16
331
32
16
331
32
16
332
32
16
331
32
16
331
32
16
332
34
17
331
32
16
331
12
6
333
16
8
331
16
8
331
16
8
331
28
14
366
28
14
366
266
Taxa
Gen. sp.1
arrhenotoky
Gen. sp.1
arrhenotoky
Gen. sp.2
arrhenotoky
Gen. sp.2
arrhenotoky
Pteromalidae
Anisopteromalus calandrae
arrhenotoky
Coelopisthia extenta
arrhenotoky
Dibrachys sp.1
arrhenotoky
Lariophagus distinguesdus
arrhenotoky
Muscidifurax zaraptor
arrhenotoky
Nasonia vitripennis
arrhenotoky
Pteromalus puparum
arrhenotoky
Pteromalus venustus
arrhenotoky
Sapygidae
Sapyga pumila
arrhenotoky
Scelionidae
Telenomus fariai
arrhenotoky
Siricidae
Sirex cyaneus
arrhenotoky
Sirex juvencus
arrhenotoky
Sirex noctilio
arrhenotoky
Urocerus augur
arrhenotoky
Urocerus gigas
arrhenotoky
Sphecidae
Carinostigmus filippovi
arrhenotoky
Chalybion japonicum
arrhenotoky
Ectemnius rubicola nipponis
arrhenotoky
Microstigmus arlei
arrhenotoky
Microstigmus brasiliensis
arrhenotoky
Microstigmus cooperi
arrhenotoky
Microstigmus crucifex
arrhenotoky
Microstigmus luederwaldti
arrhenotoky
Pemphredon diervillae
arrhenotoky
Pemphredon krombeini
arrhenotoky
Pemphredon lethifer
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
30
15
331
30
15
332
30
15
331
30
15
332
14
7
333
10
5
333
10
5
333
10
5
333
10
5
333
12
6
333
10
5
333
10
5
333
52
26
367
20
10
333
16
8
331
16
8
331
16
8
331
36
18
331
26
13
331
28
14
332
48
24
332
28
14
332
6
3
368
10
5
368
6
3
369
8
4
369
6
3
368
8
4
332
16
8
332
16
8
332
267
Sex Chrom. Source
Karyotype
Taxa
Psenulus carnifrons iwatai
arrhenotoky
Psenulus maculipes
arrhenotoky
Rhopalum pygidiale
arrhenotoky
Rhopalum watanabei
arrhenotoky
Trychofoenus sp.1
arrhenotoky
Trypoxylon albitarse
arrhenotoky
Trypoxylon asuncicola
arrhenotoky
Trypoxylon fabricator
arrhenotoky
Trypoxylon nitidum
arrhenotoky
Trypoxylon obsonator
arrhenotoky
Trypoxylon petiolatum
arrhenotoky
Trypoxylon sp.1
arrhenotoky
Trypoxylon sp.1
arrhenotoky
Trypoxylon sp.2
arrhenotoky
Tenthredinidae
Aglaostigma albicincta
arrhenotoky
Aglaostigma amoorensis
arrhenotoky
Aglaostigma aucupariae
arrhenotoky
Aglaostigma nebulosa
arrhenotoky
Aglaostigma occipitosa
arrhenotoky
Aglaostigma sapporonis
arrhenotoky
Aglaostigma sp.
arrhenotoky
Allantus luctifer
arrhenotoky
Allantus meridionalis
arrhenotoky
Allantus nakabusensis
arrhenotoky
Allantus sp.
arrhenotoky
Alphostromboceros konowi
arrhenotoky
Ametastegia geranii
arrhenotoky
Ametastegia pallipes
arrhenotoky
Aneugmenus japonicus
arrhenotoky
Aneugmenus kiotonis
arrhenotoky
Aneugmenus stramineipes
arrhenotoky
Anoplonyx sp.
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
16
8
332
42
21
332
26
13
332
26
13
332
28
14
332
32
16
331
32
16
370
32
16
370
28
14
371
28
14
332
28
14
332
32
16
370
18
9
332
26
13
332
30
15
331
32
16
331
24
12
331
20
10
331
34
17
331
18
9
331
44
22
331
16
8
331
18
9
331
20
10
331
16
8
331
14
7
331
18
9
331
12
6
331
14
7
331
14
7
331
16
8
331
16
8
331
268
Sex Chrom. Source
Karyotype
Taxa
Asiemphytus albilabris
arrhenotoky
Athalia bicolor
arrhenotoky
Athalia cordata
arrhenotoky
Athalia japonica
arrhenotoky
Athalia kashmirensis
arrhenotoky
Athalia lugens infumata
arrhenotoky
Athalia rosae rosae
arrhenotoky
Athalia rosae ruficornis
arrhenotoky
Birka carinifrons
arrhenotoky
Cladius morio
arrhenotoky
Cladius pectinicornis
arrhenotoky
Corymbus fujisana
arrhenotoky
Corymbus nipponica
arrhenotoky
Croesus japonicus
arrhenotoky
Croesus septentrionalis
arrhenotoky
Croesus varus
arrhenotoky
Dolerus aeneus
arrhenotoky
Dolerus ephippiatus
arrhenotoky
Dolerus gessneri
arrhenotoky
Dolerus hematodes
arrhenotoky
Dolerus lewisii
arrhenotoky
Dolerus niger
arrhenotoky
Dolerus nigratus
arrhenotoky
Dolerus similis japonicus
arrhenotoky
Dolerus subfasciatus
arrhenotoky
Dolerus varispinus
arrhenotoky
Dolerus yokohamensis
arrhenotoky
Elinora koehleri
arrhenotoky
Empria sp.
arrhenotoky
Empronus obsoletus
arrhenotoky
Eriocampa mitsukurii
arrhenotoky
Eutomostethus juncivorus
arrhenotoky
Hemibeleses nigriceps
Sexual
System
Ploidy
Female
2n
Male
2n
30
15
331
12
6
331
12
6
331
16
8
331
12
6
331
16
8
331
16
8
331
16
8
331
14
7
331
12
6
331
12
6
331
20
10
331
20
10
331
16
8
331
16
8
331
16
8
331
16
8
331
16
8
331
16
8
331
16
8
331
28
14
331
16
8
331
16
8
331
18
9
331
18
9
331
16
8
331
16
8
331
20
10
331
30
15
331
20
10
331
18
9
331
12
6
331
32
16
331
269
Sex Chrom. Source
Karyotype
Taxa
Sexual
System
arrhenotoky
Hemichroa alni
arrhenotoky
Heptamelus (Heptamelus) ochroleucus
arrhenotoky
Heptamelus (Pseudoheptamelus) runari
arrhenotoky
Lagidina irritans
arrhenotoky
Lagidina platycerus
arrhenotoky
Loderus eversmanni obscurus
arrhenotoky
Loderus genucinctus insulicola
arrhenotoky
Macrophya albipuncta
arrhenotoky
Macrophya annulitibia
arrhenotoky
Macrophya apicalis
arrhenotoky
Macrophya carbonaria
arrhenotoky
Macrophya coxalis
arrhenotoky
Macrophya esakii exilis
arrhenotoky
Macrophya falsifica
arrhenotoky
Macrophya fascipennis
arrhenotoky
Macrophya imitator
arrhenotoky
Macrophya infumata
arrhenotoky
Macrophya malaisei
arrhenotoky
Macrophya montana
arrhenotoky
Macrophya punctumalbum
arrhenotoky
Macrophya ribis
arrhenotoky
Macrophya rohweri
arrhenotoky
Macrophya rufipes
arrhenotoky
Macrophya sp.
arrhenotoky
Macrophya timida
arrhenotoky
Monosoma pulveratum
arrhenotoky
Nematinus fuscipennis
arrhenotoky
Nematinus luteus
arrhenotoky
Nematinus willigkiae
arrhenotoky
Nematus (Lygaeonematus) erichsonii
arrhenotoky
Nematus (Nematus) lucidus
arrhenotoky
Nematus (Pachynematus) obductus
arrhenotoky
Ploidy
22
Female
2n
Male
2n
18
9
331
20
10
331
14
7
331
36
18
331
36
18
331
28
14
331
11
Sex Chrom. Source
Karyotype
331
16
8
331
20
10
331
16
8
331
20
10
331
20
10
331
16
8
331
20
10
331
24
12
331
16
8
331
18
9
331
20
10
331
16
8
331
20
10
331
24
12
331
20
10
331
20
10
331
20
10
331
20
10
331
16
8
331
16
8
331
16
8
331
16
8
331
14
7
331
16
8
331
16
8
331
270
Taxa
Sexual
System
Nematus (Pikonema) alaskaensis
arrhenotoky
Nematus (Pikonema) dimmockii
arrhenotoky
Nematus (Pontania) sp.
arrhenotoky
Nematus (Pontania) viminalis
arrhenotoky
Nematus (Pristiphora) geniculatus
arrhenotoky
Nematus (Pristiphora) pallipes
arrhenotoky
Nematus (Pristiphora) ruficornis
arrhenotoky
Nematus (Pristiphora) rufipes
arrhenotoky
Nematus (Pristiphora) sp.1
arrhenotoky
Nematus (Pristiphora) sp.2
arrhenotoky
Nematus (Pristiphora) sp.3
arrhenotoky
Nematus (Pteronidea) leucotrochus
arrhenotoky
Nematus (Pteronidea) melanaspis
arrhenotoky
Nematus (Pteronidea) nigricornis
arrhenotoky
Nematus (Pteronidea) olfaciens
arrhenotoky
Nematus (Pteronidea) pavidus
arrhenotoky
Nematus (Pteronidea) ribesii
arrhenotoky
Nematus (Pteronidea) viridescens
arrhenotoky
Nematus (s.l.) sp.
arrhenotoky
Neostromboceros itoi
arrhenotoky
Neostromboceros nipponicus
arrhenotoky
Neostromboceros okinawaensis
arrhenotoky
Neostromboceros sinanensis
arrhenotoky
Nesoselandria morio
arrhenotoky
Pachyprotasis asteris
arrhenotoky
Pachyprotasis caerulescens kashmirica
arrhenotoky
Pachyprotasis erratica
arrhenotoky
Pachyprotasis fukii
arrhenotoky
Pachyprotasis hayasuensis
arrhenotoky
Pachyprotasis hiensis
arrhenotoky
Pachyprotasis hiyodorii
arrhenotoky
Pachyprotasis iwatai
arrhenotoky
Pachyprotasis longicornis
Ploidy
Female
2n
16
8
331
16
8
331
18
16
Male
2n
9
8
16
Sex Chrom. Source
Karyotype
331
331
8
331
16
8
331
16
8
331
16
8
331
14
7
331
16
8
331
16
8
331
18
9
331
16
8
331
16
8
331
18
9
331
16
8
331
14
18
9
331
16
8
331
18
9
331
12
6
331
14
7
331
7
331
14
7
331
12
6
331
20
10
331
18
9
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
271
Taxa
arrhenotoky
Pachyprotasis Malaise
arrhenotoky
Pachyprotasis malaisei
arrhenotoky
Pachyprotasis nigrinotata
arrhenotoky
Pachyprotasis nogusai
arrhenotoky
Pachyprotasis okutanii
arrhenotoky
Pachyprotasis pallidiventris
arrhenotoky
Pachyprotasis rapae
arrhenotoky
Pachyprotasis sasabensis
arrhenotoky
Pachyprotasis sawadai
arrhenotoky
Pachyprotasis sengaminensis
arrhenotoky
Pachyprotasis serii
arrhenotoky
Pachyprotasis sp.1
arrhenotoky
Pachyprotasis sp.10
arrhenotoky
Pachyprotasis sp.2
arrhenotoky
Pachyprotasis sp.3
arrhenotoky
Pachyprotasis sp.4
arrhenotoky
Pachyprotasis sp.5
arrhenotoky
Pachyprotasis sp.6
arrhenotoky
Pachyprotasis sp.7
arrhenotoky
Pachyprotasis sp.8
arrhenotoky
Pachyprotasis sp.9
arrhenotoky
Pachyprotasis tanakai
arrhenotoky
Pachyprotasis volatilis
arrhenotoky
Pachyprotasis yamahakkai
arrhenotoky
Pachyprotasis zukaensis
arrhenotoky
Parachractus leucopodus
arrhenotoky
Perineura esakii
arrhenotoky
Perineura japonica
arrhenotoky
Perineura okutanii
arrhenotoky
Perineura pictipennis
arrhenotoky
Perineura sp.
arrhenotoky
Phyllocolpa sp.
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
22
11
331
22
11
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
22
11
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
34
17
331
34
17
331
34
17
331
34
17
331
34
17
331
18
9
331
272
Sex Chrom. Source
Karyotype
Taxa
Sexual
System
Priophorus varipes
arrhenotoky
Propodea fentoni
arrhenotoky
Pseudohemitaxonus dryopteridis
arrhenotoky
Rocalia japonica
arrhenotoky
Rocalia longipennis
arrhenotoky
Rocalia sp.
arrhenotoky
Siobla ferox
arrhenotoky
Siobla metallica
arrhenotoky
Siobla ruficornis
arrhenotoky
Siobla sturmi
arrhenotoky
Siobla venusta apicalis
arrhenotoky
Stromboceros koebelei
arrhenotoky
Strongylogaster blechni
arrhenotoky
Strongylogaster filicis
arrhenotoky
Strongylogaster lineata
arrhenotoky
Strongylogaster macula
arrhenotoky
Strongylogaster mixta
arrhenotoky
Strongylogaster moiwana
arrhenotoky
Strongylogaster onocleae
arrhenotoky
Strongylogaster osmundae
arrhenotoky
Strongylogaster ruber
arrhenotoky
Strongylogaster secunda
arrhenotoky
Strongylogaster tambensis
arrhenotoky
Taxonus alboscutellatus
arrhenotoky
Tenthredo (Eurogaster) mesomelas
arrhenotoky
Tenthredo (Eurogaster) obsoleta
arrhenotoky
Tenthredo (Eurogaster) opaciceps
arrhenotoky
Tenthredo (Olivacedo) olivacea
arrhenotoky
Tenthredo (Olivacedo) pseudolivacea
arrhenotoky
Tenthredo (Olivacedo) viridatrix
arrhenotoky
Tenthredo (Rhogogaster) sp.1
arrhenotoky
Tenthredo (Rhogogaster) sp.2
arrhenotoky
Tenthredo (Rhogogaster) sp.3
Ploidy
10
20
Female
2n
Male
2n
12
6
331
20
10
331
5
11
331
18
9
331
16
8
331
18
9
331
18
9
331
18
9
331
18
9
331
18
9
331
14
7
331
18
9
331
16
8
331
16
8
331
14
7
331
18
9
331
18
9
331
16
8
331
14
7
331
18
9
331
18
9
331
14
7
331
18
9
331
20
10
331
10
331
10
10
20
20
331
22
20
20
Sex Chrom. Source
Karyotype
331
331
10
10
331
331
20
10
331
20
10
331
20
10
331
273
Taxa
arrhenotoky
Tenthredo (Rhogogaster) viridis
arrhenotoky
Tenthredo (s.l.) abdominalis
arrhenotoky
Tenthredo (s.l.) angustiannulata
arrhenotoky
Tenthredo (s.l.) basizonata
arrhenotoky
Tenthredo (s.l.) contusa
arrhenotoky
Tenthredo (s.l.) convergenata
arrhenotoky
Tenthredo (s.l.) crassa
arrhenotoky
Tenthredo (s.l.) cylindrica
arrhenotoky
Tenthredo (s.l.) dentina
arrhenotoky
Tenthredo (s.l.) finschi seguro
arrhenotoky
Tenthredo (s.l.) flavomandibulata
arrhenotoky
Tenthredo (s.l.) fortunei
arrhenotoky
Tenthredo (s.l.) fukaii
arrhenotoky
Tenthredo (s.l.) gifui
arrhenotoky
Tenthredo (s.l.) hokkaidonis
arrhenotoky
Tenthredo (s.l.) japonica
arrhenotoky
Tenthredo (s.l.) jozana
arrhenotoky
Tenthredo (s.l.) latifasciata
arrhenotoky
Tenthredo (s.l.) matsumurai
arrhenotoky
Tenthredo (s.l.) melanogastra
arrhenotoky
Tenthredo (s.l.) mortivaga
arrhenotoky
Tenthredo (s.l.) nigropicta
arrhenotoky
Tenthredo (s.l.) nitidiceps
arrhenotoky
Tenthredo (s.l.) opposita
arrhenotoky
Tenthredo (s.l.) ornatula
arrhenotoky
Tenthredo (s.l.) picticornis
arrhenotoky
Tenthredo (s.l.) platycera
arrhenotoky
Tenthredo (s.l.) procincta
arrhenotoky
Tenthredo (s.l.) providens
arrhenotoky
Tenthredo (s.l.) rubrocaudata
arrhenotoky
Tenthredo (s.l.) sp.
arrhenotoky
Tenthredo (s.l.) sp.1
arrhenotoky
Sexual
System
Ploidy
Female
2n
24
12
20
16
Male
2n
Sex Chrom. Source
Karyotype
331
10
8
331
331
18
9
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
20
24
12
331
20
10
331
24
12
331
20
10
331
20
10
331
18
9
331
20
10
331
38
19
331
10
331
20
10
331
20
10
331
20
10
331
18
9
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
10
331
18
9
331
20
10
331
274
Taxa
Sexual
System
Tenthredo (s.l.) sp.2
arrhenotoky
Tenthredo (s.l.) sp.3
arrhenotoky
Tenthredo (s.l.) sp.4
arrhenotoky
Tenthredo (s.l.) sp.5
arrhenotoky
Tenthredo (s.l.) sp.6
arrhenotoky
Tenthredo (s.l.) subolivacea
arrhenotoky
Tenthredo (s.l.) takeuchii
arrhenotoky
Tenthredo (s.l.) ussuriensis
arrhenotoky
Tenthredo (Temuledo) temula
arrhenotoky
Tenthredo (Tenthredella) atra
arrhenotoky
Tenthredo (Tenthredella) bipunctula
arrhenotoky
Tenthredo (Tenthredella) colon
arrhenotoky
Tenthredo (Tenthredella) decens
arrhenotoky
Tenthredo (Tenthredella) fagi
arrhenotoky
Tenthredo (Tenthredella) ferruginea
arrhenotoky
Tenthredo (Tenthredella) fulva adusta
arrhenotoky
Tenthredo (Tenthredella) fuscoterminata
arrhenotoky
Tenthredo (Tenthredella) hilaris
arrhenotoky
Tenthredo (Tenthredella) limbata
arrhenotoky
Tenthredo (Tenthredella) livida
arrhenotoky
Tenthredo (Tenthredella) mandibularis
arrhenotoky
Tenthredo (Tenthredella) solitaria
arrhenotoky
Tenthredo (Tenthredella) velox
arrhenotoky
Tenthredo (Tenthredo) arcuata
arrhenotoky
Tenthredo (Tenthredo) brevicornis
arrhenotoky
Tenthredo (Tenthredo) marginella
arrhenotoky
Tenthredo (Tenthredo) notha
arrhenotoky
Tenthredo (Tenthredo) omissa
arrhenotoky
Tenthredo (Zonuledo) amoena
arrhenotoky
Tenthredopsis carinata
arrhenotoky
Tenthredopsis litterata
arrhenotoky
Tenthredopsis nassata
arrhenotoky
Tenthredopsis sp.1
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
26
13
331
22
11
331
20
10
331
22
11
331
18
9
331
20
10
331
20
10
331
20
10
331
20
10
331
20
10
331
24
12
331
40
20
331
22
11
331
38
19
331
20
10
331
36
18
331
16
8
331
18
9
331
18
9
331
16
8
331
275
Taxa
arrhenotoky
Tenthredopsis sp.2
arrhenotoky
Tenthredopsis sp.3
arrhenotoky
Thrinax athyrii
arrhenotoky
Thrinax japonicus
arrhenotoky
Thrinax melanogyne
arrhenotoky
Thrinax minomensis
arrhenotoky
Thrinax paucipunctatus
arrhenotoky
Thrinax sasayamensis
arrhenotoky
Thrinax struthiopteridis
arrhenotoky
Thrinax tokunagai
arrhenotoky
Torymidae
Monorionlolllems clemellti
arrhenotoky
Monorionlolllems montivagus
arrhenotoky
Monorionlolllems obscurus
arrhenotoky
Monorionlolllems saltuosus
arrhenotoky
Torymus baccharidis
arrhenotoky
Torymus californicus
arrhenotoky
Torymus capillaceus
arrhenotoky
Torymus koebelei
arrhenotoky
Torymus occidentalis
arrhenotoky
Torymus tubicola
arrhenotoky
Torymus umbilicatus
arrhenotoky
Torymus vesiculi
arrhenotoky
Torymus warreni
arrhenotoky
Trichogrammatidae
Trichogramma chilonis
arrhenotoky
Trichogramma deion
arrhenotoky
Trichogramma dendrolimi
arrhenotoky
Trichogramma evanescens
arrhenotoky
Trichogramma nubialale
arrhenotoky
Trichogramma pretiosum
arrhenotoky
Trichogramma sp.1
arrhenotoky
Vespidae
Brachygastra lecheguana
Sexual
System
Ploidy
Female
2n
Male
2n
16
8
331
16
8
331
12
6
331
18
9
331
12
6
331
16
8
331
12
6
331
12
6
331
12
6
331
14
7
331
12
6
333
12
6
333
10
5
333
10
5
333
12
6
333
12
6
333
12
6
333
10
5
333
12
6
333
12
6
333
10
5
333
12
6
333
12
6
333
10
5
333
10
5
333
10
5
333
10
5
333
10
5
333
10
5
333
10
5
333
56
28
331
276
Sex Chrom. Source
Karyotype
Taxa
arrhenotoky
Liostenogaster sp.1
arrhenotoky
Metapolybia sp.
arrhenotoky
Mischocyttarus cassununga
arrhenotoky
Mischocyttarus sp.
arrhenotoky
Parachartergus smithii
arrhenotoky
Polistes apachus
arrhenotoky
Polistes canadensis
arrhenotoky
Polistes carolinus
arrhenotoky
Polistes chinensis
arrhenotoky
Polistes chinensis antennalis
arrhenotoky
Polistes cinerascens
arrhenotoky
Polistes exclamans
arrhenotoky
Polistes fuscatus
arrhenotoky
Polistes gallicus
arrhenotoky
Polistes hebraeus
arrhenotoky
Polistes jadwigae
arrhenotoky
Polistes mandarinus
arrhenotoky
Polistes metricus
arrhenotoky
Polistes nimpha
arrhenotoky
Polistes omissus
arrhenotoky
Polistes simillimus
arrhenotoky
Polistes snelleni
arrhenotoky
Polistes versicolor versicolor
arrhenotoky
Polybia occidentalis
arrhenotoky
Polybia paulista
arrhenotoky
Polybia scutellaris
arrhenotoky
Polybia sericea
arrhenotoky
Polybia sp.1
arrhenotoky
Polybia sp.2
arrhenotoky
Protonectarina sylveirae
arrhenotoky
Protopolybia exigua exigua
arrhenotoky
Protopolybia pumila
arrhenotoky
Sexual
System
Ploidy
Female
2n
Male
2n
14
7
332
38
19
331
64
32
331
68
34
331
54
27
331
46
23
331
32
16
331
38
19
331
46
23
331
46
23
372
54
27
331
66
33
331
52
26
331
42
21
331
18
9
331
62
31
332
52
26
332
52
26
331
44
22
331
28
14
331
56
28
331
60
30
331
62
31
331
34
17
331
34
17
331
34
17
331
54
27
331
32
16
331
34
17
331
58
29
331
62
31
331
42
21
331
277
Sex Chrom. Source
Karyotype
Taxa
Pseudopolybia vespiceps
arrhenotoky
Stelopolybia multipicta multipicta
arrhenotoky
Stelopolybia pallipes pallipes
arrhenotoky
Vespa crabro
arrhenotoky
Vespa mandarinia
arrhenotoky
Vespa simillima xanthoptera
arrhenotoky
Vespula flaviceps
arrhenotoky
Vespula vulgaris
arrhenotoky
Isoptera
Kalotermitidae
Bifiditermes improbus
Cryptotermes austrinus
Cryptotermes brevis
Cryptotermes cristatus
Cryptotermes cynocephalus
Cryptotermes domesticus
Cryptotermes domesticus
Cryptotermes dudleyi
Cryptotermes gearyi
homomorphic
Cryptotermes papulosus
Cryptotermes primus
Cryptotermes queenslandis
Cryptotermes riverinae
Cryptotermes secundus
Cryptotermes secundus
Cryptotermes sp.
Glyptotermes brevicornis
Glyptotermes iridipennis
Incisitermes barretti
Incisitermes sp.
Kalotermes convexus
Kalotermes flavicollis
Neotermes insularis
Neotermes insularis
Procryptotermes australiensis
Rhinotermitidae
Coptotermes acinaciformis
Reticulitermes lucifugus grassei
Reticulitermes urbis
homomorphic
Stolotermitidae
Porotermes adamsoni
Stolotermes victoriensis
Termitidae
Acidnotermes praus
Afrosubulitermes congoensis
Amitermes darwini
Amitermes eucalipti
Amitermes germanus
Amitermes parvus
Crenetermes albotarsalis
Cubitermes exiguus
Cubitermes sankurensis
Cubitermes sp. 1.
Cubitermes weissi
Drepanotermes septentrionalis
Sexual
System
Ploidy
64
Female
2n
Male
2n
16
8
32
Sex Chrom. Source
Karyotype
331
331
64
32
331
50
25
332
50
25
331
50
25
331
50
25
331
50
25
373
36
35
40
37
46
43
29
29
40
48
36
44
30
30
40
42
30
42
42
40
40
41
42
XXY
XYY
XXY
XXY
XXY
XY
XXY
40
47
42
42
42
42
48
46
67
52
60
47
42
42
42
42
XXYY
42
40
32
40
31
XO
42
XXYY
XXYY
XXYY
68
52
42
42
42
42
42
42
42
42
42
42
42
42
42
278
42
42
XXY
XXXY
XXY
XXYY
244
374
374
374
374
374
244
374
374
374
374
374
374
374
244
374
374
374
374
374
374
374
244
374
374
374
244
244
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
374
374
375
375
244
244
244
244
375
375
375
375
375
244
Taxa
Sexual
System
Ephelotermes melachoma
Ephelotermes taylori
Kalotermes approximatus
Kalotermes approximatus
Kalotermes Flavicollis
Lophotermes septentrionalis
Macrognathotermes sunteri
Macrotermes bellicosus
Mastotermes darwiniensis
homomorphic
Microcerotermes boreus
Microcerotermes fuscotibiatis
Microcerotermes nervosus
Microcerotermes parvulus
Microcerotermes sp.1
Microcerotermes sp.2
Microtermes sp.1
Nasutitermes arboreus
Nasutitermes graveolus
Nasutitermes longipennis
Nasutitermes triodiae
Noditermes lamanianus
Odontotermes redemanni
Odontotermes snyderi
Odontotermes sp.1
Ophiotermes mandibularis
Pericapritermes sp.1
Procubitermes sp. 1
Protermes minimus
Pseudacanthotermes militaris
Reticulitermes flavipes
Reticulitermes lucifugus
Reticutitermes santonensis
Schedorhinotermes lamanianus
Tenuirostritermes tenuirostris
Thoracotermes macrothorax
Tuberculitermes bycanistes
Tumulitermes pastinator
Unguitermes bouilloni
Zootermopsis angusticollis
Zootermopsis angusticollis
Zootermopsis nevadensis
Lepidoptera
Agaristidae
Seudyra subflava
Arctiidae
Abraxas grossulariata
Abraxas grossulariata dohrnii
Arctia caja
Arctia caja
Arctia caja
Arctia caja phaesoma
Arctia hebe
Cycnial mendica
Cycnial mendica
Earias fabia
Earias insulana
Hippocrita jacobaea
Lophocampa maculata
Miltochrista miniata
Pericallia ricini
Phragmatobia fuliginosa
Phragmatobia fuliginosa amurensis
Spilarctis imparilis
Spilaritia infernalis
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
42
42
34
32
61
42
42
42
98
42
42
33
XXYY
XXYY
XXY
XXYY
XXYY
XXYY
XXYY
XXYY
XY
244
244
376
375
375
244
244
375
245
42
42
42
42
44
42
42
42
42
42
42
38
48
42
42
42
42
38
42
42
42
42
42
38
42
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
XXYY
244
375
244
375
375
375
375
375
244
244
244
375
375
375
375
375
375
375
375
375
375
375
375
375
375
375
375
244
375
375
375
375
42
42
42
42
52
52
52
33
57
279
42
42
42
42
42
42
42
62
377
56
56
56
62
62
62
63
52
62
62
62
62
34
62
62
56
58
62
62
377
377
377
377
377
377
377
377
377
377
377
377
378
377
377
377
377
377
377
ZO
ZWW
Taxa
Spilosoma lubricipeda
Spilosoma lubricipeda
Spilosoma menthastri
Spilosoma menthastri
Utetheisa ornatrix
Utetheisa ornatrix bella
Utetheisa ornatrix stretchii
Utetheisa pulchella
Bombycidae
Bombyx mandarina
Bombyx mandarina
Bombyx mandarina
Bombyx mandarina
Bombyx mori
Bombyx mori
Bombyx mori
Theophila religiosae
Carposinidae
Carposina niponensis
Crambidae
Ostrinia scapulalis
Cymatophoridae
Thyatira batis
Danaidae
Euploes mulciber barsine
Eupterotidae
Apha tychoona
Gelechiidae
Exoteleia dodecella
Exoteleia pinifoliella
Phthorimaea operculella
Tachyptilia populella
Geometridae
Agathia carissima
Aperia syringaria
Ascotis selenaria cretacea
Boarmia consonaria
Calospiloa sylvata fulvobasalis
Campaea margatitata
Camptogramma bilineata
Chlorochysta miata
Coenotephria sagittata
Colostygia aptata
Colostygia didymata
Colostygia olivata
Colostygia parallelolineata
Colostygia pectinataria
Cusiala stipitaria karuizawensis
Dysstroma citrata
Dysstroma latefasciata
Dysstroma truncata
Epirrho alternata
Epirrho galiata
Eptrrho trista
Eulype hastata
Eulype subhastata
Euphyia corylata
Euphyia cucullata
Euphyia luctuata
Euphyia unangulata
Hydrelia flammeolaria
Hydrelia testaceata
Hydriomena coerulata
Hydriomena furcata
Hydriornena ruberata
Sexual
System
Ploidy
Female
2n
56
Male
2n
Sex Chrom. Source
Karyotype
62
62
60
60
62
62
62
62
377
377
377
377
377
377
377
377
54
54
56
56
56
56
56
62
377
377
377
377
377
377
377
377
ZW
62
62
23
21
58
62
377
ZW
62
377
58
377
44
377
24
22
58
58
ZO
ZO
ZW
62
58
68
62
58
378
378
378
377
377
377
377
377
377
62
60
60
24
60
54
62
60
62
60
377
377
377
377
377
377
377
377
377
377
58
58
58
62
62
60
62
62
60
62
60
56
60
26
60
56
60
280
378
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Sexual
System
Ploidy
Female
2n
Lampropteryx minna
Lampropteryx suffumata
Larentia autumnalis
Lycia hirtaria
Lycia hirtaria
Lyncometra ocellata
Mesolenca albicillata
Nyssia zonaria
Opisthograptis luteolata
Oporinia autumnata
Oporinia christyi
Oporinia filigrammaria
Ourapteryx sambucaria
Pelurga comitata
Perizoma affinitata
Perizoma alchemillata
Perizoma blandiata
Perizoma flavofasciata
Perizoma hydrata
Perizoma taeniata
Phigalia pilosaria
Plemyria bicolorata
Poecilopsis pomonaria
Pseudoboarmia punctinalis
Psychophora caesiata
Psychophora sabini frigidaria
Qporinia dilutata
Selenia bilunaria
Selenia lunaria
Selenia tetralunaria
Thera cognata
Thera firmata
Thera juniperata
Thera obeliscata
Thera variata
Xanthorho fluctuata
Xanthorho montanata
Xanthorho quadrifasciata
Xantlwrho spadicearia
Zonosoma pendularia
Gracilariidae
Gracilaria elongella
Hesperidae
Achalarus lycidas
Achalarus toxeus
Augiades venata
Bibasis aquilina
Carcharadus alceae
Cegenes pumilio
Chioides catillus
Choaspes benjaminii
Cotia otho
Daimio tethys
Erynnis baptisiae
Erynnis horatius
Erynnis icelus
Erynnis juvenalis
Erynnis lucilus
Erynnis marloyi
Erynnis montanus
Erynnis persius
Erynnis tages
Erynnis tages
Grais stigmaticus
Hesperia alveus
281
Male
2n
Sex Chrom. Source
Karyotype
34
64
60
28
28
62
62
112
62
76
62
74
62
64
50
60
60
60
50
64
224
60
102
64
62
58
60
60
62
58
40
38
60
26
26
62
59
62
58
62
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
58
377
62
32
26
58
62
48
62
62
62
60
62
62
60
60
62
62
62
62
62
62
62
48
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Sexual
System
Hesperia alveus
Hesperia malvae
Hesperia onopordi
Hesperia orbifer
Hesperia sao
Hesperia serratulae
Lavatheria lavatherae
Muschampsia proto
Nastra I'herminieri
Ochlodes ochracea
Ochlodes sylvanoides
Ochlodes venata
Ochlodes venata
Parnara guttata
Pholisora catullus
Poanes hobomok
Polytremis mathiades
Polytremis pellucida
Pyrgus bellieri
Pyrgus cacaliae
Pyrgus carlinae
Pyrgus carlinae cirsii
Pyrgus carthami
Pyrgus malvae
Reverdinus boeticus
Reverdinus floccifera
Reverdinus floccifera orientalis
Reverdinus floccijera dravira
Reverdinus stauderi ambigua
Sarangesa phidyle
Spialia orbifer
Spialia phlomidis
Thanaos tages
Thoressa varia
Thorybes pylades
Thymelicus lineola
Thymelicus lineola
Thymelicus sylvestris
Tuttia tessellum
Lasiocampidae
Dendrolinus jezoensis
Dendrolinus pini
Dendrolinus spectabilis
Epicnaptera ilicifolia
Epicnaptera tremulifolia
Malacosoma castrense
Malacosoma neustria
Malacosoma neustria
Philudoria potatoria
Philudoria potatoria
Poecilocompa populi
Trabala vishnu
Trichiura crataegi
Lycaenidae
Acrodiaetus menalcas
Agriades glandon
Agriades glandon rusticus
Agrodiaetus actis firdussi
Agrodiaetus actis pseudactis
Agrodiaetus admetus
Agrodiaetus admetus anatoliensis
Agrodiaetus alcestis
Agrodiaetus allivaganus pseudoxerces (?)
Agrodiaetus altivagans
Agrodiaetus antidolus
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
48
62
60
60
62
60
60
60
60
48
58
58
58
32
58
58
32
32
54
60
60
60
58
66
90
90
67
51
377
85
60
58
62
62
62
62
62
54
58
54
60
377
377
377
377
377
377
377
377
377
377
377
377
60
60
60
62
62
62
62
62
62
62
72
52
56
377
377
377
377
377
377
377
377
377
377
377
378
377
170
48
48
65
55
160
157
41
32
41
123
282
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
ZZW
377
377
377
377
377
377
377
377
377
377
377
Taxa
Sexual
System
Agrodiaetus ardschira
Agrodiaetus baytopi
Agrodiaetus carmon
Agrodiaetus cyanea
Agrodiaetus cyanea brandti
Agrodiaetus cyanea kermansis
Agrodiaetus cyanea paracyanea
Agrodiaetus dama
Agrodiaetus dama hamadanensis .
Agrodiaetus damone
Agrodiaetus damone maraschi (?)
Agrodiaetus damone wagneri
Agrodiaetus demavendi
Agrodiaetus dolus
Agrodiaetus dolus ainsae
Agrodiaetus dolus pseudovirgilia
Agrodiaetus dolus vittata
Agrodiaetus erschoffi tekkeana
Agrodiaetus fabressei
Agrodiaetus hopfferi
Agrodiaetus hopfferi hadjina
Agrodiaetus hopfferi sennanensis
Agrodiaetus interjectus
Agrodiaetus iphigenia
Agrodiaetus iphigenia iphidamon
Agrodiaetus mithridates saetosus
Agrodiaetus mofidii
Agrodiaetus morgani
Agrodiaetus pfeifferi
Agrodiaetus phyllides askhabadica
Agrodiaetus phyllis
Agrodiaetus phyllis vanensis
Agrodiaetus poseidon
Agrodiaetus poseidon mesopotamica
Agrodiaetus posthumus
Agrodiaetus ripartii
Agrodiaetus ripartii paralcestis
Agrodiaetus tankeri
Agrodiaetus transcaspica
Agrodiaetus transcaspica elbursica
Agrodiaetus transcaspica ninae
Albulina orbitulus
Antigius attilia
Aricia agestis
Aricia agestis
Aricia agestis
Aricia agestis
Aricia allous
Aricia anteros
Aricia isaurica
Aricia nicias
Asrodiaetus mithridates
Calastrina argiolus
Calastrina argiolus
Calastrina argiolus
Calastrina argiolus
Callophrys rubi
Callophrys rubi
Calycopis cecrops
Chrysophanus hippotho
Chrysophanus virgaureae
Chrysozephyrus aurorimus
Curetis acuta paracuta
Cyaniris semiargus
Cyaniris semiargus
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
127
54
163
37
37
44
39
377
377
377
377
377
377
377
83
43
90
32
32
377
377
377
377
377
137
148
118
216
377
377
377
377
149
28
377
377
180
31
30
58
377
377
377
377
60
27
28
43
377
377
377
377
69
84
214
124
161
156
41
40
21
180
180
377
377
377
377
377
377
377
377
377
377
377
42
105
33
70
377
46
50
46
46
48
48
46
46
48
46
53
48
50
50
50
46
46
48
48
48
48
58
46
48
283
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Cyaniris semiargus antiochena
Cyaniris semiargus persica
Eumaeus debora
Everes alcetas
Everes alcetas
Everes amyntula
Everes argiades
Everes argiades
Everes comyntas
Everes comyntas
Everes decolorata
Everes minimus
Everes minimus
Everes sebras
Favonius cognatus
Freyeria trochilus
Glaucopsyche alexis
Glaucopsyche arion
Glaucopsyche cyllarus
Glaucopsyche melanops
Heliophorus epicles matsumurae
Icaricia icarioides
Iolana iolas
Japonica saepestriata
Lycaeides idas
Lycaeides idas
Lycaeides melissa
Lycaena agyrognomon
Lycaena alciphron
Lycaena amandus
Lycaena argus
Lycaena arion
Lycaena asabinus
Lycaena dorilis
Lycaena eumedon
Lycaena helloides
Lycaena heteronea
Lycaena hippotho
Lycaena hypophleas
Lycaena icarus
Lycaena lampon
Lycaena nivalis
Lycaena ochimus
Lycaena optilete
Lycaena phlaeas
Lycaena phlaeas
Lycaena phlaeas
Lycaena phlaeas
Lycaena phoenicurus
Lycaena rubidus
Lycaena snowi
Lycaena thersamon
Lycaena thetis
Lycaena tityrus
Lysandra albicalis esteparina
Lysandra albicans
Lysandra albicans pennaelensis
Lysandra argester
Lysandra argester
Lysandra bellargus
Lysandra bellargus
Lysandra candalus
Lysandra candalus isauricoides
Lysandra candalus zuleikae
Lysandra coelestissima
Sexual
System
Ploidy
Female
2n
Male
2n
48
Sex Chrom. Source
Karyotype
377
48
48
51
52
48
48
48
48
48
50
48
48
48
48
46
46
46
46
46
50
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
48
44
50
47
48
48
48
48
46
46
46
48
48
48
48
136
48
48
46
48
48
48
48
48
48
48
48
48
76
48
48
48
48
164
164
164
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
191
282
90
90
52
76
377
63
169
284
377
377
377
377
377
377
377
Taxa
Sexual
System
Lysandra coeruleossmar
Lysandra coridon
Lysandra coridon
Lysandra coridon
Lysandra coridon hispanagallica
Lysandra coridon jurae
Lysandra coridon manleyi
Lysandra coridon maritimarum
Lysandra corldon gallica
Lysandra escheri
Lysandra hispana
Lysandra hispana constanti
Lysandra hispana galliaealbicans
Lysandra hispana pseudoalbicans
Lysandra hispana rezniceki
Lysandra myrrha
Lysandra nausithous
Lysandra nivescens
Lysandra ossmar
Lysandra punctifera
Lysandra syriaca
Lysandra thersites
Lysandra thersites
Maculinea alcon
Maculinea alcon tarannis
Meleagerta meleager
Narathura japonica
Neozephyrus taxila
Philotes baton
Philotes bavius
Philotes vicrama
Plebeius eurypilus
Plebeius loewii
Plebeius pylaon
Plebeius pylaon nicholi
Plebeius pylaon solimana
Polyommatus amandus
Polyommatus argus
Polyommatus argyrognomon energetes
Polyommatus eroides
Polyommatus eros
Polyommatus eumedon
Polyommatus hylas
Polyommatus icarus
Polyommatus icarus
Polyommatus icarus persica
Polyommatus idas armoricana
Polyommatus idas croatica
Polyommatus meleager
Satyrium sylvinus
Scolitantides orion
Strymon melinus
Strymon pruni
Strymon pruni
Syntarucus telicanus
Taraka hamada
Tarucus balcanicus
Thecla quercus
Turanana panagaea
Ussuriana stygiana
Vacciniina alcedo
Vacciniina hyrcana
Vacciniina sieversi
Zephyrus betulae
Zizeeria maha argia
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
168
176
180
180
176
377
377
377
377
377
176
176
176
377
377
377
176
46
168
168
168
377
377
377
377
377
168
168
52
48
381
168
48
48
48
48
46
46
46
48
48
48
48
48
40
48
38
42
42
48
46
48
46
46
48
48
46
46
46
48
377
48
48
82
46
48
46
46
48
30
48
48
48
94
46
44
44
32
48
285
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Lymantriidae
Dasychira pseudabietis
Dasychira pudibunda
Dasychira selenitica
Euproctis flava
Euproctis pseudoconspersa
Euproctis similis
Euproctis similiso
Leucoma salicis
Lymantria dispar
Lymantria dispar
Lymantria dispar
Lymantria dispar
Lymantria dispar japonica
Lymantria fumida
Lymantria mathura aurora
Lymantrla monacha
Orgyia antiqua
Orgyia antiqua
Orgyia antiqua
Orgyia ericae
Orgyia ericae
Orgyia leucostigma
Orgyia recens
Orgyia thyellina
Orgyia thyellina
Megathymidae
Agathymus mariae
Megathymus violae
Stallingsia maculosa
Micropterigidae
Micropterix calthella
Noctuidae
Adris tyrannus amurensis
Agrotis fimbria
Agrotis triangulum
Amathes c-nigrum
Amathes xanthographa
Amphipoea burrowsi
Anomis commoda
Apamea monoglypha
Apatele incretata
Apatele psi
Apatele psi
Arcte coerulea
Autographa nigrisigma
Calpe aureola
Cosmia camplostigma
Dichromia trigonalis
Euplexia lucipara
Gortyna flavago
Hoplotarache lunana
Hypena proboscidalis
Leucania conigera
Leucania favicolor
Leucania impura
Leucania impura
Leucania pallens
Luperina testacea
Mamestra brassicae
Mamestra persicariae
Ophiusa melicerte
Orthosia circellaris
Orthosia gracilis
Plusia intermixta
Sexual
System
Ploidy
Female
2n
62
28
23
22
Male
2n
104
174
44
44
44
44
46
60
62
62
62
62
62
60
62
62
28
28
28
60
60
56
60
22
22
Sex Chrom. Source
Karyotype
ZW
ZW
ZWW
ZW
42
54
100
57
27
286
58
62
58
58
58
68
62
62
58
62
62
62
62
62
62
62
62
62
62
64
62
62
62
58
62
62
58
62
62
62
60
28
62
377
377
377
377
377
377
377
377
377
377
377
378
377
377
377
377
377
377
378
377
377
377
377
378
378
377
377
377
ZO
ZO
378
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
378
377
Taxa
Sexual
System
Ploidy
Female
2n
Plusia peponis
Plusiodonta casta
Polytela gloriosae
Proderia littora
Pseudoips prasinana
Xylina ingrica
Notodontidae
Cerura erminea
Cerura vinula
Cerura vinula albanica
Cerura vinula delavoiei
Cerura vinula fennica
Cerura vinula germanica
Clostera anachoreta
Clostera anachoreta
Clostera anastomosis
Clostera anastomosis tristis
Clostera apicalis
Clostera curtula
Clostera pigra
Clostera timon
Drepana curvatula
Drepana falcataria
Harpyia bicuspis
Harpyia bifida
Harpyia furcula
Harpyia lanigera
Lophopteryx camelina
Notodonta phoebe
Notodonta trltophus
Notodonta ziczac
Odontosia camelina
Phalera bucephala
Phalera bucephala
Nymphalidae
Acraea bonasia
Acraea natalica pseudegina
Aglais urticae
Aglais urticae
Aglais urticae
Aglais urticae
Aglais urticae connexa
Agraulis juno
Amauris egialea hyalites
Amauris hecate
Amauris tartarea
Anaea aidea
Anartia fatima
Anartia jatrophae
Apatura ilia
Apatura ilia substituta
Araschnia burejana
Araschnia burejana strigosa
Araschnia Ievana
Araschnia Ievana obscura
Araschnia levana
Argynnis aglaia
Argynnis aglaia
Argynnis anadyomene
Argynnis anadyomene midas
Argynnis aphirape ossianus
Argynnis euphrosyne
Argynnis euphrosyne
Argynnis freija
Argynnis frigga
287
Male
2n
Sex Chrom. Source
Karyotype
62
62
62
62
64
62
377
377
377
377
377
377
56
42
40
62
42
42
60
60
50
50
60
58
46
20
62
62
60
98
58
58
62
62
62
62
62
60
60
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
64
62
58
62
62
62
62
62
58
58
77
60
62
62
62
62
62
62
62
62
62
58
58
72
74
56
62
62
62
62
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Sexual
System
Argynnis ino
Argynnis lathonia
Argynnis pales arsilache
Argynnis paphia
Argynnis paphia
Argynnis paphia
Argynnis paphia
Argynnis paphia geisha
Argynnis selene
Argynnis thore scandinavica
Argyreus hyperbius
Argyronome loadice
Argyronome loadice japonica
Argyronome ruslana
Argyronome ruslana lysippe
Asterocampa celtis
Asterocampa Ieilia
Aterica galene
Biblis hyperia
Boloria aquilonaris
Boloria eunomia caelestis
Boloria graeca
Boloria napaea
Boloria pales
Boloria selene tollandensis
Boloria titania helena
Brenthis daphne
Brenthis daphne rabdia
Brenthis heate
Brenthis ino mashurnsis
Brenthis ino tigroides
Caduga sita
Charaxes fulvescens
Charaxes zingha
Charaxes zingha
Chlosyne damoetas
Chlosyne harrish
Chlosyne palla
Clossiana thore jezoensis
Clossiana titania
Cyrestis thyodamas
Cyrestis thyodamas formosana
Damora sagana Iiane
Danaus chrysippus
Danaus chrysippus
Danaus eresimus
Danaus gilippus
Danaus limniace petiverana
Dryas julia
Euphaedra edwardsii
Euphydryas anicia eurytion
Euphydryas aurinia
Euptoieta hegesia
Euryphene mardania senegalensis
Euryphura chalcis
Euryphura plautilla
Euthalia thibetana insulae
Fabriciana adippe
Fabriciana nerippe
Fabriciana niobe
Fabriciana niobe
Fahriciana adippe
Fahriciana adippe pallescens
Hamadryas glauconome
Hestina japonica
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
60
60
58
58
58
58
58
60
60
62
62
62
52
52
62
62
45
56
60
56
62
62
60
60
60
26
28
68
28
28
94
114
52
54
62
62
62
62
62
62
62
377
62
60
62
60
58
87
62
60
62
60
62
66
60
60
28
58
58
58
58
58
58
62
60
288
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Hypolimnas bolina kezia
Hypolimnas misippus
Inachus io geisha
Junonia chorimene
Junonia coenia
Junonia evarete zonalis
Junonia hierta
Junonia oenone
Junonia stygia
Junonia westermanni
Kalkasia philyra
Kaniska canace
Ladoga camilla
Ladoga glorifica
Lampides boeticus
Limenitis anonyma
Limenitis archippus
Limenitis astyanax
Limenitis camilla
Limenitis camilla
Limenitis glorifica
Limenitis populi
Limenitis populi
Limenitis weidemeyerii
Melanargia galathea
Melitaea athalia
Melitaea aurelia
Melitaea cinxia
Melitaea diamina
Melitaea didyma
Melitaea iduna
Melitaea maturna
Melitaea maturna
Melitaea montium
Melitaea persea
Melitaea phoebe
Melitaea transcaucasica
Melitaea trivia
Mellicta athalia
Mellicta varia
Mesoacidalia charlotta
Mesoacidalia charlotta
Mesoacidalia chartotta .basalis
Metamorpha stelenes
Morpho peleides
Neptis aceris
Neptis aceris
Neptis hylas luculenta
Neptis morosa
Neptis nysiades
Nymphalis antiopa
Nymphalis antiopa
Nymphalis antiopa
Nymphalis canace
Nymphalis io
Nymphalis xanthomelas
Nymphalis xanthomelas japonica
Palla violinitens
Pandoriana maja
Paraneptis pryeri
Paraneptis rivularis
Phyciodes phaon
Phyciodes tharos
Polygnnia c-aureum
Polygonia c-album
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
62
62
62
62
62
62
62
62
66
62
60
62
60
60
48
60
60
60
60
60
60
60
60
60
48
62
64
62
62
56
62
62
62
54
55
62
58
62
62
62
58
58
58
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
62
56
60
60
60
56
88
60
62
62
62
62
62
62
377
34
58
60
60
62
62
62
62
289
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Polygonia c-album
Polygonia c-album
Polygonia c-album hamigera
Polygonia egea
Polygonia vau-album samurai
Polygonia zephyrus
Polygonla c-album
Precis almana
Precis almana
Precis iphita
Precis orithya
Precis pelarga
Precis pelarga leodice
Pseudacraea boisduvalii
Pseudacraea gottbergi
Pyrameis atalanta
Pyrameis cardui
Salamis cacta
Salamis parhassus
Sasakia charonda
Sephisa daimio
Speyeria aphrodite
Speyeria aphrodite ethne
Speyeria atlantis nikias
Speyeria callippe meadii
Speyeria coronia halcyone
Speyeria cybele
Speyeria hydaspe sakuntala
Speyeria marmonia eurynome
Speyeria zerene sinope
Thaleropis jonia
Vanessa cardui
Vanessa cardui
Vanessa indica
Vanessa indica
Vanessa kashmirensis
Vanessa polychloros
Vanessa virginiensis
Vanessa xanthomelas
Oecophoridae
Depressaria nervosa
Papilionidae
Battus philenor
Byasa alcinous
Byasa latreillei
Byasa polyeuctes letincius
Byasa polyeuctes termessus
Chilasa clytia
Graphium doson
Graphium feisthameli
Graphium phaon
Graphium podalirius
Graphium podalirius
Graphium podalirius
Graphium sarpedon
Luedorfia japonica
Luedorfia puziloi
Papilio alexanor
Papilio bianor
Papilio brucei (?)
Papilio cresphontes
Papilio dardanus
Papilio demoleus
Papilio helenus
Papilio helenus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
62
62
62
62
62
377
377
377
377
377
62
62
62
62
62
62
62
62
60
60
62
62
42
65
58
62
54
58
58
60
60
58
58
58
290
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
58
62
62
62
30
62
62
62
62
62
377
377
377
377
377
377
377
377
377
377
60
377
60
60
60
60
60
60
60
60
60
56
60
60
40
62
60
60
60
60
60
60
60
60
60
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Papilio lnachaon hippocrates
Papilio maackii
Papilio machaon
Papilio machaon
Papilio machaon
Papilio machaon
Papilio machaon emihippocrates
Papilio memnon
Papilio memnon heronus
Papilio nepheles chaon
Papilio ornythion
Papilio palamedes
Papilio paris decorosa
Papilio polyctor
Papilio polytes pasikrates
Papilio polytes romulus
Papilio polyxenes
Papilio protenor demetrius
Papilio protenor euprotenor
Papilio rutulus
Papilio thoas
Papilio troilus
Papilio xuthus
Papllio pilumnus
Parnassius apollo
Parnassius apollo
Parnassius mnemosyne
Parnassius smintheus
Pornassius eversmanni
Princeps demoleus
Troides aeacus
Zerynthia hypermnestra
Phycitidae
Anagasta khniella
Anagasta khniella
Anagasta khniella
Anagasta khniella
Anagasta khniella
Eurrhypara hortulata
Syllepte derogata
Pieridae
Anteos clorinde
Anthocharis cardamines
Anthocharis scolymus
Aporia crataegi
Aporia crataegi
Aporia crataegi
Aporia crataegi
Aporia crataegi adherbal
Aporia hippia japonica
Appias drusilla
Appias lyncida formosana
Appias sabina
Ascia monuste
Belenois aurota
Belenois calypso
Belenois creona
Belenois gidica
Belenois mesentina
Belenois theora concolor
Catopsilia crocale
Catopsilia crocale
Catopsilia florella
Catopsilia pyranthe
Catopsilia pyranthe
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
62
60
62
60
60
64
60
291
377
377
377
377
377
377
377
60
60
60
60
60
60
60
60
60
60
60
60
64
54
60
60
60
60
60
58
60
124
60
60
62
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
58
60
60
60
60
56
62
377
377
377
377
377
377
377
62
62
62
50
50
52
52
50
50
64
64
64
54
50
50
50
52
50
50
62
62
62
62
62
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Catopsllia pomona
Cepora coronis phryne
Cepora nadina eunama
Colias alexandra
Colias aurorina
Colias australis
Colias chlorocoma
Colias croceus
Colias croceus
Colias electo fieldii
Colias erate nilagiriensis
Colias erate poliographus
Colias eurytheme
Colias hecta sulitelma
Colias hyale
Colias hyale
Colias hyale
Colias meadii
Colias myrmidone
Colias noster werdandi
Colias palaeno
Colias palaeno sugitanii
Colias phicimone
Colias philodice
Colias philodice
Colias sagartia
Colias scudderii
Colias stoliczkana miranda
Colias thisoa
Colotis aurora evarne
Colotis liagore
Delias aglaia
Delias aglaia curasena
Delias descombesi
Delias eucharis
Dixeia orbona
Euchloe ausonia
Euchloe ausonides
Euchloe cardamines
Euchloe cardamines
Euchloe cardamines
Euchloe charlonia
Euchloe crameri occidentalis
Euchloe crameri romana
Euchloe gruneri
Euchloe lessei
Eurema brigitta
Eurema hecabe contubernalis
Eurema hecabe mandarina
Eurema laeta bethesba
Eurema lisa
Eurema nicippe
Eurema proterpia
Eurema senegalensis
Gonepteryx farinosa
Gonepteryx mahaguru niponica
Gonepteryx rhamni
Gonepteryx rhamni
Gonepteryx rhamni
Gonepteryx rhamni maxima
Hebomoia glaucippe
Hebomoia glaucippe formosana
Hebomoia glaucippe shorizui
Kricogonia lyside
Leptidea amurensis
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
62
46
50
62
64
62
62
62
62
62
62
62
62
62
63
62
62
62
61
62
63
62
60
62
62
64
62
62
65
30
56
50
50
59
50
48
62
62
60
62
62
62
62
62
48
55
24
62
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
48
58
62
62
62
62
64
62
377
377
377
377
377
377
377
377
63
62
62
62
34
34
377
34
62
128
292
377
377
377
377
377
377
377
377
Taxa
Leptidea duponcheli
Leptidea duponcheli
Leptidea duponcheli
Leptidea morsei
Leptidea morsei
Leptidea sinapis
Leptidea sinapis
Leptosia alcesta
lxias pyrene familiaris
lxias pyrene insignis
Mylothris hilara
Mylothris ochracea
Mylothris rhodope
Phoebis philea
Pieris beckerii
Pieris brassicae
Pieris brassicae
Pieris brassicae
Pieris brassicae
Pieris brassicae
Pieris brassicae
Pieris brassicae azoriensis
Pieris brassicae nepalensis
Pieris calyce
Pieris conidia
Pieris conidia indica
Pieris ergane
Pieris krueperi
Pieris manni
Pieris manni
Pieris melete pseudonapi
Pieris napi
Pieris napi
Pieris napi
Pieris napi bryoniae
Pieris napi bryoniae
Pieris napi meridionalis
Pieris napi nesis
Pieris napi sulphurea
Pieris occidentalis
Pieris rapae
Pieris rapae
Pieris rapae
Pieris rapae
Pieris rapae
Pieris rapae crucivora
Pinacopteryx eriphia
Pleris daplidicae
Pleris napi
Pleris napi macdunnoughli
Pontia daplidice moorei
Synchlo callidice
Synchlo callidice
Synchlo protodice
Zerene cesonia
Plutellidae
Cerostoma nemorellum
Psychidae
Apterolia helix
Fumea casta
Luffia ferchaultella
Luffia ferchaultella
Luffia lapidella
Solenobia alpicolella
Solenobia atlanca
Sexual
System
Ploidy
Female
2n
30
61
293
Male
2n
205
206
208
108
108
59
69
24
56
56
44
51
49
62
52
30
30
30
30
30
30
30
30
52
50
50
52
48
50
50
54
50
50
50
53
50
50
52
50
52
50
50
50
50
52
50
26
52
50
50
52
52
52
52
62
Sex Chrom. Source
Karyotype
ZW
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
378
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
68
377
62
61
63
62
62
61
62
377
377
377
377
378
377
377
ZO
Taxa
Solenobia clathrella
Solenobia fumosella
Solenobia generosensis
Solenobia goppensteinensis
Solenobia lichenella
Solenobia manni
Solenobia pineti
Solenobia rupicolella
Solenobia seileri
Solenobia siederi
Solenobia thomanni
Solenobia triquetrella
Solenobia triquetrella
Solenobia triquetrella
Solenobla lichenella
Talaeporia tubulosa
Pyralidae
Cadra cautella
Chino simplex
Diatraea saccharalis
Ectomyelois ceratoniae
Ephestia kuehniella
Galleria mellonella
Galleria mellonella
Galleria mellonella
Haritala ruralis
Perinephala coronata
Plodia interpunctella
Sylepta ruralis
Witlesia murana
Riodinidae
Calephelis virginiensis
Hamearis lucina
Libythea bachmanii
Libythea celtis
Saturniidae
Antherea pernyi
Antherea pernyi
Antherea yamamai
Automeris io
Caligula japonica
Callosamia promethea
Dictyoploca japonica
Platysamia cecropia
Platysamia cecropia
Platysamia columbia
Platysamia euryalis
Platysamia gloveri
Samia cynthia
Samia cynthia
Samia cynthia
Samia cynthia pryeri
Samia cynthia pryeri
Samia cynthia pryeri
Samia cynthia ricini
Samia cynthia ricini
Samia cynthia ricini
Samia cynthia ricini
Samia cynthia ricini
Samia cynthia ricini
Samia cynthia walkeri
Samia cynthia walkeri
Samia cynthia walkeri
Saturnia pavonia
Saturnia pavonia
Sexual
System
Ploidy
parth
parth
2
4
parth
4
Female
2n
61
59
60
62
60
60
60
59
62
61
25
27
26
294
Male
2n
62
62
62
62
62
62
62
62
62
62
62
62
124
62
124
60
60
58
34
62
60
60
60
60
82
20
60
62
58
Sex Chrom. Source
Karyotype
ZO
ZO
ZW
ZW
ZW
ZW
ZW
ZWW
377
377
377
377
377
377
377
377
377
377
377
377
377
378
377
378
378
377
377
378
378
377
377
378
377
377
378
377
378
90
58
62
62
377
377
377
377
66
98
62
62
62
38
62
60
62
62
62
62
26
28
26
28
28
28
28
28
28
28
28
28
26
26
26
58
58
377
377
378
377
378
377
377
377
377
377
377
377
378
378
378
377
377
377
377
377
377
377
377
377
377
377
377
377
377
ZW
ZO
ZZW
ZO
ZW
Taxa
Sexual
System
Saturnia pavonia
Saturnia pyri
Telea polyphemus
Satyridae
Agapetes galathea
Agapetes russiae
Aphantopus hyperantus
Aphantopus hyperantus
Bicylus funebris
Bicylus sandace
Bicylus vulgaris
Bicylus zinebi
Brentesia circe
Cercyonis oetus
Cercyonis pegala
Cercyonis pegala
Chazara anthe
Chazara bischoffi
Chazara briseis
Coenonympha arcania
Coenonympha arcania clorinda
Coenonympha arcania parvinsubrica
Coenonympha arcanla cephalus
Coenonympha dorus
Coenonympha gardetta darwiniana
Coenonympha gardetta lecerfi
Coenonympha gardetta philedarwiniana
Coenonympha gardetta satyrion
Coenonympha iphis
Coenonympha oedippus
Coenonympha pamphilus
Coenonympha pamphilus
Coenonympha saadi
Coenonympha tullia
Coenonympha tullia occupata
Elymnias hypermnestra hainana
Elymniopsis bammakoo
Erebia aethiopellus
Erebia aethiops
Erebia alberganus
Erebia calcarius
Erebia calcarius
Erebia callias
Erebia callias
Erebia cassfoides pseudomurina
Erebia cassioides
Erebia cassioides
Erebia cassioides aquitania
Erebia cassioides arvernensis
Erebia cassioides carmenta
Erebia cassioides carmenta
Erebia cassioides dolomitensis
Erebia cassioides illyrica
Erebia cassioides illyrica
Erebia cassioides illyromacedonica
Erebia cassioides illyromacedonica
Erebia cassioides majellana
Erebia cassioides murina
Erebia cassioides pseudocarmenta
Erebia casstoides murina
Erebia claudina
Erebia disa
Erebia discoidalis
Erebia epiphron cassiope
Erebia epipsodea
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
60
58
60
60
ZW
48
50
46
58
58
56
56
52
58
54
54
54
56
56
56
64
66
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
68
64
377
377
58
83
59
377
377
377
60
62
377
377
58
58
58
58
58
58
58
52
377
377
377
377
377
377
377
377
52
14
42
35
16
16
30
30
20
377
377
377
377
377
377
377
377
377
20
20
20
20
20
20
20
377
377
377
377
377
377
377
20
20
20
20
20
20
20
20
36
58
56
34
22
295
377
378
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Sexual
System
Ploidy
Female
2n
Male
2n
56
58
38
42
24
23
50
50
44
48
103
102
102
104
56
44
58
58
58
22
22
38
42
28
24
48
46
44
38
22
22
28
80
56
56
38
38
44
31
36
44
46
32
20
20
20
22
59
58
58
58
58
58
58
58
58
58
58
58
50
50
58
58
58
58
Erebia epistygne
Erebia eriphyle
Erebia glacialis
Erebia gorge
Erebia gorgone
Erebia hewitsonii
Erebia hispania
Erebia hispania
Erebia hispania rondoui
Erebia hispania rondoui
Erebia iranica
Erebia iranica
Erebia iranica savalanica
Erebia iranica transcaucasia
Erebia lappona
Erebia lefebyrei
Erebia ligea
Erebia magdalena
Erebia manto constans
Erebia medusa
Erebia medusa polaris
Erebia melampus
Erebia melas
Erebia meolans
Erebia mnestra
Erebia montanus
Erebia neoridas
Erebia nerine
Erebia niphonica
Erebia nivalis
Erebia nivalis
Erebia ome
Erebia ottomana
Erebia pandrose
Erebia pandrose sthennyo
Erebia pharte
Erebia prono
Erebia scipio
Erebia serotina
Erebia serotina
Erebia stirius
Erebia styx
Erebia triarius
Erebia tyndarus
Erebia tyndarus
Erebia tyndarus semimurina
Erebia warreniana
Hipparchia aelia
Hipparchia allionii
Hipparchia aristaeus
Hipparchia fatua
Hipparchia mersina
Hipparchia pisidice
Hipparchia statilinus burgeffi
Hipparchia syriaca
Hyponephele dysdora
Hyponephele lupinus
Hyponephele lycaon
Hyponephele narica
Kirina climene
Kirina roxelana
Lasiommata menava
Lethe diana
Lethe rohria daemoniaca
Lethe sicelis
296
Sex Chrom. Source
Karyotype
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Sexual
System
Ploidy
Female
2n
Maniola jurtina
Maniola jurtina
Maniola jurtina
Maniola tithonus
Melanargia lachesis
Melanitis leda
Melanitis phedima
Minois dryas
Mycalesis francisca perdiccas
Mycalesis gotama
Neope goschkevitschii
Neope muirheadi nagasawae
Oeneis asamana
Oeneis daisetsuzana
Oeneis jutta
Oeneis lucilla
Pararge aegeria
Pararge aegeria aegerides
Pararge deidamia
Pararge hiera
Pararge maera
Pararge maera adrasta
Pararge maera pannonica
Pararge megera
Pararge megera
Pararge megera lyssia
Penthema formosanum
Pseudochazara anthelea
Pseudochazara berae
Pseudochazara caucasia
Pseudochazara geyeri
Pseudochazara hippolyte
Pseudochazara mamurra
Pseudochazara mniszechii
Pseudochazara telephassa
Pseudotergumia fidia
Pyronia cecilia
Pyronia pasiphae
Pyronia tithonus
Satyrus actaea
Satyrus bryce
Satyrus hermione
Satyrus semete
Yphthima argus
Yphthima doleta
Yphthima motschulskyi
Sphingidae
Celerio euphorbiae
Celerio euphorbiae
Celerio euphorbiae
Celerio euphorbiae
Celerio galii
Celerio galii
Celerio hippophaes
Celerio lineata
Celerio livornica
Ceterio vespertilio
Deilephila elpenor
Deilephila elpenor
Deilephila elpenor lewisii
Deilephila porcellus
Laothoe populi
Laothoe populi austauti
Laothof! populi
Marumba gaschkewitschii echephron
297
Male
2n
Sex Chrom. Source
Karyotype
58
58
58
58
48
56
59
56
58
56
56
56
58
58
64
58
54
56
58
58
56
56
56
54
58
58
56
56
54
54
54
56
56
56
56
60
56
66
58
54
54
58
58
58
48
54
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
57
56
56
58
56
58
58
58
58
58
58
58
58
58
56
56
56
56
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
377
Taxa
Mimas tiliae
Mimas tiliae
Psilogramma menephron increta
Smerinthus ocellata
Smerinthus ocellata
Smerinthus ocellata
Smerinthus ocellata planus
Sphinx ligustri
Sphinx ligustri
Theretra japonica
Theretra oldenlandiae
Talaeporiidae
Talaeporia lubulosa
Tischeriidae
Tischeria angusticolella
Tischeria ekebladella
Tortricidae
Adoxophyes orana
Archips breviplicanus
Archips fuscocupreana
Bactra lacteana
Choristoneura cerasiorana
Choristoneura cerasiorana
Choristoneura fumiferana
Choristoneura pinus
Cydia pomonella
Homona magnanima
Homona menciana
Pandemis heparana
Yponomeutidae
Yponomeuta cagnagellus
Yponomeuta evonymella
Yponomeuta evonymella
Yponomeuta gigas
Yponomeuta kitabatakei
Yponomeuta malinellus
Yponomeuta padellus
Yponomeuta polistictus
Yponomeuta rorellus
Yponomeuta vigintipunctatus
Zygaenidae
Elcysma westwoodii
Illiberis nigra
Illiberis psychina
Zygaena achilleae
Zygaena ephialtes
Zygaena ephialtes coronillae
Zygaena ephialtes peucedani
Zygaena lonicerae
Zygaena purpuralis
Zygaena trifolii
Mantodea
Amorphoscelidae
Amorphoscelis indica
Cliomantis cornuta
Glabromantis nebulosa
Empusidae
Empusa egena
Empusa pauperata
Empusa spinosa
Gongylus gongyloides
Eremiaphilidae
Didymocorypha lanceolata
Didymocorypha lanceolata
Humbertiella indica
Sexual
System
Ploidy
Female
2n
58
48
46
59
56
61
61
61
61
61
61
62
Male
2n
58
58
Sex Chrom. Source
Karyotype
ZW
377
377
378
54
54
56
54
56
56
58
62
377
377
377
377
377
377
377
377
60
377
42
46
60
60
60
58
60
60
60
60
56
60
60
60
62
62
62
62
60
62
62
60
62
ZW
ZWW
ZW
ZZW
ZZW
ZZW
ZZW
ZZW
ZZW
377
64
50
50
60
60
60
60
60
60
60
377
378
377
377
377
378
377
377
377
377
378
377
377
377
378
377
378
378
377
378
378
377
378
377
377
377
377
377
377
377
377
377
377
34
26
26
33
25
25
XO
XO
XO
243
243
243
28
28
28
28
27
27
27
27
XO
XO
XO
XO
243
243
243
243
16
18
24
15
17
23
XO
XO
XO
243
243
243
298
Taxa
Humbertiella n.sp.
Humbertiella similis
Humbertiella sp.
Hymenopodidae
Acanthops falcata
Acanthops godmani
Acontiothespis cordillerae vitrea
Acontiothespis multicolor
Acontiothespis sp.
Antemna rapax
Creobroter gemmatus
Creobroter laevicollis
Creobroter urbanus
Euantissa ornata
Harpagomantis tricolor
Hestiasula brunneriana
Pseudacanthops medusa
Tithrone roseipennis
Mantidae
Aethalochroa ashmoliana
Ameles abjecta
Ameles heldreichi
Ameles sp.
Angela guianensis
Antistia sp.
Transvaal
Apterornantis bolivari
Archimantis quinquelobata
Archimantis sobrina
Arneles cypria
Bisanthe pulchripennis
Bolbe nigra
Bolbe pallida
Brunneria borealis
Callimantis antillarum
Cheddikulama straminea
Choeradodis rhombicollis
Compsothespis anomala
Compsothespis natalica
Deiphobe brunneri
Deiphobe indica
Dystacta alticeps
Haldwania liliputana
Hierodula coarctata
Hierodula patellifera
Hierodula sp.
Hierodula tenuidentata
Hierodula venosa
Hierodula ventralis
Holaptilon pusillulurn
Hoplocorypha macra
Hoplocorypha sp.
Ima fusca
Iris aratoria
Kongobatha diadernata
Leptomantis parva
Ligaria quadripunctata
Liturgusa actuosa
Liturgusa cursor
Liturgusa maya
Liturgusa sp.
Mantis octospilota
Mantis religiosa
Melliera brevipes
Miomantis sp.
Sexual
System
Ploidy
16
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
40
32
22
39
31
21
XO
XO
XO
243
243
243
20
20
15
16
16
28
28
28
28
34
26
28
20
16
19
19
XO
XO
XO
243
243
243
15
27
27
27
27
33
25
27
XO
XXY
XO
XO
XO
XO
XO
XO
15
XO
30
30
29
21
29
29
28
20
19
27
XO
XO
XO
XO
29
27
27
26
21
25
25
XO
complex XY
complex XY
XO
XO
XO
XO
17
27
31
23
23
19
27
25
15
27
27
27
27
27
27
29
35
27
34
25
25
39
23
23
33
17
21
27
27
27
15
XO
XXY
XXY
XXY
XXY
XO
XXY
XO
XO
complex XY
XXY
XXY
complex XY
XXY
complex XY
XO
XO
XO
XY
XO
XO
XO
XO
XO
XO
XO
XO
XXY
XXY
XXY
XO
28
30
28
28
27
22
30
30
28
18
28
32
30
30
20
28
26
16
28
28
28
28
28
28
30
36
28
34
26
30
40
24
24
34
18
22
28
28
28
16
299
XO
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
Taxa
Nullabora flavoguttata
Oligonyx dohrnianus
Orthodera gunnii
Orthodera ministralis
Orthoderina straminea
Oxyopsis rubicunda
Paratenodera sinensis
Parathespis humbertiana
Phyllovates tripunctata
Polyspilota aeruginosa
Polyspilota sp.
Promiopteryx granadensis
Pseudomiopteryx infuscata
Rhodamantis pulchella
Rhodamantis sp3'
Schizocephala bicornis
Sp. Sp
Sphodromantis gastrica
Sphodropoda sp4
Sphodropoda tristis
Sphodrornantis viridis
Stagomantis heterogamia
Stagornantis carolina
Stagrnatoptera septentrionalis
Statilia maculata
Tenodera aridifolia
Tenodera australasiae
Tenodera superstitiosa
Thesprotia filum
Thesprotia graminis
Toxomantis sinensis
Vates pectinicornis
Mantoididae
Mantoida schraderi
Mecoptera
Bittacidae
Bittacus italicus
Bittacus pilocornis
Bittacus stigmaterus
Boreidae
Boreus brumalis
Choristidae
Chorista australis
Nannochoristidae
Nannochorista dipteroides
Panorpidae
Neopanorpa lui
males are achiasmatic
Panorpa acuta
Panorpa anomala
Panorpa cognata
Panorpa communis
Panorpa dubia
males are achiasmatic
Panorpa emarginata
males are achiasmatic
Panorpa germanica
Panorpa sp.
males are achiasmatic
Megaloptera
Corydalidae
Chauliodes japonicus
Corydalus cornutus
Neohermes filicornis
Protohermes grandis
Sexual
System
Ploidy
28
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
30
26
26
26
28
28
32
28
28
28
20
18
28
28
30
30
28
28
28
24
28
28
27
28
28
28
28
30
30
28
28
27
19
25
25
25
27
27
31
27
27
27
19
17
27
27
27
25
27
27
27
23
27
27
complex XY
27
27
27
27
23
23
27
27
complex XY
XO
XXY
XXY
XXY
XXY
XXY
XO
XXY
XXY
XXY
XO
XO
complex XY
XXY
XO
XO
XXY
XXY
complex XY
XXY
complex XY
complex XY
243
XXY
XXY
XXY
XXY
XO
XO
XO
XXY
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
243
38
37
XO
243
26
30
32
25
29
31
XO
XO
XO
379
380
380
26
25
XXY
381
58
57
XO
382
22
243
243
243
243
243
243
243
243
382
42
41
XO
383
24
24
44
46
40
23
23
43
45
39
XO
XO
XO
XO
XO
380
380
384
384
383
40
39
XO
383
42
48
41
47
XO
XO
384
383
20
24
22
24
20
24
22
24
Xy+
Xy+
Xxyyp
Xy+
385
386
386
387
300
Taxa
Sexual
System
Microcoryphia
Machilidae
Dilta littoralis
Machilis noctis
Neuroptera
Chrysopidae
Chrysopa alba
heteromorphic
Chrysopa aspersa
heteromorphic
Chrysopa carnea
heteromorphic
Chrysopa cognatella
heteromorphic
Chrysopa flava
heteromorphic
Chrysopa flavifrons
heteromorphic
Chrysopa formosa
heteromorphic
Chrysopa intima
heteromorphic
Chrysopa japana
heteromorphic
Chrysopa japonica
heteromorphic
Chrysopa kurisakiana
heteromorphic
Chrysopa matsumurae
heteromorphic
Chrysopa parabola
heteromorphic
Chrysopa perla
heteromorphic
Chrysopa sapporensis
heteromorphic
Chrysopa septempunctata
heteromorphic
Chrysopa septempunctata cognata
heteromorphic
Chrysopa sp.
heteromorphic
Chrysopa venosa
heteromorphic
Chrysopa ventralis
heteromorphic
Chrysopa vittata
heteromorphic
Chrysopa vividana
heteromorphic
Chrysopa vulgaris
heteromorphic
Chrysopa yamamuvae
heteromorphic
Hemerobiidae
Hemerobius marginatus
heteromorphic
Mantispidae
Climaciella semihyalina
heteromorphic
Climaciella sp.
heteromorphic
Entanoneura limbata
heteromorphic
Entanoneura phithisica
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
32
30
388
388
12
12
XY
389
12
12
XY
389
14
14
XY
389
12
12
XY
389
14
14
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
10
10
XY
389
10
10
XY
389
10
10
XY
389
12
12
XY
389
12
12
XY
389
14
14
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
389
12
12
XY
390
20
20
XY
391
20
20
XY
392
20
20
XY
391
20
XXXY
391
301
Taxa
Eumantispa harmandi
heteromorphic
Leptomantispa pulchella
heteromorphic
Mantispa fuscicornis
heteromorphic
Mantispa interrupta
heteromorphic
Mantispa japonica
heteromorphic
Mantispa sayi
heteromorphic
Mantispa styriaca
heteromorphic
Mantispa uhleri
heteromorphic
Nolima pinal
heteromorphic
Plega dactyloya
Plega signata
heteromorphic
Tuberonotha stenua
heteromorphic
Zeugomantispa minuta
heteromorphic
Myrmeleontidae
Acanthaclisis japonica
heteromorphic
Brachynemurus coquilletti
heteromorphic
Brachynemurus dissimilis
heteromorphic
Brachynemurus hubbardi
heteromorphic
Brachynemurus mexicanus
heteromorphic
Brachynemurus niger
heteromorphic
Brachynemurus nigrilabris
heteromorphic
Brachynemurus schwarzi
heteromorphic
Centroclisis brachygaster
heteromorphic
Creoleon plumbea
heteromorphic
Macronemurus appendiculatus
heteromorphic
Macronemurus sp.
heteromorphic
Morter hyalinus
heteromorphic
Myrmeleon alcestris
heteromorphic
Myrmeleon californicus
heteromorphic
Myrmeleon europaeus
heteromorphic
Myrmeleon exitalis
heteromorphic
Myrmeleon formicarius
heteromorphic
Myrmeleon immaculatus
heteromorphic
Myrmeleon mexicanum
Sexual
System
Ploidy
16
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
18
XY
391
22
22
XY
391
22
22
XY
391
22
22
XY
391
20
20
XY
391
22
22
XY
391
18
18
XY
391
22
22
XY
391
18
18
XY
391
22
18
21
18
XO
XY
391
391
20
20
XY
391
20
20
XY
391
14
14
XY
393
14
14
XY
394
16
16
XY
394
14
14
XY
394
14
14
XY
394
16
16
XY
394
16
16
XY
394
14
14
XY
394
14
14
XY
393
18
18
XY
393
16
XY
393
16
16
XY
393
14
14
XY
393
14
14
XY
393
14
14
XY
394
14
14
XY
393
14
14
XY
394
14
14
XY
393
14
14
XY
394
14
14
XY
394
302
Taxa
heteromorphic
Myrmeleon obscurus
heteromorphic
Myrmeleon sagax
heteromorphic
Neuroleon sp.
heteromorphic
Palpares libelluloides
heteromorphic
Palpares sobrinus
heteromorphic
Palpares sp.
heteromorphic
Palparidius concinnus
heteromorphic
Psammoleon arizonensis
heteromorphic
Vella texana
heteromorphic
Osmylidae
Osmylus decoratus
Sisyridae
Climacia areolaris
Sisyra vicaria
heteromorphic
Odonata
Aeshnidae
Acanthaeschna anacantha
micro chromosomes present
Acanthaeschna multipunctata
micro chromosomes present
Aeshna bonariensis
Aeshna canadensis
Aeshna cf. unicolor
micro chromosomes present
Aeshna clepsydra
micro chromosomes present
Aeshna coerulea
Aeshna confusa
Aeshna cornigera planaltica
Aeshna crenata
micro chromosomes present
Aeshna cyanea
micro chromosomes present
Aeshna diffinis diffinis
micro chromosomes present
Aeshna grandis
Aeshna grandis
Aeshna intricata
micro chromosomes present
Aeshna juncea
also XY
Aeshna juncea
Aeshna mixta
Aeshna palmate
micro chromosomes present
Aeshna peralta
micro chromosomes present
Aeshna serrata fennica
micro chromosomes present
Aeshna squamata
Aeshna subartica elisabethae
micro chromosomes present
Aeshna umbrosa occidentalis
Aeshna umbrosa umbrosa
Sexual
System
Ploidy
28
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
14
14
XY
393
14
14
XY
393
16
16
XY
393
26
26
XY
393
22
22
XY
393
24
24
XY
393
18
18
XY
393
16
16
XY
394
14
14
XY
394
16
15
XXY
395
14
14
13
14
XO
XY
396
396
28
27
XO
397
27
XO
397
26
28
28
26
27
27
NeoXY
XO
XO
398
399
397
28
27
XO
397
24
28
16
28
24
27
16
27
NeoXY
XO
XY
XO
398
398
398
397
28
27
XO
397
22
21
XO
397
26
26
20
26
25
19
NeoXY
XO
XO
398
400
397
27
27
XO
397
26
28
28
26
27
27
NeoXY
XO
XO
398
401
397
28
27
XO
397
26
26
NeoXY
397
24
28
24
27
NeoXY
XO
398
397
28
28
27
27
XO
XO
399
399
303
Taxa
Aeshna verticalis
micro chromosomes present
Aeshna virdis
micro chromosomes present
Aeshna walkeri
micro chromosomes present
Anax imperator
micro chromosomes present
Anax junius
micro chromosomes present
Anax longipes
micro chromosomes present
Anax parthenope
micro chromosomes present
Basiashna janata
Boyeria maclachlani
micro chromosomes present
Boyeria vionsa
Castoraeschna castor
micro chromosomes present
Coryphaeschna adnexa
Gynacantha japonica
micro chromosomes present
Hemianax papuensis
micro chromosomes present
Hemianax ephippiger
micro chromosomes present
Oplonaeshna armata
micro chromosomes present
Planaeschna milnei
micro chromosomes present
Agrionidae
Agrion aequabile
Agrion maculatum
Hetaerina americana
Calopterygidae
Anaciagrion cornelia
micro chromosomes present
Calopteryx aequabile
micro chromosomes present
Calopteryx atrata
micro chromosomes present
Calopteryx maculata
micro chromosomes present
Calopteryx splendens caprai
micro chromosomes present
Calopteryx splendens splendens
Calopteryx virgo japonica
micro chromosomes present
Calopteryx virgo meridionalis
Calopteryx virgo padana
micro chromosomes present
Calopteryx virgo virgo
micro chromosomes present
Matrona basalaris
Mnais costalis
micro chromosomes present
Mnais strigata
micro chromosomes present
Coenagrionidae
Acanthagrion ascendens
micro chromosomes present
Acanthagrion chacoense
micro chromosomes present
Aeolagrion foliaceum
Sexual
System
Ploidy
26
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
27
XO
397
26
26
NeoXY
397
28
27
XO
399
28
27
XO
399
28
27
XO
399
28
27
XO
399
28
27
XO
397
26
28
25
27
XO
XO
399
397
28
28
27
27
XO
XO
399
397
28
28
27
27
XO
XO
397
397
28
27
XO
401
14
13
XO
397
28
27
XO
397
28
27
XO
401
26
26
26
25
25
25
XO
XO
XO
399
399
399
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
25
26
XO
25
397
XO
397
27
26
26
25
XO
XO
397
397
26
25
XO
397
26
26
25
25
XO
XO
397
397
26
25
XO
397
28
27
XO
397
28
27
XO
397
28
27
XO
397
304
Taxa
Agrion funebris
Agrion sedula
Agrion violacea
Agrion vivida
Amphiagrion abbreviatum
Argia violacea
Argia vivida
Ceratura capreola
Cercion lindeni
micro chromosomes present
Ceriagrion cerinorubellum
micro chromosomes present
Ceriagrion coromandeliamum
micro chromosomes present
Ceriagrion fallax
micro chromosomes present
Ceriagrion rubiae
micro chromosomes present
Ceriagrion tenellum tenellum
micro chromosomes present
Chromagrion conditum
Coenagrion armatum
Coenagrion hastulatum
Coenagrion hieroglyphicum
micro chromosomes present
Coenagrion pulchellum
Coenagrion resolutum
Coenagrion sp.
micro chromosomes present
Diceratobasis macrogaster
micro chromosomes present
Enallagma aspersum
Enallagma boreale
Enallagma carunculatum
Enallagma civile
Enallagma cyathigerum
Enallagma cyathigerum
Enallagma cyathigerum
single specimen with fission
Enallagma ebrium
Enallagma praevarum
Erythromma najas
Ischnura cervula
Ischnura cf. ultima
Ischnura denticollis
Ischnura elegans
Ischnura fluviatilis
Ischnura perparva
Ischnura senegalensis
micro chromosomes present
Ischnura verticalis
Leptagrion macrurum
Magalagrion oahuense
micro chromosomes present
Mortonagrion selenion
micro chromosomes present
Nehalennia irene
Nehalennia speciosa
Pseudagrion australasiae
micro chromosomes present
Pseudagrion decorum
micro chromosomes present
Pseudagrion microcephalum
micro chromosomes present
Pseudagrion rubriceps
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
28
28
28
28
28
28
28
28
27
27
27
27
27
27
27
27
27
XO
XO
XO
XO
XO
XO
XO
XO
XO
397
397
397
397
399
399
399
397
397
28
27
XO
397
28
27
XO
399
28
27
XO
397
28
27
XO
397
28
27
XO
397
28
28
28
28
27
27
27
27
XO
XO
XO
XO
399
397
397
397
28
28
28
27
27
27
XO
XO
XO
397
399
397
28
27
XO
397
28
28
28
28
28
28
27
27
27
27
27
27
28
XO
XO
XO
XO
XO
XO
399
399
399
399
399
399
402
28
28
28
28
28
28
28
28
28
28
27
27
27
27
27
27
27
27
27
27
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
399
399
397
399
397
399
397
397
399
397
28
30
28
27
30
27
XO
NeoXY
XO
399
403
397
28
27
XO
397
28
28
28
27
27
27
XO
XO
XO
399
397
397
28
27
XO
397
28
27
XO
397
28
27
XO
397
305
Taxa
micro chromosomes present
Pseudagrion spencei
micro chromosomes present
Pyrrhosoma nymphula
Tigriagrion aurantinigrum
Zoniagrion exclamationis
Cordulegasteridae
Antogaster sieboldii
micro chromosomes present
Cordulegaster boltoni
micro chromosomes present
Cordulegaster deserticola
micro chromosomes present
Cordulegaster diastatops
micro chromosomes present
Cordulegaster diastatops
Cordulegaster dorsalis
micro chromosomes present
Cordulegaster maculatus
micro chromosomes present
Cordulidae
Cordulia shurtleffi
Didymops transversa
micro chromosomes present
Dorocordulia libera
Dorocordulia libera
single specimen with fusion
Ephitheca canis
Ephitheca cynosura
Ephitheca cynosura
single specimen with fusion
Ephitheca semiaquea
Ephitheca spinigera
Macromia magnifica
micro chromosomes present
Somatochlora metallica
Somatochlora semicircularis
Corduliidae
Cordulia aenea
Epicodulia princeps
micro chromosomes present
Epitheca canis
micro chromosomes present
Epitheca cynosura
Epitheca semiaqua
Epitheca spinigera
micro chromosomes present
Epopthalmia frontalis frontalis
micro chromosomes present
Somatochlora flavomaculata
Somatochlora metallica
Somatochlora uchidai
micro chromosomes present
Somatochlora viridiaenea
Tetragoneuria petechialis
Tetragoneuria spinigera
Epallagidae
Epallage fatime
Epiophlebiidae
Epiophlebia superstes
Gomphidae
Anisogomphus bivittatus
micro chromosomes present
Aphylla edentata
Aphylla producta
Sexual
System
Ploidy
26
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
28
27
XO
397
28
28
28
27
27
27
XO
XO
XO
397
397
399
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
26
25
25
XO
XO
399
399
26
25
XO
399
26
26
25
25
XO
XO
399
399
14
13
14
XO
399
402
26
22
25
21
22
XO
XO
399
399
402
26
26
26
25
25
25
XO
XO
XO
399
399
399
26
26
25
25
XO
XO
400
399
26
26
25
25
XO
XO
397
397
26
25
XO
397
21
26
26
20
25
25
XO
XO
XO
397
397
397
25
XO
397
26
26
26
25
25
25
XO
XO
XO
397
397
397
26
22
28
25
21
27
XO
XO
XO
397
397
397
26
25
XO
404
26
25
XO
397
24
23
XO
401
24
24
23
23
XO
XO
401
401
306
Taxa
Dromogomphus spinosus
micro chromosomes present
Dromogomphus spoliatus
micro chromosomes present
Epigomphus llama
Erpetogomphus ophibolus
micro chromosomes present
Gomphoides sp.
Gomphus graslini
micro chromosomes present
Gomphus confraternus
micro chromosomes present
Gomphus exilis
micro chromosomes present
Gomphus exilis
Gomphus lentulus
Gomphus lividus
micro chromosomes present
Gomphus melaenops
single specimen with fusion
Gomphus melampus bifasciatus
single specimen with fusion
Gomphus militaris
Gomphus plagiatus
micro chromosomes present
Gomphus scudderi
Gomphus spicatus
micro chromosomes present
Gomphus submedianus
Ictinogomphus rapax
Octogomphus specularis
Onychogomphus forcipatus
Onychogomphus forcipatus
Ophiogomphus bison
Ophiogomphus bison
Ophiogomphus bison
single specimen with fission
Ophiogomphus colubrinus
Ophiogomphus rupinsulensis
Ophiogomphus serpentinus
Phyllocycla sp.
Progomphus borealis
Progomphus borealis
Progomphus intricatus
Progomphus obscurus
Progomphus obscurus
Progomphus phyllochromus
micro chromosomes present
Sieboldius albardae
micro chromosomes present
Trigolnphus melampus bifasciatus
Trigomphus citimus tabei
micro chromosomes present
Trigomphus interruptus
micro chromosomes present
Trigomphus melampus
Trigomphus unifasciatus
Gomphidae
Davidius nanus
Dromogomphus spinosus
Dromogomphus spoliatus
Erpetogomphus designatus
micro chromosomes present
Erpetogomphus diadophis
Gomphus confraternus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
23
XO
399
24
23
XO
399
20
24
19
23
XO
XO
401
397
24
24
23
23
XO
XO
401
401
24
23
XO
399
24
23
XO
399
24
24
23
23
XO
XO
402
399
399
24
24
20
402
402
24
24
23
23
XO
XO
399
399
24
24
23
23
XO
XO
399
399
24
24
24
24
23
23
23
24
24
25
23
24
24
XO
XO
XO
XO
XO
XO
XO
399
401
399
401
397
399
397
402
24
24
24
24
24
24
24
24
24
24
23
23
23
23
23
23
23
23
23
23
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
399
399
397
401
399
401
401
399
401
401
24
23
XO
401
19
22
XO
21
401
XO
397
20
19
XO
397
20
22
19
21
XO
XO
401
397
24
24
24
24
23
23
23
23
XO
XO
XO
XO
401
401
401
401
24
24
23
23
XO
XO
401
401
307
Taxa
Gomphus exilis
Gomphus graslini
Gomphus lentulus
Gomphus lividus
Gomphus melaenops
Gomphus militaris
Gomphus pallidus
Gomphus plagiatus
Gomphus postocularis
micro chromosomes present
Gomphus scudderi
Gomphus spicatus
Gomphus submedianus
Nihonogomphus viridis
micro chromosomes present
Octogomphus specularis
Onychogomphus forcipatus
Ophiogolnphus serpentinus
Ophiogomphus colubrinus
Ophiogomphus occidentalis
Ophiogomphus rupinsulensis
Ophiogonlphus bison
Stylogomphus suzukii
micro chromosomes present
Trigomphus citimus tabei
Hetaerinidae
Hetaerina americana
micro chromosomes present
Hetaerina charca
micro chromosomes present
Hetaerina rosea
micro chromosomes present
Hetaerina sanguinea
Hetaerina titia
micro chromosomes present
Hetaerina tricolor
micro chromosomes present
Hetaerina vulnerata
micro chromosomes present
Lestidae
Chalcolestes viridis
micro chromosomes present
Lestes virens
Lestes viridis
Lestes congener
micro chromosomes present
Lestes disjunctus
Lestes dryas
micro chromosomes present
Lestes forcipatus
Lestes forficula
micro chromosomes present
Lestes rectangularis
micro chromosomes present
Lestes simulatrix
micro chromosomes present
Lestes sponsa
micro chromosomes present
Lestes stultus
micro chromosomes present
Lestes vidua
micro chromosomes present
Lestes virens vestallis
micro chromosomes present
Sympecma fusca
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
24
24
24
24
24
24
24
24
23
23
23
23
23
23
23
23
23
NeoXY
XO
XO
XO
XO
XO
XO
XO
XO
401
401
401
401
401
401
401
401
401
24
24
24
24
23
23
23
23
XO
XO
XO
XO
401
401
401
401
24
24
24
24
24
24
24
24
23
25
23
23
23
23
25
23
XO
NeoXY
XO
XO
XO
XO
NeoXY
XO
401
401
401
401
401
401
401
401
22
21
XO
401
26
25
XO
397
26
25
XO
397
28
27
XO
397
26
26
25
25
XO
XO
397
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
26
26
25
25
25
XO
XO
XO
401
401
399
26
26
25
25
XO
XO
399
399
22
26
21
25
XO
XO
399
397
26
25
XO
399
26
25
XO
397
26
25
XO
397
26
25
XO
399
26
25
XO
397
26
25
XO
397
26
25
XO
397
308
Taxa
Sexual
System
micro chromosomes present
Libellulidae
Acisoma panorpoides panorpoides
micro chromosomes present
Brachydiplax chalybea
micro chromosomes present
Brachydiplax farinosa
micro chromosomes present
Brachydiplax sobrina
micro chromosomes present
Brachythemis contaminata
micro chromosomes present
Bradinopyga geminata
micro chromosomes present
Brechmorhoga mendax
Brechmorrhoga nubecula
micro chromosomes present
Brechmorrhoga pertinax peruviana
Canaphila vibex
micro chromosomes present
Cannacria elisa
micro chromosomes present
Cannacria fasciata
micro chromosomes present
Cannacria gravida
micro chromosomes present
Cannacria herbida
micro chromosomes present
Celithemis elisa
Celithemis fasciata
Crocothemis erythraea
micro chromosomes present
Crocothemis servilia
also XY
Dasythemis esmeralda
micro chromosomes present
Dasythemis venosa
micro chromosomes present
Diastatops intensa
micro chromosomes present
Diastatops obscura
micro chromosomes present
Diplacodes bipunctata
Diplacodes haematodes
Diplacodes leferbvrei
micro chromosomes present
Diplacodes nebulosa
micro chromosomes present
Diplacodes trivialis
micro chromosomes present
Dythemis cannacroides
micro chromosomes present
Dythemis fugax
micro chromosomes present
Dythemis rufinervis
micro chromosomes present
Dythemis velox
micro chromosomes present
Erythemis attala
Erythemis collocata
micro chromosomes present
Erythemis plebeja
Erythemis simplicicollis
micro chromosomes present
Erythrodiplax basalis basalis
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
25
XO
397
26
25
XO
397
26
26
NeoXY
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
26
25
25
XO
XO
399
397
26
26
25
25
XO
XO
397
397
26
25
XO
397
26
25
XO
397
26
25
XO
399
26
25
XO
397
26
26
26
25
25
25
XO
XO
XO
399
399
397
25
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
28
25
26
27
24
25
XO
XO
XO
397
397
397
26
25
XO
397
26
25
XO
397
24
23
XO
397
26
25
XO
399
26
25
XO
397
26
25
XO
397
26
26
25
25
XO
XO
397
399
26
26
25
25
XO
XO
397
399
26
25
XO
397
309
Taxa
Sexual
System
Erythrodiplax berenice
micro chromosomes present
Erythrodiplax castanea
Erythrodiplax connata connata
micro chromosomes present
Erythrodiplax connata fusca
Erythrodiplax fervida
micro chromosomes present
Erythrodiplax justiniana
micro chromosomes present
Erythrodiplax media
micro chromosomes present
Erythrodiplax melanorubra
micro chromosomes present
Erythrodiplax paraguayensis
micro chromosomes present
Erythrodiplax umbrata
micro chromosomes present
Erythrodiplax unimaculata
micro chromosomes present
Ladona julia
micro chromosomes present
Lathrecista asiatica
micro chromosomes present
Lepthemis vesiculosa
Leucorrhinia albifrons
micro chromosomes present
Leucorrhinia dubia
Leucorrhinia frigida
Leucorrhinia frigida
Leucorrhinia frigida
single specimen with fusion
Leucorrhinia glacialis
micro chromosomes present
Leucorrhinia hudsonica
micro chromosomes present
Leucorrhinia intacta
micro chromosomes present
Leucorrhinia pectoralis
micro chromosomes present
Leucorrhinia proxima
micro chromosomes present
Leucorrhinia rubicunda
Libellula angelina
micro chromosomes present
Libellula axilena
Libellula composita
micro chromosomes present
Libellula croceipennis
micro chromosomes present
Libellula cyanea
Libellula depressa
Libellula flavida
micro chromosomes present
Libellula florensis
micro chromosomes present
Libellula incesta
Libellula luctuosa
Libellula pulchella
micro chromosomes present
Libellula quadrimaculata
Libellula quadrimaculata quadrimaculata
micro chromosomes present
Libellula saturata
micro chromosomes present
Ploidy
26
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
25
XO
399
25
25
24
XO
XO
397
397
26
26
25
25
XO
XO
399
397
26
25
XO
397
22
21
XO
397
26
25
XO
397
24
23
XO
397
26
25
XO
399
26
25
XO
397
26
25
XO
399
26
25
XO
397
26
26
25
25
XO
XO
397
397
26
24
23
25
23
22
24
XO
XO
XO
397
399
397
402
26
25
XO
399
26
25
XO
399
26
25
XO
399
26
25
XO
397
26
25
XO
399
26
26
25
25
XO
XO
397
397
24
26
23
25
XO
XO
397
399
26
25
XO
399
26
25
26
25
24
25
XO
XO
XO
399
397
399
26
25
XO
399
26
26
26
25
25
25
XO
XO
XO
399
397
399
26
26
25
25
XO
XO
399
397
26
25
XO
399
310
Taxa
Libellula semifasciata
micro chromosomes present
Libellula vibrans
micro chromosomes present
Lyriothemis pachygastra
micro chromosomes present
Macrothemis hemichlora
Macrothemis mortoni
micro chromosomes present
Macrothemis musiva
micro chromosomes present
Macrothemis declivata
micro chromosomes present
Macrothemis imitans imitans
micro chromosomes present
Miathyria marcella
micro chromosomes present
Micrathyria atra
micro chromosomes present
Micrathyria cf. eximia
Micrathyria didyma
micro chromosomes present
Micrathyria hageni
micro chromosomes present
Micrathyria iheringi
micro chromosomes present
Micrathyria laevigata
micro chromosomes present
Micrathyria ocellata dentiens
micro chromosomes present
Micrathyria sp.
Micrathyria spuria
micro chromosomes present
Nannothemis bella
micro chromosomes present
Nesciothemis farinosum
micro chromosomes present
Nesogonia blackburni
micro chromosomes present
Neurothemis tullia tullia
micro chromosomes present
Orthemis levis
single specimen with fission
Orthemis biolleyi
Orthemis cultiformis
micro chromosomes present
Orthemis ferruginea
Orthemis ferruginea
micro chromosomes present
Orthemis ferruginea
micro chromosomes present
Orthemis levis
NeoXY probable
Orthetrum albistylum albistylum
micro chromosomes present
Orthetrum albistylum speciosum
micro chromosomes present
Orthetrum azureum
micro chromosomes present
Orthetrum brachiale
Orthetrum brunneum
micro chromosomes present
Orthetrum cancellatum
micro chromosomes present
Orthetrum coerulescens
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
25
XO
399
26
25
XO
399
26
25
XO
397
6
26
6
25
NeoXY
XO
398
398
26
25
XO
398
24
23
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
22
26
21
25
XO
XO
397
397
26
25
XO
397
24
23
XO
397
26
25
XO
397
26
25
XO
397
24
26
23
25
XO
XO
397
397
26
25
XO
399
26
25
XO
397
26
25
XO
397
28
28
NeoXY
397
6
402
24
24
23
23
XO
XO
397
397
24
24
23
23
XO
XO
399
397
10
10
NeoXY
397
6
402
26
25
XO
397
26
25
XO
397
26
25
XO
397
22
26
21
25
XO
XO
397
397
26
25
XO
397
26
25
XO
397
311
Taxa
Sexual
System
Orthetrum glaucum
micro chromosomes present
Orthetrum japonicum
micro chromosomes present
Orthetrum pruinosum neglectum
micro chromosomes present
Orthetrum sabina
micro chromosomes present
Orthetrum taeniolatum
micro chromosomes present
Orthetrum triangulare melania
micro chromosomes present
Orthetrum triangulare triangulare
micro chromosomes present
Pachydiplax longipennis
micro chromosomes present
Pantala flavescens
micro chromosomes present
Pantala hymenea
micro chromosomes present
Perithemis cornelia
Perithemis domitia
micro chromosomes present
Perithemis electra
Perithemis lais
Perithemis mooma
micro chromosomes present
Perithemis seminole
micro chromosomes present
Perithemis sp.
Planiplax sanguineventris
micro chromosomes present
Plathemis lydia
micro chromosomes present
Potamarcha obscura
micro chromosomes present
Pseudothemis zonata
Rhodopygia cardinalis
micro chromosomes present
Rhyothemis fuliginosa
Rhyothemis variegata
micro chromosomes present
Scapanea frontalis
micro chromosomes present
Sympetrum costiferum
micro chromosomes present
Sympetrum danae
micro chromosomes present
Sympetrum eroticum eroticum
Sympetrum flaveolum
micro chromosomes present
Sympetrum frequens frequens
Sympetrum madidum
micro chromosomes present
Sympetrum meridionale
micro chromosomes present
Sympetrum obtrusum
micro chromosomes present
Sympetrum parvulum
micro chromosomes present
Sympetrum pedemontanum elatum
micro chromosomes present
Sympetrum robicundulum
micro chromosomes present
Sympetrum sanguineum
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
25
XO
397
26
25
XO
397
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
399
26
25
XO
397
26
25
XO
399
26
26
25
25
XO
XO
397
397
26
18
26
25
17
25
XO
XO
XO
397
397
397
26
25
XO
397
26
26
25
25
XO
XO
397
399
26
25
XO
399
26
25
XO
397
24
26
23
25
XO
XO
397
397
26
26
25
XO
402
397
26
25
XO
397
26
25
XO
399
26
25
XO
399
21
26
XO
25
397
XO
397
23
26
XO
25
397
XO
399
26
25
XO
397
26
25
XO
399
26
25
XO
397
26
25
XO
397
26
25
XO
399
26
25
XO
397
26
22
24
312
Taxa
micro chromosomes present
Sympetrum semicinctum
micro chromosomes present
Sympetrum striolatum
Sympetrum vicinum
micro chromosomes present
Sympetrum vulgatum
micro chromosomes present
Tarnetrum corruptum
micro chromosomes present
Tarnetrum illotum
micro chromosomes present
Tauriphila australis
micro chromosomes present
Tauriphila azteca
micro chromosomes present
Tramea abdominalis
Tramea basilaris
Tramea burmeisteri
micro chromosomes present
Tramea carolina
micro chromosomes present
Tramea cophysa
micro chromosomes present
Tramea lacerata
Tramea limbata
micro chromosomes present
Tramea virginia
micro chromosomes present
Trithemis aurora
micro chromosomes present
Trithemis pallidinervis
micro chromosomes present
Urothemis signata signata
micro chromosomes present
Zenithoptera viola
micro chromosomes present
Macrodiplactidae
Aethriamanta brevipennis
micro chromosomes present
Megapodagrionidae
Heteragrion flavidorsum
Heteragrion inca
micro chromosomes present
Megapodagrion contortum
Megapodagrion macropus
Megapodagrion setigerum
Philogenia carrilica
micro chromosomes present
Petaluridae
Tachopteryx thoreyi
micro chromosomes present
Tanypteryx hageni
micro chromosomes present
Tanypteryx pryeri
micro chromosomes present
Uropetala carovei
micro chromosomes present
Platycnemididae
Copera annulata
micro chromosomes present
Platycnemis pennipes
Platystictidae
Palaemnema paulina
micro chromosomes present
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
26
25
XO
399
26
26
25
25
XO
XO
397
399
26
25
XO
397
26
25
XO
399
26
25
XO
399
26
25
XO
397
26
25
XO
399
26
26
26
25
25
25
XO
XO
XO
397
399
397
26
25
XO
399
26
25
XO
397
26
26
25
25
XO
XO
399
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
399
26
25
XO
397
26
25
XO
397
26
26
25
25
XO
XO
397
397
27
26
26
26
26
25
25
25
XO
XO
XO
XO
397
397
397
397
20
19
XO
397
18
17
XO
399
18
17
XO
397
18
17
XO
397
26
25
XO
397
26
25
XO
397
26
25
XO
397
313
Taxa
Polythoridae
Cora irene
micro chromosomes present
Polythore boliviana
micro chromosomes present
Protoneuridae
Epipleoneura sp.
Neoneura rubiventris
micro chromosomes present
Pseudolestidae
Hypolestes clara
Pseudostigmatidae
Mecistogaster sp. 1
micro chromosomes present
Mecistogaster sp. 2
Orthoptera
Acrididae
Abracris dilecta
Abracris sp. B
Adimantus cubiceps
Adimantus ornatissimus
Albretchia palpata
Aleuas gracilis
Aleuas lineatus
Aleuas sp. 1
Aleuas sp. 2
Aleuas sp. 3
Aleuas vitticollis
Allotruxalis sp.
Allotruxalis strigata
Amblytropidia australis
Apacris rubrithorax
Apacris sp. 1
Apolobamba prope pulchra
Atrachelacris olivaceus
Atrachelacris unicolor
Belosacris coccineipes
Bucephalacris bohlsii
Carbonellacris grossa
Chlorus bolivianus
Chlorus borrelli
Chlorus sp. 1
Chlorus vittatus
Cocytolettix argentina
Cocytolettix intermedia
Cocytolettix pulchripennis
Cocytolettix sp. 1
Cornops aquaticum
Cornops frenatum
Covasacris sp.
Dichromatos corupa
Dichromatos lilloanus
Dichromatos montanus
Dichromatos schrottkyi
Dichromorpha australis
Dichroplus alejomesai
Dichroplus auriventris
Dichroplus bergi
Dichroplus conspersus
Dichroplus democraticus
Dichroplus dubius
Dichroplus elongatus
Dichroplus exilis
Dichroplus fuscus
Dichroplus fuscus.2
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
23
XO
397
24
23
XO
397
28
28
27
27
XO
XO
397
397
18
17
XO
397
30
29
XO
397
12
12
NeoXY
397
24
22
24
24
24
20
20
20
22
20
20
24
24
24
24
24
24
22
22
24
22
24
20
22
20
24
24
24
24
24
24
24
24
23
21
23
23
23
20
20
20
22
20
19
23
23
23
23
23
23
22
22
23
21
23
19
21
19
23
23
23
23
23
23
23
23
24
24
24
22
24
24
22
24
24
24
20
23
23
23
22
23
23
21
23
23
23
19
XO
XO
XO
XO
XO
XY
XY
XY
XY
XY
XO
XO
XO
XO
XO
XO
XO
XY
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XXY
XXY
XXY
XXY
XO
XO
XO
XY
XO
XO
XXY
XO
XO
XO
XO
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
406
406
406
406
405
405
405
405
405
405
405
405
405
405
405
314
Taxa
Dichroplus maculipennis
Dichroplus mantiqueirae
Dichroplus misionensis
Dichroplus obsurus
Dichroplus paraelongatus
Dichroplus paraguayensis
Dichroplus patruelis
Dichroplus piceomaculatus
Dichroplus porteri
Dichroplus pratensis
Dichroplus pseudopunctulatus
Dichroplus punctulatus
Dichroplus robustulus
Dichroplus robustus
Dichroplus schulzi
Dichroplus silveiraguidoi
Dichroplus sp. 11
Dichroplus sp. 12
Dichroplus sp. 13
Dichroplus sp. 14
Dichroplus sp. 15
Dichroplus sp. 16
Dichroplus vittatus
Dichroplus vittatus.2
Dichroplus vittigerum
Eucephalacris borellii
Eujivarus fusiformis
Eujivarus sp. A
Eujivarus sp. B
Eujivarus sp. C
Eujivarus vittatus
Eulampiacris leucoptera
Euplectrotettix sp. 1
Euplectrotettix sp. 2
Euplectrotettix sp. 3
Eurotettix lilloanus
Eurotettix minor
Eurotettix schrottkyi
Eurotettix sp. 1
Eurotettix sp. 2
Eusitalces sp. A
Eusitalces vulneratus
Eutryxalis sp.
Fenestra bohlsii
Haroldgrantia lignosa
Hyalopteryx rufipennis
Isonyx paragauyensis
Isonyx sp. 1
Jodacris chapadensis
Jodacris ferrugineus
Jodacris furcillata
Lamiacris migroguttata
Laplatacris dispar
Laplatacris sp. 1
Leiotettix flavipes
Leiotettix politus
Leiotettix politus
Leiotettix politus.2
Leiotettix pulcher
Leiotettix sanguineus
Leiotettix sp. 1
Leiotettix sp. 2
Leiotettix sp. 3
Leiotettix viridis
Leptysma dorsalis
Sexual
System
Ploidy
24
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
24
24
18
24
20
22
22
22
20
23
24
24
22
24
8
24
20
20
22
20
22
20
18
18
24
22
22
22
22
24
24
24
24
24
22
22
22
22
22
24
24
24
24
24
24
24
24
20
20
20
24
24
24
22
23
23
23
18
23
20
21
22
22
19
XO
23
23
21
23
8
23
20
20
22
20
22
20
18
18
23
21
21
21
21
23
23
23
23
23
21
22
21
21
21
23
23
23
23
23
23
23
23
19
19
19
23
23
23
22
14
14
22
24
16
18
24
24
24
14
13
22
23
15
18
23
23
23
XO
XO
XO
XY
XO
XY
XO
XY
XY
XO
405
XO
XO
XXY
XO
XY
XO
XY
XY
XY
XY
XY
XY
XY
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XXY
XY
XXY
XXY
XXY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XXY
XY
XXY
XY
XO
XXY
XY
XO
XO
XO
315
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
406
405
405
405
405
405
405
405
405
405
Taxa
Leptysmina pallida
Leptysmina sp. 1
Machaeropeles rostratum
Mastusia quadricarinata
Meloscirtus montanus
Meloscirtus sp. 1
Metaleptea brevicornis adspersa
Nahuelia rubriventris
Neopedies brunneri
Neopedies sp. 1
Neopedies sp. 2
Neopedies sp. 3
Neopedies sp. 4
Notopomala glaucipes
Omalotettix obliquum
Ommatolampis perspicillata
Orphula sp.
Orphulella concinnula
Orphulella punctata
Orphulella sp.
Orphulina pulchella
Osmilia flavolineata
Oxyblepta sp.
Oxybleptella sagitta
Paraorphula graminea
Parapellopedon instabilis
Parapellopedon sp.
Parascopas exertus
Parascopas obesus
Parascopas sanguineus
Parascopas similis
Paratylotropidia morsei
Paropaon laevifrons
Paropaon pilosus tingomariae
Pedies andeanus
Pedies sp. 1
Pedies sp. 2
Pedies sp. 3
Pedies sp. 4
Propedies bilobus
Propedies bipunctatus
Propedies fusiformis
Propedies olivaceus
Propedies sanguineus
Propedies sp. 1
Pseudoscopas nigrigena
Pseudoscopas sp. 1
Pseudoscopas sp. 2
Pseudoscopas sp. 3
Pseudoscopas sp. 4
Pseudoscopas sp. 5
Pseudoscopas sp. 6
Pseudoscopas sp. 7
Pseudoscopas sp. 8
Psiloscirtus bolivianus
Psiloscirtus olivaceus
Psiloscirtus sp. A
Pycnosarcus atavus
Ronderosia dubius
Ronderosia robustus
Schistocerca cancellata
Schistocerca flavofasciata
Schistocerca pallens
Schistocerca paraensis
Schistocerca sp.
Sexual
System
Ploidy
24
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
24
24
24
24
24
23
24
24
24
24
24
24
24
22
24
24
24
24
24
24
24
24
24
24
24
24
22
24
24
24
23
23
23
23
23
23
XO
23
23
23
23
23
23
23
21
23
23
23
23
23
23
23
23
23
23
23
23
21
23
23
23
24
24
22
22
22
22
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
18
23
23
21
21
21
21
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
17
24
24
24
24
24
23
23
23
23
23
XO
XO
XO
XO
XO
XO
405
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XXY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XXY
XXY
XO
XO
XO
XO
XO
316
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
406
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
406
406
405
405
405
405
405
Taxa
Scotussa cliens
Scotussa daguerrei
Scotussa daguerrei
Scotussa delicatula
Scotussa impudica
Scotussa lemniscata
Scotussa liebermanni
Scotussa sp. 1
Scyllina humilis
Scyllina signatipennis
Scyllina sp.
Scyllinops brunneri
Scyllinops pallida
Scyllinops sp. 1
Scyllinops sp. 2
Silvitettix concolor
Sinipta acuta
Sinipta dalmani
Sinipta maldonadoi
Sitalces dorsalis
Sitalces infuscatus
Sitalces volxemi
Staurorhectus longicornis
Stenopola bohlsii
Stenopola boliviana
Stenopola dorsalis
Stenopola pallida
Stenopola rubrifons
Stereotettix sp. 1
Tetrataenia surinama
Trimerotropis ochraceipennis
Trimerotropis pallidipennis
Xiphiola borellii
Zygoclistron falconinum
Zygoclistron nasicum
Zygoclistron trachystictum
Lentulidae
Karruacris browni
Ommexechidae
Aucacris bullocki
Calcitrena maculosa
Clarazella bimaculata
Clarazella patagona
Conometopus sulcaticollis
Cumainocloidus cordillerae
Descampsacris serrulata
Graea horrida
Neuquina fictor
Ommexecha sp.
Ommexecha virens
Ommexechia germari
Pachyosa signata
Spathalium audouini
Tetrixocephalus chilensis
Tetrixocephalus micropterus
Tetrixocephalus sergioi
Tetrixocephalus sp.
Tetrixocephalus willemsei
Pauliniidae
Marellia remipes
Paulinia acuminata
Pyrgomorphidae
Omura congrua
Romaleidae
Alcamenes clarazianus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
22
21
22
16
24
24
22
24
24
22
24
24
24
24
24
24
24
24
24
24
24
20
24
24
24
24
22
24
24
20
24
24
24
20
20
20
21
16
23
23
21
23
23
22
23
23
23
23
23
23
23
23
23
23
23
19
23
23
23
23
21
23
23
19
23
23
23
20
20
20
XO
XXY
XXY
XY
XO
XO
XO
XO
XO
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XY
XY
405
406
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
XXY
406
24
24
24
24
26
24
24
24
22
24
24
22
22
22
24
24
24
24
22
23
23
23
23
25
23
23
23
22
23
23
21
22
22
23
23
23
23
22
XO
XO
XO
XO
XO
XO
XO
XO
XY
XO
XO
XO
XY
XY
XO
XO
XO
XO
XY
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
24
24
23
23
XO
XO
405
405
20
19
XO
405
24
23
XO
405
317
Taxa
Antandrus viridus
Chariacris miniacea
Chromacris miles
Chromacris peruviana
Chromacris speciosa
Coryacris angustipennis
Diponthus clarazianus
Diponthus communis
Diponthus dispar
Diponthus electus
Diponthus maculiferus
Diponthus prope communis
Diponthus sp.
Elaeochlora basalis
Elaeochlora brachyptera
Elaeochlora sp.
Elaeochlora trilineata
Elaeochlora viridicata
Eutropidacris collares
Prionolopha serrata
Procolpia minor
Securigera acutangula
Xestotrachelus robustus
Xyleus attenuatus
Xyleus discoideus
Xyleus gracilis
Xyleus insignis
Xyleus laevipes
Xyleus modestus
Xyleus sp. 1
Xyleus sp. 2
Xyleus sp. 3
Zoniopoda hempeli
Zoniopoda iheringi
Zoniopoda juncorum
Zoniopoda omnicolor
Zoniopoda similis
Zoniopoda tarsata
Tristiridae
Atacamacris diminuta
Elysiacris angusticollis
Illapelia penai
Peplacris recutita
Tropidostethus bicarinatus
Parasitiformes
Polyaspidoidea
Iphidinychus geieri
Iphiduropoda penicillata
Antennophoridae
Antennophorus grandis
haplodiploid type unspecified
Argasidae
Argas brumpti
Argas cooleyi
Argas hermanni
Argas japonicus
Argas persicus
Argas radiatus
Argas reflexus
Argas sanchezi
Argas tridentatus
Argas vespertilionis
Argas zumpti
Ornithodoros alactogalis
Ornithodoros asperus
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
24
24
24
24
24
24
24
22
22
22
22
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
22
24
24
24
24
24
22
24
24
24
24
23
23
23
23
23
23
23
22
21
21
21
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
22
23
23
23
23
23
22
23
23
23
23
XO
XO
XO
XO
XO
XO
XO
XY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XO
XO
XO
XO
XO
XY
XO
XO
XO
XO
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
405
10
22
24
22
22
10
21
23
21
21
XY
XO
XO
XO
XO
405
405
405
405
405
parth
407
407
24
24
26
26
26
26
26
26
26
26
26
26
33
16
318
213
XY
XY
XY
XY
XY
XY
XY
XY
XY
XY
213
213
213
213
213
213
213
213
213
213
213
213
213
Taxa
Ornithodoros capensis
Ornithodoros gurneyi
Ornithodoros lahorensis
Ornithodoros macmillani
Ornithodoros moubata
Ornithodoros nereensis
Ornithodoros savignyi
Ornithodoros tartakovskyi
Ornithodoros tholozani
Otobius lagophilus
Otobius megnini
Ascidae
Asca aelhiopica
Asca afroaphidioides
Asca aphidioides
Asca cranela
Asca evansi
Asca garmani
Asca muma
Asca piloja
Asca quinqueselosa
Cheiroseius sp.
Gamasellodes bicolor
Gamasellodes rectiventris
haplodiploid type unspecified
Gamasellodes sp.
haplodiploid type unspecified
Gamasellodes vermivorax
haplodiploid type unspecified
Protogamasellus brevicornis
Protogamasellus hibernicus
Protogamasellus massula
Protogamasellus mica
Protogamasellus sp.
Rhinoseius colwelli
haplodiploid type unspecified
Blattisociidae
Blattisocius patagiorum
haplodiploid type unspecified
Lasioseius berlesi
Lasioseius denlalus
Lasioseius subterraneus
haplodiploid type unspecified
Lasioseius youcefi
Cercomegistidae
Cercoleipus coelonotus
Dermanyssidae
Dermanyssus gallinae
haplodiploid type unspecified
Dermanyssus prognephilus
haplodiploid type unspecified
Dermanyssus prognephilus.2
haplodiploid type unspecified
Dinychidae
Urodiaspis pannonica
Urodiaspis stammeri
Urodiaspis tecta
Discourellidae
Discourella baloghi
Discourella modesta
Eviphididae
Pelelhiphis berlesei
Pelelhiphis insignis
Pelelhiphis rufeseens
Halolaelapidae
Sexual
System
Ploidy
Female
2n
20
12
26
16
20
25
20
21
16
20
20
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
Male
2n
Sex Chrom. Source
Karyotype
XY
XY
XY
XY
XY
XY
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
408
213
213
213
parth
parth
parth
parth
parth
213
213
213
213
213
213
6
213
parth
parth
213
213
213
parth
213
26
213
6
213
10
211
6
213
parth
parth
407
407
407
parth
407
407
parth
parth
parth
213
213
213
319
Taxa
Sexual
System
Leitneria granulata
parth
Ixodidae
Amblyomma americanum
Amblyomma cajennense
Amblyomma darwini
Amblyomma dissimile
Amblyomma helvolum
Amblyomma inornotum
Amblyomma limbatum
Amblyomma maculalum
Amblyomma moreliae
Amblyomma sp.
Amblyomma testudinarium
Amblyomma triguttatum
Amblyomma tubereulalum
Aponomma concolor
Aponomma fimbriatum
Aponomma hydrosauri
Aponomma undatum
Boophilus annulatus
Boophilus micropius
Dermacentor albipictus
Dermacentor anderson
Dermacentor hunteri
Dermacentor nitens
Dermacentor occidentalis
Dermacentor parumapertus
Dermacentor silvarum
Dermacentor sp.
Dermacentor sp.2
Dermacentor variabilis
Haemaphysalis bancrofti
Haemaphysalis bispinosa
Haemaphysalis bremneri
Haemaphysalis campanulata
Haemaphysalis flava
Haemaphysalis formosensis
Haemaphysalis hystricis
Haemaphysalis japonica
Haemaphysalis kitaokai
Haemaphysalis lagrangei
Haemaphysalis leachii
Haemaphysalis leporispalustris
Haemaphysalis longicornis
Haemaphysalis megaspinosa
Haemaphysalis pentalagi
Hyalomma aegyptium
Hyalomma anatolicum
Hyalomma anatolicum excavatum
Hyalomma asiaticum
Hyalomma asiaticum excavatum
Hyalomma detritum
Hyalomma dromedarii
Hyalomma franchinii
Hyalomma impeltarum
Hyalomma marginatum
Hyalomma rhipicephaloides
Hyalomma rufipes
Ixodes cornuatus
Ixodes hexagonus
Ixodes holocyclus
Ixodes kingi
Ixodes laysanensis
Ixodes nipponensis
Ixodes ricinus
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
213
22
22
20
22
22
22
22
22
22
22
22
20
22
20
22
18
20
22
22
22
22
22
22
22
22
22
20
22
22
22
XO
XO
XY
XO
XO
XO
XXY
XO
XXY
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
22
22
22
22
20
22
20
22
16
22
27
22
22
22
22
22
XO
XO
XO
22
22
22
22
22
22
22
22
22
24
26
24
26
28
28
28
320
XO
XO
XO
XO
XO
XO
213
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XY
XY
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
213
Taxa
Ixodes tasmani
Rhipicephalus bursa
Rhipicephalus evertsi
Rhipicephalus sanguineus
Rhipicephalus secundus
Laelapidae
Androlaelaps casalis
haplodiploid type unspecified
Cosmolaelaps vacua
Dicrocheles phalaenodectes
PGE
Euandrolaelaps sp.
haplodiploid type unspecified
Eulaelaps shanghaiensis
haplodiploid type unspecified
Geolaelaps aculeifer
haplodiploid type unspecified
Geolaelaps oreithyiae
Geolaelaps sp.1
haplodiploid type unspecified
Geolaelaps sp.2
haplodiploid type unspecified
Haemogamasus centrocarpus
haplodiploid type unspecified
Haemogamasus longipes
haplodiploid type unspecified
Hypoaspis lubrica
haplodiploid type unspecified
Laelaspis sp.
haplodiploid type unspecified
Stratiolaelaps miles
haplodiploid type unspecified
Varroa destructor
PGE
Varroa jacobsoni
PGE
Laelapidae
Cosmolaelaps gurabensis
haplodiploid type unspecified
Cosmolaelaps sp.1
haplodiploid type unspecified
Cosmolaelaps sp.2
haplodiploid type unspecified
Cosmolaelaps vacua
Pseudoparasitus sp.
haplodiploid type unspecified
Macrochelidae
Areolaspis bifoliatus
haplodiploid type unspecified
Areolaspis sp.
haplodiploid type unspecified
Geholaspis alpinus
Geholaspis berlesei
Geholaspis longispinosus
Geholaspis longulus
Geholaspis mandibularis
Geholaspis pauperior
Glyptholaspis americana
haplodiploid type unspecified
Glyptholaspis confusa
haplodiploid type unspecified
Glyptholaspis fimicola
haplodiploid type unspecified
Glyptholaspis pontina
haplodiploid type unspecified
Sexual
System
Ploidy
Female
2n
24
24
22
22
22
Male
2n
Sex Chrom. Source
Karyotype
XO
XO
XO
213
213
213
213
213
14
213
6
213
213
parth
213
16
213
18
213
parth
213
213
213
213
213
14
213
213
14
213
409
14
409
12
213
213
213
parth
213
213
10
211
213
parth
parth
parth
parth
parth
parth
213
213
213
213
213
213
213
213
213
213
321
Taxa
Holostaspella sp.
haplodiploid type unspecified
Macrocheles lerreus
Macrocheles penicilliger
Macrocheles peniculalus
Macrocheles similis
Macrochelidae
Macrocheles boudreauxi
haplodiploid type unspecified
Macrocheles glaber
haplodiploid type unspecified
Macrocheles insignitus
haplodiploid type unspecified
Macrocheles mammifer
haplodiploid type unspecified
Macrocheles matrius
haplodiploid type unspecified
Macrocheles merdarius
haplodiploid type unspecified
Macrocheles muscadomesticae
haplodiploid type unspecified
Macrocheles parapisentii
haplodiploid type unspecified
Macrocheles peregrinus
haplodiploid type unspecified
Macrocheles perglaber
haplodiploid type unspecified
Macrocheles pisentii
haplodiploid type unspecified
Macrocheles robustulus
haplodiploid type unspecified
Macrocheles rodriguezi
haplodiploid type unspecified
Macrocheles schaeferi
haplodiploid type unspecified
Macrocheles scutatus
haplodiploid type unspecified
Macrocheles sp.
haplodiploid type unspecified
Macrocheles subbadius
haplodiploid type unspecified
Macrocheles vernalis
haplodiploid type unspecified
Macronyssidae
Ophionyssus bacoti
haplodiploid type unspecified
Ophionyssus natricis
haplodiploid type unspecified
Ophionyssus sylviarum
haplodiploid type unspecified
Ornithonyssus bacoti
haplodiploid type unspecified
Ornithonyssus natrids
haplodiploid type unspecified
Ornithonyssus silviarum
haplodiploid type unspecified
Melicharidae
Proctolaelaps krimsei
haplodiploid type unspecified
Proctolaelaps longipilis
haplodiploid type unspecified
Nenteriidae
Nenteria breviunguiculata
Nenteria stylifera
Ologamasidae
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
213
parth
parth
parth
parth
213
213
213
213
10
213
213
213
213
213
213
10
213
213
213
213
10
213
213
213
213
213
213
213
10
213
16
211
18
211
18
211
16
213
18
213
18
213
213
213
407
407
parth
322
Taxa
Gamasellus vibrissae
Oplitidae
Oplitis alophora
Oplitis franzi
Oplitis wasmanni
Uroplitella conspicua
Uroplitella paradoxa
Parasitidae
Amblygamasus septentrionalis
Pergamasus brevicornis
Parasitidae
Eugamasus kraepelini
Eugamasus magnus
Phorytocarpais hyalinus
Phytoseiidae
Amblyseius aberrans
haplodiploid type unspecified
Amblyseius agrestris
Amblyseius barkeri
haplodiploid type unspecified
Amblyseius bibens
PGE
Amblyseius brevipes
haplodiploid type unspecified
Amblyseius chiapensis
haplodiploid type unspecified
Amblyseius chilensis
haplodiploid type unspecified
Amblyseius cucumeris
haplodiploid type unspecified
Amblyseius deleoni
haplodiploid type unspecified
Amblyseius deleoni.2
Amblyseius herbarius
Amblyseius hibisci
haplodiploid type unspecified
Amblyseius judaicus
haplodiploid type unspecified
Amblyseius largoensis
haplodiploid type unspecified
Amblyseius masiaka
haplodiploid type unspecified
Amblyseius messor
haplodiploid type unspecified
Amblyseius parasundi
Amblyseius rotundus
haplodiploid type unspecified
Amblyseius rubini
haplodiploid type unspecified
Amblyseius salish
Amblyseius sp.
haplodiploid type unspecified
Amblyseius swirskii
haplodiploid type unspecified
Amblyseius vazimba
haplodiploid type unspecified
Clavidromus jackmickleyi
haplodiploid type unspecified
Euseius hibisci
PGE
Euseius quetzali
PGE
Iphiseius degenerans
haplodiploid type unspecified
Metaseiulus occidentalis
Sexual
System
Ploidy
Female
2n
parth
Male
2n
Sex Chrom. Source
Karyotype
213
407
407
407
407
407
parth
12
12
213
213
12
10
213
213
213
8
213
8
213
213
8
213
8
213
8
213
8
213
8
213
8
213
8
213
213
213
8
213
8
213
8
213
8
213
8
213
213
8
213
8
213
213
8
213
8
213
8
213
parth
parth
parth
parth
parth
parth
410
410
8
213
6
411
323
Taxa
PGE
Neoseiulus californicus
PGE
Neoseiulus setulus
Paragignathus tamaricis
haplodiploid type unspecified
Phytoseiulus persimilis
PGE
Phytoseius amba
haplodiploid type unspecified
Phytoseius finitimus
haplodiploid type unspecified
Phytoseius sp.1
haplodiploid type unspecified
Phytoseius sp.2
haplodiploid type unspecified
Seiulus isolrichus
haplodiploid type unspecified
Typhlodromus athiasae
haplodiploid type unspecified
Typhlodromus caudiglans
haplodiploid type unspecified
Typhlodromus chazeaui
haplodiploid type unspecified
Typhlodromus contiguus
haplodiploid type unspecified
Typhlodromus drori
haplodiploid type unspecified
Typhlodromus fallacis
haplodiploid type unspecified
Typhlodromus guatemalensis
Typhlodromus gutierrezi
haplodiploid type unspecified
Typhlodromus phialatus
haplodiploid type unspecified
Typhlodromus porathi
haplodiploid type unspecified
Typhlodromus pyri
PGE
Typhlodromus rhenanus
haplodiploid type unspecified
Typhlodromus sp.
haplodiploid type unspecified
Typhlodromus stemlichti
haplodiploid type unspecified
Typhlodromus transvaalensis
Podocinidae
Podocinum pacificum
Podocinum sagax
haplodiploid type unspecified
Polyaspidae
Polyaspis sansonei
Polyaspididae
Apionoseius infirmus
Polyaspidoidea
Uroseius hunzikeri
Rhodacaridae
Rhodacarellus silesiacus
Rhodacarus denticulatus
Trachytidae
Polyaspinus cylindricus
Polyaspinus patavinus
Trachytes aegrota
Trachytes irenae
Trachytes lamda
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
412
parth
8
213
213
8
213
8
213
8
213
8
213
8
213
8
213
8
213
8
213
8
213
8
211
8
211
8
211
8
413
211
8
211
8
211
parth
414
8
211
8
211
8
211
parth
parth
213
10
10
211
213
407
407
407
parth
parth
213
213
parth
407
407
407
407
407
parth
parth
324
Taxa
Trachytes minima
Trachytes montana
Trachytes pauperior
Trachytes pi
Trachyuropodidae
Trachyuropoda coccinea
Trachyuropoda pulchella
Trachyuropoda pyriformis
Trematuridae
Leiodinychus orbicularis
Oodinychus karawaiewi
Oodinychus obscurasimilis
Oodinychus ovalis
Oodinychus spatulifera
Trematurella elegans
Trichouropoda schweizeri
Trichouropoda sociata
Urodinychidae
Fuscouropoda appendiculata
Phaulodiaspis borealis
Phaulodiaspis rackei
Uroobovella advena
Uroobovella arcuatus
Uroobovella cordieri
Uroobovella fracta
Uroobovella inermis
Uroobovella obovata
Uroobovella perforatus
Uroobovella woelkei
Uropodidae
Cilliba cassidea
Cilliba erlangensis
Cilliba minor
Cilliba sopronensis
Cilliba sp. 1
Cilliba sp. 2
Neodiscopoma pulcherrima
Neodiscopoma splendida
Olodiscus minimus
Pseudouropoda calcarata
Pseudouropoda structura
Pseudouropoda tuberosa
Uropoda hamulifera
Uropoda italica
Uropoda kargi
Uropoda minima
Uropoda misella
Uropoda orbicularis
Uropoda undulata
Urotrachytes formicarius
Veigaiidae
Gamasolaelaps whartoni
Veigaia cerva
Veigaia exigua
Veigaia kochi
Veigaia nemorensis
Veigaia parlita
Veigaia planieola
Veigaia pusilla
Veigaia tranisalae
Veigaia uneata
Phasmatodea
Phasmatidae
Acanthoxyla fasciata
thelytoky
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
407
407
213
213
parth
parth
parth
407
407
407
parth
407
407
407
407
407
407
407
407
407
407
407
407
407
407
407
407
407
407
407
407
407
213
407
407
407
407
407
213
407
407
407
407
407
407
407
407
407
407
407
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
parth
213
213
213
213
213
213
213
213
213
213
parth
415
325
Taxa
Acanthoxyla geisovii
thelytoky
Acanthoxyla huttoni
thelytoky
Acanthoxyla inermis
Acanthoxyla intermedia
thelytoky
Acanthoxyla prasina
thelytoky
Acanthoxyla speciosa
thelytoky
Acanthoxyla suteri
thelytoky
Clitarchus hookeri
Phasmidae
Acrophylla titan
Aplopus mayeri
Baculum artemis
Bostra sp.
Carausius furcillatus
Carausius furcillatus
Carausius juvenilis
Carausius morosus
Carausius morosus
Carausius rotundato-lobatus
Carausius sp.
Carausius theiseni
Carnacia telesphorus
Clitumnus extradentatus
Clonistria exornata
Ctenomorpha chronus
Ctenomorpha chronus
Ctenomorphodes sp.2
Ctenomorphodes tessulatus
Didymuria violescens
also XY
Didymuria violescens
race_3
Didymuria violescens
race_4
Didymuria violescens
race_5
Didymuria violescens
race_6
Didymuria violescens
race_7
Didymuria violescens
race_8
Didymuria violescens
Didymuria violescens
Didymuria violescens
Didymuria violescens
Dubreuilia lineata
Dyme krugiana
Extatosoma tiaratum
Extatosoma tiaratum
Genus7 longiceps
Genus8 sp1
Lampionus sp.?
Lampionus sp.I
Lampionus sp.III
Lampionus sp.V
Menexenus semiarmatus
Oncotophasma sp.
Orxines macklotti
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
parth
415
parth
415
parth
36
416
415
parth
415
parth
415
parth
415
36
36
36
72
36
parth
416
37
33
XO
XO
297
297
297
297
297
297
297
417
297
297
297
297
297
297
297
297
297
297
297
297
38
37
XO
297
36
35
XO
297
32
31
XO
297
33
32
XY
297
31
30
XY
297
29
28
XY
297
29
27
40
40
26
50
36
38
42
46
24
30
36
36
46
42
38
28
26
39
39
25
49
35
37
XY
XY
XO
XO
XO
XO
XO
XO
45
23
29
35
35
45
41
37
XO
XO
XO
XO
XO
XO
XO
XO
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
42
64
22
42
44
38
34
30
32
40
38
33
326
35
35
72
35
66
87
41
65
21
41
41
43
37
33
30
32
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XY
Taxa
Sexual
System
Pachymorpha simplicipes
Pachymorpha sp.3
Pachymorpha squalida
Parasipyloidea annulatus
Parasipyloidea cercata
Parasipyloidea granulosa
Parasipyloidea sp2
Parasipyloidea sp3
Parasipyloidea sp4
Parasipyloidea sp5
Parasosibia parva
Phibalosoma phyllinum
Phobaeticus sinetyi
Podacanthus typhon
Podacanthus viridiroseus
Podacanthus wilkinsoni
Pseudobacteria sp.
Sipyloidea panaeticus
Sipyloidea sipylus
Sipyloidea sp6
Tropiderus childrenii
Vetilia enceladus
Phylliidae
Acanthoderus grandis
Acanthoderus inermis
Bacillus atticus
Bacillus grandii benazzii
Bacillus grandii grandii
Bacillus libanicus
Bacillus lynceorum
parth
Bacillus rossius
Bacillus rossius
Bacillus rossius-grandii benazzii
Bacillus rossius-grandii grandii
Bacillus whitei
parth
Clonopsis androgenes
Clonopsis felicitatis
Clonopsis gallica
parth
Clonopsis gallica
Clonopsis maroccana
Clonopsis soumiae
parth
Clonopsis sp. 1
Clonopsis sp. 2
Epibacillus lobipes
Isagoras schraderi
Isagoras sp.
Isagoras subaquilus
Leptynia attenuata
Leptynia attenuata
Leptynia caprai
Leptynia montana
Leptynia sp.
Phalces longiscaphus
Phyllium bioculatum
Pijnackeria barbarae
Pijnackeria hispanica
Moved from Genus Leptynia Scali 2009
Pijnackeria hispanica
Moved from Genus Leptynia Scali 2009
Pijnackeria hispanica
Pijnackeria hispanica
Moved from Genus Leptynia Scali 2009
Pijnackeria lelongi
Pijnackeria lucianae
Pijnackeria masettii
Ploidy
Female
2n
34
30
34
30
40
36
40
28
39
39
54
42
52
28
36
36
48
22
80
38
34
38
44
44
34
34
34
3
2
3
51
36
35
35
35
44
36
54
56
22
72
36
72
3
34
48
28
36
36
40
38
40
36
34
38
57
4
76
3
Male
2n
Sex Chrom. Source
Karyotype
29
33
29
39
35
39
27
38
38
53
41
51
28
35
35
46
21
80
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XY
XO
XO
XO
XO
33
38
XO
XY
43
43
XO
XO
33
33
XO
XO
XY
35
36
XO
35
XO
56
21
XO
35
XO
36
34
47
27
36
35
39
37
39
35
33
37
XY
XO
XO
XY
XO
XO
XO
XO
XO
XO
XO
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
297
418
418
418
297
418
297
297
418
418
418
419
419
419
297
419
419
420
420
297
297
297
297
421
297
421
421
421
297
297
422
423
423
38
54
37
54
XO
423
297
38
38
57
37
37
XO
XO
XO
422
422
422
327
Taxa
Pijnackeria originis
Prisopus ariadne
Prisopus berosus
Pseudophasma menius
Timematidae
Timema poppensis
Timema shepardi
Phthiraptera
Goniodidae
Chelopistes meleagridis
achiasmatic male meiosis
Unknown sp.
achiasmatic male meiosis
Gyropidae
Gyropus ovalis
achiasmatic male meiosis
Haematopinidae
Haematopinus asini
achiasmatic male meiosis
Haematopinus consobrinus
achiasmatic male meiosis
Haematopinus suis
achiasmatic male meiosis
Heptapsogasteridae
Unknown sp.
achiasmatic male meiosis
Hoplopleuridae
Hoplopleura sp.
achiasmatic male meiosis
Linognathidae
Linognathus tenuirostris
achiasmatic male meiosis
Menoponidae
Menacanthus stramineus
achiasmatic male meiosis
Pediculidae
Pediculus capitis
achiasmatic male meiosis
Pediculus corporis
achiasmatic male meiosis PGE
Pediculus vestimenti
achiasmatic male meiosis
Philopteridae
Goniodes stylifer
achiasmatic male meiosis
Lipeurus baculus
Polyplacidae
Polyplax serrata
achiasmatic male meiosis
Trichodectidae
Bovicola caprae
achiasmatic male meiosis
Bovicola limbata
achiasmatic male meiosis
Plecoptera
Perlidae
Calineuria jezoensis
Perla abdominalis
Perla baetica
Perla bipunctata
Perla cephalotes
Perla immarginata
Perla marginata
Perla maxima
Perlodidae
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
38
28
50
24
37
28
49
23
XO
XY
XO
XO
422
297
297
297
424
424
parth
10
12
425
11
426
4
427
18
427
14
427
10
427
11
426
16
427
12
427
10
10
427
12
12
427
12
12
427
10
10
427
24
427
12
427
16
427
14
428
14
14
XY
428
26
25
26
26
21
26
10
22
19
XO
XXO
XXO
XO
XXO
XY
XXO
XO
429
430
430
430
430
430
431
430
22
10
24
20
328
Taxa
Isogenus alpinus
Isogenus fontium
Isogenus imhoffi
Isoperla grammatica
Isoperla rivulorum
Perlodes intricala
Perlodes jurassica
Perlodes microcephala
Protura
Acerentomidae
Acerentomon maius
Acerentomon meridionale
Acerentomon microrhinus
Eosentomidae
Eosentomon sakura
Eosentomon transitorium
homomorphic
Psocoptera
Amphientomidae
Seopsis sp.
holocentric
Seopsocus acuminatus
holocentric
Amphipsocidae
Amphipsocus japonicus
holocentric
Dasypsocus japonicus
holocentric
Kolbia quisquiliarum
holocentric
Archipsocidae
Archipsocus sp.
holocentric
Caeciliusidae
Asiocaecilius singaporensis
holocentric
Caecilius fuscopterus
holocentric
Caecilius sp.1
holocentric
Caecilius sp.2
holocentric
Dypsocus sp.
holocentric
Enderleinella obsoleta
holocentric
Valenzuela burmeisteri
holocentric
Valenzuela flavidus
apomictic
Valenzuela gynapterus
holocentric
Valenzuela labinae
holocentric
Valenzuela oyamai
holocentric
Valenzuela piceus
holocentric
Valenzuela sp.
apomictic
Ectopsocidae
Ectopsocopsis cryptomeriae
holocentric
Ectopsocus briggsi
holocentric
Sexual
System
Ploidy
Female
2n
Male
2n
24
26
26
26
26
33
31
27
parth
3
parth
XXO
XXO
XXO
complex XO
12
XY
432
432
432
10
18
10
18
XY
XY
433
433
16
15
XO
434
18
17
XO
434
16
16
XY
434
16
15
XO
435
16
16
XY
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
27
434
17
XO
27
3
430
430
430
430
430
430
430
430
12
9
16
18
parth
Sex Chrom. Source
Karyotype
434
434
18
17
XO
434
18
17
XO
434
27
436
18
17
XO
434
18
17
XO
437
329
Taxa
Ectopsocus briggzi
holocentric
Ectopsocus maindroni
holocentric
Ectopsocus meridionalis
apomictic
Elipsocidae
Elipsocus abdominalis
holocentric
Elipsocus moebiusi
holocentric
Elipsocus pumilis
holocentric
Elipsocus pumilis
holocentric
Elipsocus westwoodi
holocentric
Hemineura dispar
holocentric
Reuterella helvimacula
holocentric
Epipsocidae
Epipsocus lucifugus
holocentric
Hemipsocidae
Hemipsocus sp.
holocentric
Lachesillidae
Lachesilla pedicularia
holocentric
Lachesilla pedicularia
holocentric
Lachesilla quercus
holocentric
Lachesilla tanaidana
holocentric
Lachesilla tanaidana
holocentric
Liposcelidae
Liposcelis bostrychophilus
holocentric
Liposcelis divergens
holocentric
Mesopsocidae
Mesopsocus honkongensis
holocentric
Mesopsocus laticeps
holocentric
Mesopsocus unipunctatus
holocentric
Peripsocidae
Diplopsocus fasciatus
holocentric
Peripsocus alboguttatus
holocentric
Peripsocus golubae
holocentric
Peripsocus parvulus
holocentric
Peripsocus phaeopterus
holocentric
Peripsocus quercicola
holocentric
Peripsocus sp.
holocentric
Sexual
System
parth
Ploidy
3
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
17
XO
434
18
17
XO
434
27
434
18
17
XO
434
14
13
XO
434
18
17
XO
437
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
437
18
17
XO
434
18
17
XO
434
18
17
XO
437
18
17
XO
434
parth
16
434
parth
18
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
330
Taxa
Peripsocus subfasciatus
apomictic
Philotarsidae
Aaroniella badonneli
apomictic
Haplophallus orientalis
holocentric
Philotarsus picicornis
holocentric
Pseudocaecilliidae
Pseudocaecilius hirsutus
holocentric
Pseudocaecilius maculosus
holocentric
Psocidae
Amphigerontia bifasciata
holocentric
Amphigerontia bifasciata
holocentric
Amphigerontia contaminata
holocentric
Amphigerontia jezoensis
holocentric
Amphigerontia jezoensis
holocentric
Atlantopsocus adustus
holocentric
Atrichadenotecnum nudum
holocentric
Blaste conspurcata
holocentric
Brachinodiscus cf. lepidus
holocentric
Cerastipsocus fuscipennis
holocentric
Cerastipsocus venosus
holocentric
Hyalopsocus contrarius
holocentric
Loensia moesta
holocentric
Loensia picicornis
holocentric
Loensia variegata
holocentric
Loensia variegata
holocentric
Metylophorus nebulosus
holocentric
Neopsocopsis hitricornis
holocentric
Psococerastis gibbosa
holocentric
Psococerastis interrupta
holocentric
Psococerastis reticulata
holocentric
Psococerastis sinensis
holocentric
Psocus leidyi
holocentric
Ptycta incurvata
holocentric
Sigmatoneura kolbei
holocentric
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
parth
3
27
434
parth
2
18
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
437
18
17
XO
434
18
17
XO
434
16
15
XO
437
16
15
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
14
13
XO
437
14
13
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
437
331
Taxa
Sexual
System
Sigmatoneura kolbei
holocentric
Trichadenotecnum majus
parth
apomictic
Trichadenotecnum medium
holocentric
Trichadenotecnum sexpunctatum
holocentric
Trichadenotecnum sexpunctatum
holocentric
Trichadenotecnum sinuatum
holocentric
Psoquillidae
Psoquilla marginepunctata
holocentric
Psyllipsocidae
Dorypteryx domestica
holocentric
Dorypteryx domestica
holocentric
Psocatropos sp.
holocentric
Ptiloneuridae
Triplocania caudata
holocentric
Stenopsocidae
Graphopsocus cruciatus
holocentric
Stenopsocus aphidiformis
holocentric
Stenopsocus aphidiformis
holocentric
Stenopsocus immaculatus
holocentric
Stenopsocus lachlani
holocentric
Trichopsocidae
Trichopsocus brincki
holocentric
Trichopsocus clarus
holocentric
Trichopsocus dalii
holocentric
Trogiidae
Lepinotus inquilinus
holocentric
Lepinotus reticulatus
holocentric
Trogium pulsatorium
holocentric
Raphidioptera
Raphidiidae
Agulla astuta
Agulla bicolor
Agulla bractea
Rhaphidia major
Rhaphidia nigricollis
Rhaphidia xanthostigma
Siphonaptera
Leptopsyllidae
Leptopsylla musculi
Pulicidae
Ctenocephalus canis
Ctenocephalus orientis
Xenopsylla astia
Ploidy
2
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
17
XO
18
434
434
18
17
XO
434
18
17
XO
437
18
17
XO
434
18
17
XO
434
20
19
XO
434
30
29
XO
434
30
29
XO
434
30
29
XO
434
18
17
XO
434
18
17
XO
434
24
23
XO
437
24
23
XO
434
18
17
XO
434
24
23
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
17
XO
434
18
434
22
21
XO
434
26
26
26
26
26
26
26
26
26
26
Xy+
Xy+
Xy+
XY
Xy+
Xy+
438
438
438
439
438
440
22
22
XY
405
14
16
20
14
16
18
XY
XYY
XXXYY
405
441
442
332
Taxa
Xenopsylla cheopis
Xenopsylla prasadi
Strepsiptera
Myrmecolacidae
Stichotrema dallatorreanum
Stylopidae
Xenos peckii
orig. Acroschismus wheeleri
Xenos sp
heteromorphic
Thysanoptera
Phlaeothripidae
Gynaikothrips uzeli
arrhenotoky
Gynaikothrips uzeli
arrhenotoky
Gynaikothrips ficorum
arrhenotoky
Haplothrips statices
arrhenotoky
Haplothrips tritici
arrhenotoky
Liothrips sp
arrhenotoky
Neoheegeri verbasci
arrhenotoky
Thripidae
Aptinothrips rutua
arrhenotoky
Aptinothrips rutua
arrhenotoky
Frankliniella insularis
arrhenotoky
Frankliniella schultzei
arrhenotoky
Heliothrips haemorhroidalis
arrhenotoky
Heliothrips haemorrhoidalis
arrhenotoky
Heliothrips haemorrhoidalis
arrhenotoky
Heliothrips haemorrhoidalis
arrhenotoky
Limothrips dentricornis
arrhenotoky
Parthenothrips dracaenae
arrhenotoky
Retithrips syriacus
arrhenotoky
Selenothrips rubrocinctus
arrhenotoky
Sericothrips staphilinua
arrhenotoky
Taeniothrips iconsequene
arrhenotoky
Taeniothrips iconsequene
arrhenotoky
Taeniothrips iconsequene
arrhenotoky
Taeniothrips simplex
arrhenotoky
Trichoptera
Goeridae
Goera pilosa
Hydropsychidae
Sexual
System
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
18
10
17
10
XXY
XXYY
parth
442
442
443
16
444
8
8
26
13
446
30
15
446
30
15
446
30
15
446
20
10
446
24
12
446
24
12
446
100
50
446
106
53
446
28
14
446
34
17
446
42
21
446
32
16
446
52
26
446
56
28
446
38
19
446
30
15
446
38
19
446
36
18
446
28
14
446
32
16
446
36
18
446
40
20
446
20
10
446
44
447
333
XY
445
Taxa
Sexual
System
Amphipsyche senegalensis
parth
Hydropsyche pellucidula
Hydropsyche sp.
Hydroptilidae
Oxyethira fagesii
Leptoceridae
Athripsodes aterrimus
Athripsodes excisus
Limnephilidae
Allogamus auricollis
Anabolia furcata
Anabolia soror
Apatania zonella
parth
Chaetopteryx villosa
Glyphotaelius pellucidus
Halesus tesselatus
Hesperophylax designatus
Hydatophylax infumatus
Limnephilus affinis
Limnephilus auricula
Limnephilus centralis
Limnephilus flavicornis
Limnephilus lunatus
Limnephilus nigriceps
Limnephilus politus
Limnephilus rhombicus
Limnephilus stigma
Potamophylax stellatus
Molannidae
Molanna angustata
Phryganeidae
Agrypnetes crassicornis
Dasystegia obsoleta
Dasystegia varia
Phryganea bipunctata
Phryganea grandis
Trichostegia minor
Polycentropidae
Plectronemia conspersa
Polycentropus flavomaculatus
Psychomyiidae
Psychomyia flavida
parth
Rhyacophilidae
Rhyacophila cf. obtusidens
Rhyacophila nubila
Rhyacophila sp.
Stenopsychidae
Stenopsyche griseipennis
Stenopsyche marmorata
Zoraptera
Zorotypidae
Zorotypus hubbardi
heteromorphic
Zygentoma
Lepismatidae
Lepisma (Thermobia) domestica
Lepisma domestica
Lepisma saccharina
Lepisma saccharina_2
Ploidy
Female
2n
Male
2n
Sex Chrom. Source
Karyotype
29
29
30
30
ZO
ZO
448
449
449
27
28
ZO
447
50
50
447
447
59
59
59
60
60
60
ZO
ZO
ZO
59
59
60
60
42
60
60
12
58
26
60
26
32
60
60
60
60
ZO
ZO
57
25
25
25
ZO
ZO
450
449
449
448
447
451
447
447
447
447
447
447
447
447
447
447
447
447
447
54
447
100
56
56
56
56
38
447
447
447
447
447
447
26
26
ZO
ZO
449
449
448
45
46
46
46
ZO
447
447
449
25
26
ZO
ZO
447
452
38
38
XY
453
36
36
34
34
34
58
XO
XO
248
248
454
248
60
334
XO
Taxa
Sexual
System
Ploidy
Female
2n
335
Male
2n
Sex Chrom. Source
Karyotype
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
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351
Appendix B
Genbank accesion numbers
352
Sequences retrieved from GenBank. Chimeric OTUs that represent a taxonomic unit
higher than species are indicated with a †. Chimeric OTUs that function as “anchors” are
indicated with an *.
Species
Adephaga
Carabidae
Abacetus†
Abax ovalis
Abax parallelepipedus
Acinopus picipes
Agonum marginatum
Agonum muelleri
Amara aenea
Amara apricaria
Amara aulica
Amblycheila baroni
Amblystomus†
Amblytelus curtus
Amphasia sericea
Anchomenidius astur
Anisodactylus hispanus
Anisodactylus virens
Anthia†
Apenes†
Aptinus displosor
Arthropterus†
Asaphidion curtum
Bembidion tibiale
Bembidion balli
Bembidion inaequale inaequale
Bembidion lapponicum
Bembidion levettei carrianum
Bembidion levettei levettei
Bembidion assimile
Bembidion concolor
Bembidion californicum
Bembidion levigatum
Bembidion lampros
Bembidion properans
Bembidion insulatum
Bembidion nigripes
Bembidion rapidum
Bembidion umbratum
Bembidion bifossulatum
Bembidion aenulum
Bembidion confusum
Bembidion coxendix
Bembidion americanum
Bembidion mexicanum
Bembidion sejunctum semiaureum
Bembidion tetracolum
Bembidion transversale
Bembidion decorum
Bembidion obtusum
Bembidion planatum
Bembidion punctulatum
Bembidion antiquum
Bembidion arenobilis
Bembidion bellorum
Bembidion chalceum
Bembidion honestum
Bembidion integrum
Bembidion louisella
Bembidion rothfelsi
Bembidion rufotinctum
Bembidion ruficorne
Bembidion articulatum
Bembidion tairuense
Bembidion quadrimaculatum dubitans
Brachinus †
Bradybaenus†
Calathus brevis
Calathus fuscipes
Calathus abaxoides
Calathus ambiguus chevrolati
Calathus ascendens
Calathus asturiensis
Calathus auctus
Calathus circumseptus
Calathus erratus
Calathus freyi
Calathus granatensis
Calathus melanocephalus
28s
16s
18s
COI
ArgK
AF398681.1
GU347362.1
Elf1a
AF398635.1
AF533283.1
AF533280.1
GU347370.1
FJ173081.1
FJ173093.1
GU347386.1
EU797338.1
Wng
L42908.1
GU347714.1
FJ173119.1
FJ173123.1
AF002774.1
GU348075.1
AF423057.1
GU347012.1
GU347018.1
AJ583309.1
GU347026.1
GU347034.1
AF398565.1
EU710807.1
EU797284.1
AJ583325.1
AF398683.1
AF012484.1
AF398566.1
EU710812
FJ173090.1
AJ583290.1
AJ583292.1
AF398696.1
AF398713.1
AF398644.1
GU556078.1
GU347476.1
EF648838.1
EU677692.1
EF648841.1
AF398647.1
AF012480.1
AF012516.1
AF002792.1
GU347820.1
EF648613.1
AF437906.1
AF398567.1
AF398569.1
AF398570.1
GU556027.1
GU347132.1
EF648695.1
EU677521.1
EF648619.1
AF002791.1
EF648620.1
EF648702.1
EF649474.1
EU677672.1
EF649479.1
AF398571.1
EF649480.1
DQ155770.1
EF648833.1
EF648832.1
GU556083.1
GU347437.1
GU347455.1
GU556081.1
EU677690.1
EU677691.1
EF648835.1
GQ424256.1
EF648840.1
EF648837.1
EF648834.1
GU454739.1
GU454738.1
EF648611.1
EF648610.1
GU347781.1
GU347799.1
EF649472.1
EF649471.1
GU556032.1
GU347093.1
GU347116.1
GU556030.1
DQ059789.1
DQ059790.1
EF648614.1
EU677518.1
EU677520.1
EF648696.1
GQ424227.1
EF648617.1
EF648618.1
EF648612.1
AF012490.1
GU347814.1
EU677688.1
GU347423.1
GU556085.1
GU556086.1
GU347480.1
EF648843.1
EF648963.1
EF648867.1
EF648875.1
EF648994.1
EF649055.1
EF648902.1
EF648927.1
EF649099.1
GU347463.1
GU347409.1
GU556089.1
GU556087.1
AF398693.1
EF648693.1
EF648692.1
EF648699.1
EF648700.1
EF648694.1
GU454769.1
GU454768.1
DQ155825.1
EU677517.1
GU347767.1
GU348171.1
EF648621.1
EF648630.1
EF648639.1
EF648644.1
EF648649.1
EF648659.1
EF648666.1
EF648678.1
EF648687.1
GU347806.1
GU347753.1
GU556134.1
GU556133.1
AF012478.1
GU347079.1
DQ155728.1
EU677668.1
EU677671.1
EF649475.1
GQ424283.1
EF649478.1
EF649481.1
EF649473.1
GU556033.1
EU677667.1
GU556034.1
GU556035.1
GU347136.1
EF648703.1
EF648716.1
EF648728.1
EF648733.1
EF648739.1
EF648751.1
EF648757.1
EF648763.1
EF648772.1
EF649482.1
EF649503.1
EF649534.1
EF649550.1
EF649553.1
EF649608.1
EF649644.1
EF649656.1
EF649670.1
GU347118.1
GU347068.1
DQ155801.1
GU556038.1
GU556036.1
AF398572.1
AJ583311
GU254349.1
FJ173072.1
GU254368.1
GU254401.1
GU254378.1
GU254390.1
GU254381.1
FJ173086.1
GU254355.1
GU254379.1
GU254340.1
GU254342.1
FJ173118.1
353
GU323022.1
GU254437.1
FJ173134.1
GU254456.1
GU254482.1
GU254464.1
GU254474.1
GU254466.1
FJ173180.1
GU254443.1
GU254465.1
GU254473.1
GU254432.1
Species
Calathus micropterus
Calathus mollis
Calathus rectus
Calathus rotundatus
Calosoma inquisitor
Carabus abbreviatus
Carabus cancellatus
Carabus granulatus
Carabus monticola
Carabus nemoralis
Carabus auronitens auronitens
Carabus blaptoides blaptoides
Carabus fruhstorferi
Carabus lineatus lateralis
Carabus punctatoauratus
Carabus solieri
Carabus splendens
Carabus nitens
Carabus problematicus
Carabus hortensis hortensis
Carabus caelatus
Carabus violaceus germari
Carabus creutzeri kircheri
Carabus hispanus
Carabus melancholicus costatus
Carabus rutilans
Carabus blaptoides
Carabus fruhstorferi
Carenum interruptum
Carterus fulvipes
Carterus rotundicollis
Chlaenius †
Cicindela sedecimpunctata
Cicindela marginata
Cicindela marutha
Cicindela dorsalis
Cicindela circumdata
Cicindela aurulenta
Cicindela hybrida
Cicindela lunulata
Cicindela punctulata
Cicindela repanda
Cicindela sexguttata
Cicindela splendida
Cicindela tranquebarica
Cicindela undulata
Clivina†
Colliuris pennsylvaniea
Craspedophorus†
Ctenostoma †
Cychrus caraboides
Cylindera cognata
Cymindis bedeli
Demetrias atricapillus
Dicrochile brevicollis
Diplocheila†
Dixus capito obscuroides
Dromius†
Drypta†
Dyschirius†
Egadroma marginatum
Euthenarus promptus
Galerita bicolor
Graphipterus†
Harpalus aesculans
Harpalus affinis
Harpalus anxius subcylindricus
Harpalus attenuatus
Harpalus contemptus
Harpalus decipiens
Harpalus dimidiatus
Harpalus distinguendus
Harpalus ebeninus
Harpalus honestus
Harpalus pennsylvanicus
Harpalus rubripes
Harpalus serripes
Harpalus wagnerii
Lebia†
Lecanomerus†
Licinus†
Lophyra†
Loricera pilicornis
Loxandrus†
Macrocheilus†
Mantichora amigdaloides
28s
GU254350.1
FJ173071.1
GU254369.1
GU254387.1
AB031419.1
16s
18s
COI
ArgK
Wng
FJ173117.1
Elf1a
GU254438.1
FJ173133.1
GU254457.1
GU254472.1
AY183580.1
AF231686.1
AF219428.1
GU347495.1
GU347487.1
AF012507.1
GU348178.1
L42910.1
GU347140.1
U75561.1
AF190040.1
GU347501.1
AY183653.1
AF219564.1
AY183630.1
EU797291.1
AY183598.1
AY183625.1
AY183609.1
AY183601.1
AY183602.1
AY183605.1
AY183604.1
GU348190.1
DQ155753.1
AY183633.1
AY183651.1
AY183595.1
AY183592.1
AY183584.1
AY183600.1
AY183654.1
AY183603.1
AY183607.1
AY183609.1
AF012491.1
AJ583283.1
AJ583284.1
AF438039.1
AF201404.1
AF012518.1
AF132989.1
AF133000.1
AF438894.1
AF437921.1
AF398579.1
AJ514980.1
U75564.1
AJ514976.1
AY523657.1
AJ879182.1
AJ515105.1
AF438891.1
AF438884.1
AF438886.1
AF438861.1
AF438889.1
AJ515118.1
GU347505.1
AF398712.1
AF438045.1
EU797350.1
GU347513.1
GU983749.1
AF398651.1
DQ337114.1
AJ583538.1
AJ514961.1
AY165729.1
U75563.1
GU347849.1
EU797405.1
GU347857.1
AY165643.1
EU723580.1
DQ923390.1
AJ514992.1
GU347163.1
AF398581.1
AF437928.1
EU797297.1
AB109837.1
GU983830.1
AF002773.1
AF398583.1
DQ155934.1
AF438058.1
AF438061.1
AF437936.1
AF437939.1
AF438063.1
AF438064.1
GU347528.1
GU347871.1
AF398686.1
AF398711.1
AF002780.1
AF201401.1
FJ173087.1
FJ173121.1
GU347567.1
GU347911.1
AF438084.1
AF438088.1
GU347576.1
GU983704.1
GU347593.1
AF398661.1
AF438092.1
GU348264.1
EF517579.1
GU348281.1
AF002778.1
AF423056.1
354
AJ583278.1
DQ155916.1
AF437942.1
GU347184.1
AJ583270.1
AJ583258.1
AF398590.1
AF398593.1
AJ583350.1
AJ583348.1
AJ583353.1
AJ583357.1
AJ583344.1
AJ583335.1
AJ583342.1
AJ583347.1
AJ583356.1
AJ583341.1
DQ059798.1
AJ583354.1
AJ583352.1
AJ583339.1
DQ059805.1
AJ583255.1
GU347229.1
AJ514975.1
FN868618.1
FJ173145.1
AF437959.1
AF437963.1
EF588668.1
AF398600.1
AF437965.1
Species
Mecyclothorax†
Megacephala sobrina
Megacephala brasiliensis
Megacephala whelani
Microlestes†
Mystropomus subcostatus
Nebria brevicollis
Nesarpalus sanctaecrucis
Notagonum†
Notiobia†
Notiophilus†
Olisthopus†
Omus californicus
Oodes†
Ophonus azureus
Ophonus brevicollis
Ophonus longicollis
Ophonus stictus
Orthomus velocissimus pardoi
Oxycheila†
Pamborus alternans
Pamborus opacus
Pamborus tropicus
Paraphonus †
Penetretus†
Pentacomia†
Percus politus
Perileptus areolatus
Pheropsophus†
Platyderus lusitanicus
Platyderus varians
Platymetopus†
Platynus †
Poecilus chalcites
Poecilus cupreus
Poecilus lucublandus
Poecilus versicolor
Pogonus chalceus
Promecoderus†
Prothyma†
Pterostichus diligens
Pterostichus angustatus
Pterostichus nemoralis
Pterostichus aterrimus nigerrimus
Pterostichus interruptus
Pterostichus melanarius
Pterostichus ziegleri
Pterostichus niger
Pterostichus niger
Pterostichus anthracinus
Pterostichus minor
Pterostichus nigrita
Pterostichus flavofemoratus
Pterostichus morio
Pterostichus spinolae
Pterostichus alacer
Pterostichus alacer
Pterostichus globosus ebenus
Pterostichus globosus ebenus
Pterostichus madidus
Pterostichus madidus
Pterostichus herculaneus
Pterostichus oblongopunctatus
Sarothrocrepis corticalis
Scarites laevigatus
Scarites terricola
Scarites hespericus
Scarites occidentalis
Scarites buparius
Scarites eurytus
Scarites subterraneus
Siagona†
Sphallomorpha†
Stenolophus lecontei
Stenolophus mixtus
Stenolophus teutonus
Stomis†
Syntomus†
Tachys †
Therates†
Trechus obtusus
Trechus quadristriatus
Zabrus†
Dytiscidae
Agabus confinis
Agabus conspersus
28s
AF398648.1
DQ152064.1
DQ152044.1
DQ152041.1
16s
18s
AF012482
DQ152123.1
DQ152106.1
DQ152100.1
EF517580.
AF201395.1
AF438095.1
EF694843.1
AF337390.1
AF438101.1
EU797369.1
FJ173098.1
COI
AF438907.1
Wng
AF398601.1
Elf1a
DQ155976.1
EF694889.1
FN868614.1
AJ583310.1
DQ337161.1
AF002804.1
FJ173127.1
AF012519.1
ArgK
AJ583303.1
DQ155829.1
AF438104.1
AF337564.1
AF437972.1
EU797312.1
AF437973.1
FJ173155.1
AF437975.1
AJ583367.1
AJ583372.1
AJ583365.1
AJ583361.1
FJ173088.1
EU797376.1
AB109869.1
FJ173146.1
AF201393.1
EU797318.1
AY722961.1
AY722981.1
AY722971.1
AY847575.1
AJ583322.1
GU556108.1
L42928.1
AY334312.1
GQ293625.1
AF398678.1
FR729593.1
FJ173091.1
GU556142.1
AF423039.1
FJ173128.1
GQ293503.1
AF012477.1
GU556052.1
FJ173160.1
GQ293688.1
FJ173122.1
AY551835.1
FJ173211.1
AJ583319.1
GU348308.1
EU142291.1
DQ059807.1
AY574578.2
DQ059816.1
AY734990.1
GU556054.1
AF398619.1
FJ173149.1
AF438123.1
GU347618.1
EU142440.1
AB243479.1
GU556114.1
GU556144.1
AF012499.1
EU142321.1
AB243533.1
GU556057.1
AJ514997.1
AB243463.1
EU142393.1
FJ173089.1
GU347624.1
AB243487.1
AF398707.1
GU347687.1
AB243470.1
AB243470.1
GU347621.1
EU142534.1
FJ173147.1
AF190039.1
AB253869.1
GU347663.1
AB243493.1
GU347968.1
GU347280.1
AF002779.1
GU348031.1
GU347991.1
GU347993.1
GU347965.1
DQ063219.1
GU347343.1
GU347304.1
GU347311.1
GU347278.1
DQ155942.1
DQ155787.1
GU348003.1
AB243541.1
AF398623.1
AB243524.1
AB243524.1
AB243547.1
AB243548.1
AB243549.1
AB243542.1
AB243542.1
AB243495.1
AB243488.1
FJ173114.1
FJ173114.1
FJ173132.1
FJ173132.1
EU142343.1
GU347667.1
AF398670.1
GU348011.1
FJ173234.1
FN868607.1
FN868607.1
EU142482.1
GU347323.1
FJ173170.1
FJ173170.1
AB243534.1
AF398624.1
AF042676.1
AF042678.1
AF042679.1
AF042677.1
AF042674.1
AF042675.1
AF398708.1
AF002795.1
AF012493.1
AF398717.1
AF398679.1
GU347691.1
FJ173094.1
EU142459.1
AF438140.1
GU556120.1
GU348379.1
FJ173124.1
AF398625.1
AF398636.1
DQ063220.1
DQ155940.1
DQ155970.1
DQ155941.1
DQ155915.1
AF423047.1
GQ293608.1
GQ293619.1
FJ173096.1
GQ293726.1
GQ293743.1
GQ293534.1
FJ173125.1
AY138608.1
AY138610.1
355
DQ155831.1
FN868615.1
AY551887.1
FJ173151.1
AB243535.1
AF437962.1
GU556063.1
Species
Agabus biguttatus
Agabus bipustulatus
Agabus ramblae
Agabus sturmi
Colymbetes paykulli
Colymbetes striatus
Cybister japonicus
Cybister lateromarginalis
Cybister sugillatus
Cybister tripunctatus
Dytiscus circumcinctus
Dytiscus dauricus
Dytiscus marginalis*
Dytiscus marginalis czerskii
Eretes strictus
Hydaticus†
Hydroporus humilis
Hydrovatus†
Hygrobia hermanii
Ilybius aenescens
Ilybius albarracinensis
Ilybius ater
Ilybius chalconatus
Ilybius erichsoni
Ilybius fuliginosus
Ilybius guttiger
Ilybius montanus
Ilybius quadriguttatus
Ilybius subaeneus
Ilybius vittiger
Ilybius wasastjernae
Nebrioporus lanceolatus*
Platambus maculatus*
Rhantus exsoletus
Stictotarsus griseostriatus
Stictotarsus ibericus
Gyrinidae
Dineutes†
Gyrinus†
Haliplidae
Haliplus flavicollis
Haliplus immaculatus
Haliplus lineatocollis
Haliplus mucronatus
Haliplus ruficollis
Peltodytes†
Noteridae
Canthydrus†
Noterus clavicornis
Trachypachidae
Trachypachus holmbergi
Trachypachus slevini
Archostemata
Micromalthidae
Micromalthidae†
Myxophaga
Torridincolidae
Torridincolidae†
Polyphaga
Anobiidae
Gibbium psylloides
Lasioderma sp
Ptinus†
Stegobium paniceum
Anthicidae
Anthicidae†
Anthribidae
Araecerus fasciculatus
Platystomus†
Ptychoderes†
Attelabidae
Apoderus coryli
Deporaus †
Larinus†
Liparus†
Bostrichidae
Bostrychopsis bengalensis*
Sinoxylon†
Brentidae
Ceratapion penetrans
Eutrichapion†
Rhopalapion longirostre
Brentus anchorago
Nanophyes marmoratus
Nemocephalus†
Buprestidae
28s
16s
AY039258.1
AF428198.1
AY138626.1
AY138633.1
AY334104.1
AJ850360.1
AJ850361.1
AJ850362.1
AY368226.1
AY138644.1
18s
AJ318687.1
FN257262.1
FN257254.1
AF201408.1
AJ850502.1
AJ318702.
FN257263.1
AY138646.1
AJ318704.1
AF518261.1
EF056679.1
AY071780.1
AF428202.1
AF201409.1
AJ318716.1
AJ318673.1
FJ818198.1
EU797361.1
EU797362.1
AF428200.1
AY138661.1
EF417072.1
AY250927.1
AY138675.1
AJ850354.1
EF670030.1
EU677683.1
EU797355.1
AJ850481.1
AJ850434.1
FN257256.1
AJ850497.1
COI
AY039267.1
AF309332.1
AY138716.1
AY138723.1
AY334220.1
ArgK
Wng
Elf1a
AF392005.1
FN256314.1
DQ813718.1
DQ813680.1
DQ813681.1
DQ813686.1
DQ813687.1
AY368230.1
AY138732.1
FR751067.1
FR751063.1
AY138734.1
DQ431209.1
AF518291.1
EF056609.1
AY071806.1
AF309350.1
AF309334.1
AF309343.1
AF309335.1
AY138748.1
AF309349.1
AF309341.1
AY138753.1
AF309339.1
AF309342.1
AF309336.1
AY138759.1
AY250967.1
AF309330.1
DQ813722.1
AF392010.1
DQ813725.1
AF392019.1
EU797307.1
EU797308.1
AF392029.1
FN256316.1
AJ850606.1
EF670064.1
EU677511.1
EF517575.1
AF201412.1
AY071777.1
AY071778.1
AJ318666.1
AJ318667.1
AF201406.1
EU797410.1
EU797302.1
EF417063.1
EF417065.1
EU797379.1
FJ818125.1
EU797368.1
AY071803.1
AY071804.1
AY071816.1
FJ819590.1
AY071814.1
AY071788.1
AF201416.1
EU797395.1
EU797396.1
EF517582.1
AF002807.1
EU797418.1
EU797367.1
EF517576.1
EF517587.1
EU797393.1
DQ202611.1
DQ337168.1
HM804293.1
DQ202685.1
EF213912.1
DQ202568.1
EF213843.1
DQ202557.1
EF363013.1
AY748105.1
EF362997.1
AY748103.1
DQ221970.1
DQ222030.1
EF214362.1
DQ221964.1
DQ202590.1
AY748188.1
DQ221962.1
AY424406.1
EU677506.1
EU677658.1
GQ503344.1
HM156718.1
EF484370.1
AJ495454.1
AJ495520.1
AJ495580.1
DQ202653.1
EU797332.1
EU797333.1
AJ849976.1
FJ867781.1
FJ000457.1
HQ883528.1
EU797321.1
DQ202589.1
AJ496336.1
AJ849981.1
AJ496357.1
AY748107.1
AJ849987
AJ850031.1
AJ849988.1
356
HQ883842.1
HQ883854.1
HM002623
HM002618.1
FJ621369.1
DQ487090.1
AJ495468.1
AJ495465.2
AJ495479.1
FJ859959.1
HQ883609.1
DQ155774.1
FJ867887.1
Species
Acmaeodera†
Agrilus angustulus
Anthaxia hungarica
Buprestis†
Castiarina adelaidae
Castiarina rufipennis
Chrysobothris†
Coraebus†
Meliboeus†
Byrrhidae
Byrrhus†
Cantharidae
Cantharis fusca
Cantharis livida
Cantharis rufa
Cantharis rustica
Chauliognathus expansus
Chauliognathus fallax
Chauliognathus flavipes
Chauliognathus lineatus
Chauliognathus octomaculatus*
Chauliognathus riograndensis
Chauliognathus tetrapunctatus
Malthinus†
Oedemera podagrariae*
Oedemera virescens
Rhagonycha fulva
Rhagonycha lignosa
Stenotrachelidae
Cephaloon lepturides
Cerambycidae
Acalolepta fraudatrix
Acalolepta luxuriosa
Acmaeops†
Agapanthia daurica
Anoplophora chinensis†
Anoplophora malasiaca
Apomecyna†
Apriona japonica
Asemum striatum
Batocera rubus
Callidium violaceum
Chloridolum thaliodes
Chlorophorus annularis
Clytus arietis
Cyrtoclytus†
Distenia gracilis
Eburia†
Elaphidon†
Epiglenea comes
Gaurotes†
Glenea†
Leiopus†
Leptura†
Macrotoma†
Mecynippus pubicornis
Mesechthistatus binodosus
Mesosa myops japonica
Monochamus galloprovinciallis
galloprovinciallis*
Monochamus galloprovinciallis pistor
Monochamus saltuarius
Monochamus sartor
Monochamus sutor
Monochamus urussovi
Oberea†
Phaea†
Phymatodes†
Phytoecia†
Plagionotus†
Pogonocherus hispidus
Prionus†
Pterolophia†
Purpuricenus†
Rhagium mordax
Rosalia†
Saperda†
Spondylis buprestoides
Stenygra†
Strangalia†
Stromatium†
Tetraopes femoratus
Tetraopes tetropthamus
Tetropium castaneum
Tetropium fuscum
Tetropium gabrieli
28s
EU797336.1
DQ198702.1
16s
AJ937901.1
DQ198623.1
EF209425.1
DQ202658.1
AB232645.1
DQ402092.1
DQ198705.1
DQ198625.1
18s
AF423771.1
COI
DQ100484.1
EF209485.1
DQ337150.1
DQ337152.1
FJ000486.1
DQ198545.1
EF209545.1
ArgK
Wng
EU797282.1
Elf1a
DQ222000.1
FN298878.1
DQ198548.1
AB298810.1
DQ198767.1
DQ198684.1
AF451940.1
EU677679.1
HM156710.1
DQ198775.1
GU073729.1
DQ198692.1
GU073829.1
AF451938.1
EF209976.1
EF209972.1
DQ198770.1
DQ198687.1
AF451939.1
EF490135.1
EF209980.1
AB533612.1
AB533611.1
GU323028.1
DQ156055.1
DQ198607.1
DQ156062.1
AY095220.1
AY095214.1
AY095216.1
AY095218.1
AY095217.1
AY095215.1
AY095219.1
DQ198615.1
DQ221991.1
EU677654.1
X88963.1
DQ198610.1
AB533643.1
AB533642.1
AF267400
HM046526.1
HM034774.1
AB533608.1
AF332929.1
AB533627.1
AF267398.1
HM046531.1
HM034779.1
HM046533.1
AJ841654.1
HM034781.1
DQ417760.1
DQ417764.1
AJ841404.1
AF267402.1
AJ841525.1
AJ841661.1
HQ832599.1
AF267401.1
HM046547.1
HM034797.1
AY748117.1
AB533598.1
HQ832603.1
EU373304.1
AB533600.1
AB533603.1
HQ832608.1
AF423773.1
AJ841658.1
DQ202535.1
AJ841407.1
AF332932.1
AY748116.1
AJ841529.1
FJ000425.1
HM034784.1
AF267413.1
DQ417769.1
HM062967.1
AY174165.1
AF332945.1
FJ559021
AB533658.1
AY389494.1
FJ559005.1
GU003918
AB457202.1
HM062975.1
GU003932.1
HM062990.1
AM283242.1
AB457192.1
AM283248.1
AB457034.1
EU436853.1
HM062969.1
FJ559020.1
AB533629.1
AB278221.1
HQ890340.1
AY260835.1
AY260836.1
AY260841.1
AY260838.1
AY260843.1
AY260844.1
AB457194.1
AF267467.1
AM283245.1
AF332948.1
DQ155997.1
HM062974.1
FJ559037.1
DQ861317.1
AY748118.1
AJ841659.1
HM034791.1
HM156701.1
EU734906.1
U65185.1
AJ841530.1
AF267399.1
AF267406.1
HM156709.1
U65131.1
DQ861331.1
357
AB457199.1
HM062986.1
EU839772.1
FJ558998.1
AF267481.1
AF267478.1
GU003935.1
GU003936.1
EU883935.1
HM156719.1
Species
Uraecha bimaculata
Xenolea†
Xylotrechus†
Chrysomelidae
Acanthoscelides obtectus
Bruchidius aureus
Bruchidius saundersi
Bruchidius urbanus
Bruchus pisorum
Bruchus rufimanus
Callosobruchus analis
Callosobruchus chinensis
Callosobruchus maculatus
Callosobruchus subinnotatus
Conicobruchus indicus
Pachymerus†
Spermophagus†
Zabrotes subfasciatus
Acalymma blandulum
Acalymma fairmairei
Acalymma trivittatum
Acalymma vittatum
Aedmon†
Agathomerus†
Agelastica coerulea
Agroiconota propinqua
Alagoasa arcifera
Alagoasa bipunctata
Alagoasa cruxnigra
Alagoasa decemguttata
Alagoasa extrema
Alagoasa libentina
Altica lythri
Aphthona lutescens
Apophylia†
Argopus punctipennis
Asiorestia†
Asphaera abdominalis
Asphaerina unicolor
Aspidomorpha difformis
Aulacophora†
Aulacoscelis melanocera
Baliosus†
Basilepta balyi
Blepharida rhois
Bromius obscurus
Calligrapha alni
Calligrapha alnicola
Calligrapha amator
Calligrapha apicalis
Calligrapha bidenticola
Calligrapha californica corepsivora
Calligrapha confluens
Calligrapha fulvipes
Calligrapha multipunctata bigsbyana
Calligrapha philadelphica
Calligrapha polyspila
Calligrapha rowena
Calligrapha scalaris
Calligrapha vicina
Capraita†
Cassida fuscorufa
Cassida rubiginosa
Cerotoma atrofasciata
Cerotoma ruficornis
Chaetocnema chlorophana
Chaetocnema concinna
Chalcophana†
Chalepus†
Charidotella sexpunctata
Chelymorpha cassidea
Chlamisus†
Chrysolina 'aurichalcea'
Chrysolina curvilinea
Chrysolina affinis
Chrysolina americana
Chrysolina bicolor
Chrysolina carnifex
Chrysolina cerealis
Chrysolina coerulans
Chrysolina colasi
Chrysolina diluta
Chrysolina fastuosa
Chrysolina femoralis
Chrysolina gemina
Chrysolina geminata
28s
16s
AB533616.1
AY676680.1
DQ524352.1
DQ524353.1
AY826458.1
DQ307644.1
DQ307645.1
FJ389501.1
AY625366.1
AY625367.1
AY625369.1
HQ178427.1
HQ178551.1
HQ178237.1
18s
EU553813.1
FJ389506.1
FJ000512.1
HQ178212.1
HQ178242.1
AY881194.1
AY243708.2
AY646317.1
AY271867.1
FJ000431.1
AY243678.2
AY927699.1
AY313817.1
AY533638.1
AY533639.1
AY533641.1
AY244857.1
FJ000507.1
AY313827.1
AJ841631.1
AY171444.1
AJ841642.1
AJ781631.1
AY171435.1
AJ781677.1
AY242413.1
EF421506.1
AY308979.1
AF479464.1
AF479438.1
AY308967.1
AY308969.1
AF479471.1
DQ155917.1
DQ155776.1
AY244850.1
FJ973801.1
AJ841348.1
FJ973895.1
AJ841466.1
AJ841383.1
AJ841502.1
AY244864.1
AY533621.1
AJ841392.1
AJ781514.1
AJ841350.1
AJ781551.1
AM160793.1
AM160794.1
AM160806.1
AM160807.1
AM160809.1
AM160810.1
AM160811.1
AM160816.1
AM160822.1
AM160826.1
AM160828.1
AM160833.1
AM160835.1
AM160844.1
AJ841513.1
AJ781568.1
AJ841468.1
AJ781614.1
AY243658.2
AJ841643.1
AY927709.1
AY927701.1
AY171429.1
AJ841548.1
AM283186.1
AF479444.1
AF479448.1
AM283217.1
AY171417.1
AY533565.1
AM283227.1
AM283167.1
AY171408.1
AM160947.1
AM160960.1
AJ575190.1
AJ841418.1
AJ841419.1
AY313828.1
AJ841632.1
AY646323.1
AY646322.1
AJ841599.1
AY676622.1
DQ524356.1
DQ524359.1
DQ524362.1
EF484372.1
AY390695.1
FJ465152.1
AY265224.1
EF570085.1
AY625419.1
HQ177486.1
AY390668.1
AY390669.1
AY881214.1
AF278543.1
AY242442.1
AY533584.1
AY533586.1
AF479421.1
EF421504.1
AY313806.1
AY313803.1
AY313805.1
AY313813.1
AY646304
FJ973693.1
AJ841595.1
COI
AB533647.1
FJ559039
JN000003.1
AY055520.1
AJ575194.1
AJ575192.1
AF479449.1
AB104502.1
AJ841384.1
AY533642.1
AJ841393.1
AF097080.1
AJ008053.1
AJ008037.1
AJ008034.1
AJ008054.1
AJ008058.1
AJ008055.1
AF097083.1
AJ008049.1
AJ008057.1
AJ008056.1
AJ008060.1
AJ008051.1
AJ008042.1
AY676687.1
AM283218.1
AY533587.1
AJ841470.1
AM283190.1
DQ155710.1
AY242399.1
AM283228.1
AM283219.1
AY244842.1
AJ841514.1
AJ841504.1
AF267433.1
AJ622062.1
AJ841421.1
358
AF092680.1
AY796210.1
AJ008086.1
AJ008079.1
AJ008080.1
AJ008082.1
AJ008083.1
AJ008084.1
AY171403.1
AJ008085.1
AJ008087.1
AJ008088.1
AJ008089.1
AJ008091.1
AJ008092.1
ArgK
Wng
Elf1a
AY997367.1
AF466312.1
EF421464.1
AF466334.1
AF466336.1
AF466338.1
EF421466.1
AF466323.1
AY171468.1
AY171460.1
AM160847.1
AM160849.1
AM160859.1
AM160860.1
AM160863.1
AM160864.1
AM160867.1
AM160873.1
AM160876.1
AM160879.1
AM160883.1
AM160885.1
AM160893.1
AF466324.1
AY171454.1
Species
Chrysolina graminis
Chrysolina grossa
Chrysolina haemoptera
Chrysolina herbacea
Chrysolina lucida
Chrysolina marginata
Chrysolina obsoleta
Chrysolina peregrina
Chrysolina polita
Chrysolina timarchoides
Chrysolina varians
Chrysolina veridana
Chrysomela interrupta
Chrysomela lapponica
Chrysomela mainensis interna
Chrysomela populi
Chrysomela schaefferi
Chrysomela tremulae
Colaspis†
Colasposoma auripenne
Coptocephala†
Coptocycla adamantina
Crioceris asparagi
Crioceris duodecimpunctata
Crioceris paracenthesis
Cryptocephalus venustus
Cyrtonus arcasi
Cyrtonus contractus
Cyrtonus cupreovirens
Cyrtonus cylindricus
Cyrtonus dufouri
Cyrtonus elegans
Cyrtonus fairmarei
Cyrtonus majoricensis
Cyrtonus pardoi
Cyrtonus plumbeus
Cyrtonus puncticeps
Cyrtonus rotundatus
Cyrtonus ruficornis
Dactylispa†
Deloyala guttata
Desmogramma†
Diabrotica adelpha
Diabrotica balteata
Diabrotica cristata
Diabrotica limitata 15-punctata
Diabrotica longicornis
Diabrotica nummularis
Diabrotica porracea
Diabrotica scutella
Diabrotica sexmaculata
Diabrotica speciosa
Diabrotica tibialis
Diabrotica undecimpunctata
Diabrotica undecimpunctata howardi
Dibolia†
Dicladispa testacea
Diorhabda†
Disonycha†
Donacia bicolor
Donacia biimpressa
Donacia clavipes
Donacia hirticollis
Donacia subtilis
Donacia vulgaris
Doryphora†
Entomoscelis†
Epitrix fasciata
Eumolpus†
Exema canadensis
Exora encaustica narensis
Galeruca†
Galerucella bimarnica
Galerucella calmariensis
Galerucella lineola
Galerucella nymphae
Galerucella placida
Galerucella pusilla
Galerucella tenella
Gastrophysa cyanea
Gastrophysa poligoni
Gastrophysa viridula
Glyptina†
Gonioctena linnaeana
Gonioctena sibirica*
Gonioctena springlovae
28s
AJ841549.1
AJ841550.1
AY171423.1
AJ781655.1
AJ781648.1
AJ841582.1
AY243649.2
AJ781684.1
FJ000446.1
16s
AF097086.1
AF097087.1
AJ008044.1
AJ008052.1
AJ008059.1
AJ008062.1
AJ008041.1
AJ008038.1
AJ008039.1
AJ008040.1
AJ008043.1
AJ008036.1
AY027728.1
AY027738.1
AY027735.1
AY027744.1
AY027756.1
AY027758.1
AJ781532.1
AJ781527.1
AJ841336.1
AJ781558.1
18s
COI
AF097030.1
ArgK
Wng
Elf1a
AJ841422.1
AJ841423.1
FJ000527.1
AJ781592.1
AJ781585.1
AJ841453.1
AY244836.1
AF267426.1
FJ000520.1
AJ008093.1
AJ008095.1
AJ008097.1
AJ008099.1
AJ008101.1
AJ008102.1
AJ008103.1
AJ008105.1
AJ008106.1
AJ008108.1
AY027610.1
AY027620.1
AY027617.1
AY242404.1
AY171397.1
AM283171.1
AY171449.1
AM283156.1
AY242390.1
AM283148.1
EU880722.1
AJ231163.1
AJ841589.1
AM403398.1
AM403402.1
AM403403.1
AM403405.1
AM403406.1
AM403408.1
AM403410.1
AM403416.1
AM403417.1
AM403422.1
AM403423.1
AM403424.1
AM403426.1
AJ841644.1
AJ841635.1
AJ841551.1
AY243735.2
AY243731.2
AJ841460.1
AM403365.1
AM403369.1
AM403370.1
AM403372.1
AM403373.1
AM403375.1
AM403377.1
AM403383.1
AM403384.1
AM403389.1
AM403390.1
AM403391.1
AM403393.1
AJ841394.1
AJ841386.1
AJ841308.1
AJ841515.1
AJ841506.1
AJ841424.1
AY533625.1
AF195200.1
AY533624.1
AY533627.1
AY533623.1
AY533622.1
AY533635.1
AY533632.1
AY533628.1
AJ781555.1
EF421514.1
AJ231164.1
AJ781618.1
FJ973917.1
AJ841516.1
AY243747.2
AY243737.2
AY646319.1
AY243746.2
AY243738.2
AY171442.1
AY171446.1
AY171434.1
AJ841472.1
EF532415.1
EF532418.1
EF532550.1
EF532553.1
AY232588.1
AJ841552.1
AJ841309.1
AJ008077.1
AY243668.2
AJ841579.1
AY243762.2
AJ841333.1
AJ841367.1
EF421521.1
EF116927.1
EF421518.1
EF421516.1
FJ973717.1
AY243664.2
FJ973721.1
AY748122.1
AJ841425.1
AY244848.1
AF267460.1
AJ841450.1
AJ841486.1
AJ841488.1
AF267443.1
FJ973920.1
AY027751.1
EF421520.1
AY027752.1
FJ973824.1
AF014604.1
AB104501.1
AB104498.1
AM283221.1
AM283125.1
AY242469.1
AY533569.1
AY533580.1
AY242481.1
AF278547.1
AY533568.1
AY533571.1
AY533567.1
AY533566.1
AY533579.1
AY533576.1
AY533572.1
AM283202.1
EF421472.1
EF421474.1
AM283229.1
AY171419.1
AY171407.1
EU880600.1
EF532505.1
EF532508.1
EU880682.1
EU880714.1
EU880631.1
AM283126.1
AJ008110.1
AY242409.1
AY171469.1
AY171459.1
EU880749.1
EU880809.1
EU880789.1
EU880757.1
EU880781.1
AM283153.1
AY242497.1
EF421481.1
EF177824.1
AY247725.1
AY247726.1
AY247717.1
EF133454.1
AY247724.1
AY247723.1
EF421478.1
EF421476.1
AY244843.1
EF421480.1
FJ973924.1
FJ973925.1
DQ156057.1
AJ622061.1
FJ346979.1
359
FJ346980.1
Species
Gonioctena variabilis
Gonioctena viminalis
Gynandrobrotica lepida
Gynandrobrotica nigrofasciata
Heikertingerella†
Hemipyxis†
Hermaeophaga†
Hilarocassis exclamationis
Hypolampsis†
Isotes multipunctata
Labidomera clivicollis
Laccoptera†
Lachnaia†
Lema†
Leptinotarsa haldemani
Leptinotarsa juncta
Leptinotarsa texana
Lilioceris†
Linaeidea aenea
Lochmaea capreae
Longitarsus exsoletus
Luperus†
Lysathia ludoviciana
Macrohaltica†
Macrolenes dentipes
Microrhopala vittata
Monolepta†
Monoxia†
Neocrepidodera†
Nodonota†
Notosacantha†
Octotoma scabripennis
Oedionychus cinctus
Oides†
Omophoita octoguttata
Omophoita personata
Omophoita sexnotata
Ophraella communa
Ophraella notulata*
Ophraella pilosa
Ophraella sexvittata
Ophrida†
Oreina alpestris
Oreina cacaliae
Oreina elongata
Oreina ganglbaueri
Oreina intricata
Oreina variabilis
Oreina virgulata
Oulema melanopus
Pachybrachis†
Paranaita bilimbata
Paratriarius†
Paria†
Paridea†
Paropsis atomaria
Paropsis porosa*
Paropsisterna beata
Phaedon†
Philhydronopa†
Phratora laticollis
Phratora tibialis
Phyllotreta†
Plagiodera versicolora
Plagiometriona†
Platyphora†
Podagrica malvae
Podontia†
Prasocuris†
Psylliodes affinis
Psylliodes chrysocephalus
Psylliodes cupreus
Psylliodes dulcamarae
Psylliodes marcidus*
Psylliodes napi
Pyrrhalta viburni
Sagra femorata
Sermylassa halensis
Smaragdina†
Sphaeroderma†
Stolas angulata
Syphraea†
Systena†
Timarcha aurichalcea
Timarcha balearica
Timarcha calceata
28s
AY243718.2
AY243717.2
FJ973723.1
FJ973724.1
FJ973725.1
AY927718.1
16s
AY027750.1
AF014601.1
AY533643.1
AY533644.1
FJ973826.1
FJ973827.1
EF421522.1
18s
FJ973927.1
FJ973928.1
FJ973929.1
AJ841312.1
AJ841388.1
AJ841338.1
AJ841557.1
AJ841314.1
AJ841430.1
AJ841574.1
AJ781682.1
FJ973738.1
AJ841604.1
AY646336.1
AJ841328.1
AJ781556.1
EF421524.1
AJ841356.1
AF267425.1
AJ622058.1
FJ973942.1
AJ841475.1
AY748123.1
AJ841605.1
AJ841357.1
AJ231165.1
AJ841400.1
EF421526.1
U20682.1
AJ841476.1
AJ781661.1
FJ000447.1
AJ841651.1
FJ973757.1
AY646292.1
AY313838.1
AY313839.1
AY243783.2
U20713.1
FJ973759.1
AJ841558.
AJ841577.1
AJ841592.1
AY313840.1
AY243728.2
AJ781640.1
AY243696.2
AJ781537.1
AJ841401.1
FJ973853.1
EF421528.1
U20694.1
U20710.1
AF267437.1
AJ841508.1
AJ841455.1
AY676686.1
AJ841521.1
AJ841494.1
EF362972.1
AJ781598.1
FJ000521.1
AF267431.1
FJ973962.1
AF267438.1
HM036738.1
AJ841495.1
U20715.1
FJ973854.1
GQ392138.1
HQ435544.1
GQ220082.1
GQ392195.1
AF097102.1
AF097106.1
AF097107.1
AJ841331.1
AJ841346.1
AJ841448.1
AJ841463.1
AJ781522.1
AJ781578.1
AY171438.1
FJ973964.1
AJ841431.1
AJ841432.1
AJ841560.1
AY027760.1
AJ841433.1
AJ841561.1
AY027753.1
AY027754.1
FJ973861.1
AJ841319.1
AF267435.1
FJ973766.1
AJ841562.1
ArgK
Wng
Elf1a
AY533588.1
AY533589.1
AY243723.2
AJ841555.1
AJ841637.1
AJ841584.1
AY243659.2
AY243650.2
AJ841623.1
AY243778.2
COI
FJ973971.1
AJ841435.1
AF466316.1
EF421482.1
DQ155806.1
AF479423.1
AY242457.1
AM283129.1
AM283223.1
AM283157.1
AY242400.1
DQ459377.1
HQ605771.1
HQ605774.1
AM283150.1
AY027616.1
AY247728.1
AM283194.1
AF466314.1
EF421483.1
EF421485.1
EU117150.1
AM283195.1
AM283234.1
AM283209.1
AY242512.1
EF421487.1
AM283235.1
AF479429.1
AY796204.1
AF479430.1
AF479431.1
AF479441.1
HQ664676.1
AY242517.1
EF421489.1
AF466317.1
AF466318.1
FJ977975.1
GQ392324.1
GQ392437.1
GQ220210.1
GQ392378.1
AF097040.1
AF097044.1
AF097045.1
AM283164.1
AF479481.1
AY533591.1
AF466343.1
DQ335220.1
AY171411.1
FM209242.1
DQ155742.1
FM209237.1
AM283136.1
FJ977982.1
AY171463.1
FM209275.1
FM209270.1
EU420057.1
EF421494.1
GQ268947.1
AY055518.1
AJ841609.1
FJ973769.1
AJ841563.1
AJ841361.1
AJ841480.1
FJ973974.1
AJ622059.1
AJ841610.1
EF421531.1
AJ841481.1
AJ841626.1
HQ178449.1
AY243676.2
AJ841585.1
FJ973776.1
AY927738.1
AJ841611.1
AY171432.1
AJ841378.1
HQ178243.1
EF421535.1
AJ841339.1
EF421537.1
AJ841497.1
FJ000515.1
FJ973978.1
AJ841456.1
FJ973981.1
FJ977985.1
AM283138.1
DQ155877.1
EU110830.1
EU110835.1
EU110836.1
EU110850.1
EU110853.1
AY247731.1
HQ177510.1
FJ977988.1
AM283158.1
FJ977991.1
AJ841363.1
FJ973874.1
AJ231136.1
AJ236387.1
Y18828.1
AJ841482.1
FJ973986.1
AM283199.1
AY171405.1
AM403397.1
AJ841320.1
360
EU110898.1
EU110901.1
EU110927.1
EU110928.1
EF421492.1
EU110914.1
EF421496.1
EF421497.1
EF421499.1
AY171457.1
Species
Timarcha cornuta
Timarcha cyanescens
Timarcha erosa vermiculata
Timarcha espanoli
Timarcha fallax
Timarcha geniculata
Timarcha goettingensis
Timarcha gougeleti
Timarcha granadensis
Timarcha hispanica
Timarcha insparsa
Timarcha intermedia
Timarcha interstitialis
Timarcha intricata
Timarcha lugens
Timarcha marginicollis
Timarcha maritima
Timarcha metallica
Timarcha monserratensis
Timarcha perezi
Timarcha pimelioides
Timarcha recticollis
Timarcha rugosa
Timarcha scabripennis
Timarcha sinuatocollis
Timarcha strangulata
Timarcha tenebricosa
Timarcha ventricosa
Trirhabda canadensis
Trirhabda virgata
Walterianella†
Xanthogaleruca luteola
Yingaresca†
Zygogramma bigenera
Ciidae
Cis†
Octotemnus†
Sulcasis†
Cleridae
Enoclerus moestus
Thanasimus†
Trichodes apiarius
Trichodes nutalli
Trichodes ornatus
Coccindellidae
Adalia bipunctata
Aiolocaria†
Anatis†
Azya†
Calvia quatruordecimguttata
Chilocorus bipustulatus
Chilocorus renipustulatus
Chilocorus rubidus
Chilocorus stigma
Chnoodes†
Coccinella septempunctata
Coelophora†
Coleomegilla maculata lengi
Cryptognatha†
Cryptolaemus montrouzieri
Curinus coeruleus
Cycloneda sanguinea
Epilachna†
Exochomus quadripustulatus
Hippodamia convergens
Hippodamia parenthesis
Hippodamia quinquesignata ambigua
Hyperaspis lateralis lateralis
Illeis†
Micraspis cardoni
Microweisea†
Olla†
Orcus†
Prodilis†
Psyllobora taedata
Rhyzobius†
Rodolia cardinalis
Scymnus nubilis
Sumnius†
Curculionidae
Cactopinus†
Cryphalus abietis
Crypturgus pusillus
Dendroctonus adjunctus
Dendroctonus approximatus
Dendroctonus brevicomis
28s
FJ000452.1
16s
AJ278896.1
AJ231139.1
AJ622029.1
AJ231140.1
Y18827.1
AJ236380.1
AJ231142.1
AJ236381.1
AJ231143.1
AJ231144.1
AJ231145.1
AJ236384.1
AJ236368.1
AJ622026.1
AJ231149.1
AJ231150.1
AJ622030.1
Y18826.1
AJ236374.1
AJ008078.1
Y18824.1
AJ231154.1
AJ231155.1
AJ622038.1
AJ236372.1
AJ231157.1
AJ231158.1
AJ622039.1
AJ841628.1
AY313794.1
18s
COI
AJ841324.1
DQ202638.1
FJ000412.1
FJ000528.1
AY171412.1
AJ841499.1
AY244853.1
AY515056.1
AY515115.1
AF479456.1
AJ841441.1
AY515034.1
AY171413.1
AY748191.1
EF209887.1
DQ337129.1
EF517585.1
Wng
Elf1a
AJ008111.1
EF421538.1
AJ841568.1
ArgK
AY748128.1
AY748129.1
EF209693.1
AY171464.1
AF466327.1
EF421500.1
AY171465.1
DQ156020.1
FM877949.1
AY790473.1
GU013585.1
EU145713.1
AF423775.1
AJ272139.1
GU073731.1
FJ687707.1
GU073732.1
GU073768.1
GU073769.1
GU073833.1
AM779600.1
GU073875.1
GU073876.1
GU073676.1
FJ687666.1
GU073677.1
GU073718.1
GU073719.1
X88933.1
EF192080.1
GU073920.1
EF192097.1
GU073959.1
GU073960.1
EF192085.1
EU145610.1
JF763542.1
DQ202668.1
FJ687721.1
GU073740.1
JF763548.1
FJ687709.1
FJ687723.1
AB353862.1
FJ687736.1
EU164644.1
FJ687715.1
FJ687726.1
JF763569.1
FJ687720.1
JF763575.1
GU073748.1
FJ687740.1
JF763589.1
EU145666.1
GU073773.1
GU073776.1
GU073764.1
JF763613.1
GU073845.1
DQ202558.1
AY748147.1
FJ687679.1
GU073688.1
GU073865.1
FJ687668.1
AB354089.1
AM779604.1
EU164588.1
FJ687681.1
EU145616.1
EF209855.1
EU164617.1
AJ313071.1
AY615732.1
GU073908.1
AY283622.1
AB002178.1
DQ155759.1
EU164681.1
JF296238.1
FJ687673.1
FJ687685.1
FJ687680.1
FJ687678.1
EF192090.1
EF192086.1
GU073855.1
FJ687675.1
FJ687699.1
EU164680.1
JF763665.1
GU073859.1
EF512342.1
GU073883.1
GU073871.1
EU145604.1
EF512320.1
GU073726.1
GU073714.1
GU073944.1
DQ155761.1
GU073967.1
GU073955.1
AF375247.1
AF187109.1
EU011813.1
AF067992.1
AF068000.1
AF068002.1
EU090343
AF308380.1
AF308332.2
361
EU011826.1
AF308423.1
Species
Dendroctonus frontalis
Dendroctonus jeffreyi
Dendroctonus murrayanae
Dendroctonus ponderosae
Dendroctonus pseudotsugae
Dendroctonus punctatus
Dendroctonus rufipennis
Dendroctonus simplex
Dendroctonus terebrans
Dendroctonus valens
Dryocoetes affaber
Dryocoetes autographus
Hylastes†
Hylurgops†
Ips acuminatus
Ips avulsus
Ips bonanseai
Ips borealis
Ips calligraphus calligraphus
Ips confusus
Ips cribricollis
Ips emarginatus
Ips hoppingi
Ips hunteri
Ips integer
Ips knausi
Ips lecontei
Ips montanus
Ips paraconfusus
Ips perroti
Ips pertubatus
Ips pilifrons
Ips pini
Ips plastographus maritimus
Ips sexdentatus
Ips spinifer
Ips tridens
Ips typographus
Ips woodi
Orthotomicus caelatus
Orthotomicus erosus
Orthotomicus laricis
Orthotomicus latidens
Orthotomicus suturalis
Phloeotribus†
Pityogenes chalcographus
Pityogenes hopkinsi*
Polygraphus grandiclava
Polygraphus poligraphus*
Scolytus†
Trypodendron lineatum
Xyleborus†
Xylosandrus compactus
Xylosandrus germanus
Acalles camelus
Acalles echinatus
Adexius scrobipennis
Anthonomus grandis
Apion abruptum
Apoderus jekeli
Auletobius†
Baris arthemisiae
Barynotus moerens
Barypeithes†
Bothrometopus†
Brachyderes incanus
Callirhopalus bifasciatus
Centricnemus leucogrammus
Ceutorrhynchus erysimi
Chlorophanus†
Cionus†
Cleonus†
Cosmopolites sordidus
Curculio dentipes
Curculio distinguendus
Curculio nucum*
Cylas formicarius
Diocalandra†
Dorytomus†
Dyslobus†
Echinocnemus marmoreus
Eusomus ovulum
Foucartia squamulata
Gymnetron antirrhini
Gymnetron tetrum
Hylobius abietis
28s
AF308381.1
AF308382.1
AF308384.1
AF308385.1
AF308374.2
16s
AF308386.1
18s
AF308333.1
AF308334.1
AF308336.1
AF308337.1
AF308327.1
AF308338.1
AJ850046.1
HQ883565.1
HQ883927.1
AF308364.1
EU090296.1
AJ495569.1
AF397470.1
AF397472.1
AF308390.1
EU090302.1
AF397474.1
AF397475.1
AF397477.1
AF397478.1
AF397480.1
AF397482.1
AF397483.1
AF397484.1
AF397485.1
AF397486.1
AF397488.1
AF397489.1
AF397490.1
AF397491.1
AF397493.1
AF397494.1
AF397492.1
AF397496.1
AF397506.1
AF397498.1
AF397499.1
AF397500.1
AF397501.1
AF397502.1
EU090300.1
AF397503.1
AJ496371.2
AJ850040.1
AF308343.1
AJ850041.1
AF308372.1
AF308363.1
AF375306.1
AF308394.1
GU808581.1
GU808591.1
GU808598.1
EU286365.1
EU286428.1
EU286372.1
AF308325.1
AF397509.1
AF389045.1
AJ495567.1
AF308316.1
AJ850043.1
AF250076.2
AJ850045.1
EU286282.1
EU286346.1
EU286289.1
FJ423738.1
AJ495474.1
HQ883529.1
FJ000465.1
AF250091.2
COI
AF067986.2
AF067994.2
AF067989.1
AF067987.1
AF375318.1
AF067998.1
AF067996.1
AF067985.1
AF375315.1
AF067997.1
AF187113.1
AF438517.1
HQ883660.1
AF113325.1
AF113330.1
AF113332.1
AF113334.1
AF113335.1
AF113341.1
AF113343.1
AF113347.1
AF113353.1
AF113355.1
AF113356.1
AF113357.1
AF113360.1
AF113365.1
AF113367.1
AF113369.1
AF113370.1
AF113373.1
AF113376.1
AF113378.1
AF113380.1
AF113381.1
AF113384.1
AF113385.1
AF113388.1
AF187124.1
U82236.1
AF113392.1
AF113358.1
EU191854.1
EU191863.1
DQ515997.1
AF187128.1
EU428829.1
EU428842.1
AF375329.1
AF187132.1
HM064121.1
GU808706.1
GU808714.1
EU286447.1
EU286510.1
EU286454.1
AY266610.1
DQ155878.1
HQ883610.1
ArgK
Wng
Elf1a
AF308424.1
AF308425.1
AF308427.1
AF308428.1
AF308418.1
AF308429.1
HQ883880.1
AF186661.2
AF259873.1
HQ883732.1
AF308408.1
AF397612.1
AF397615.1
AF397616.1
AF397617.1
AF397618.1
AF397620.1
AF397621.1
AF397623.1
AF397625.1
AF397626.1
AF397627.1
AF397628.1
AF397629.1
AF397630.1
AF397631.1
AF397632.1
AF397633.1
AF397635.1
AF397636.1
AF397634.1
AF397637.1
AF397644.1
AF397638.1
AF397639.1
AF397640.1
AF186672.1
AF397642.1
AF397643.1
EU191886.1
EU191895.1
AF186676.1
HQ883910.1
HQ883923.1
HM064385.1
GU808668.1
GU808674.1
AF308407.1
HQ883746.1
AF186682.1
AF508870.1
GU808743.1
GU808751.1
HQ883843.1
HQ883698.1
HQ883697.1
HQ883882.1
GU565646.1
DQ058708.1
HQ883727.1
FJ000541.1
AJ495565.1
AJ495512.1
AJ495500.1
AJ496356.2
AJ496350.1
DQ155809.1
GQ132012.1
AJ495503.1
AF250079
DQ155881.1
HQ883651.1
HQ883566.1
AJ495550.1
AJ495524.1
AJ496366.1
AJ496359.1
AY131082.1
FJ867675.1
AF389059.1
AF515703.1
AY131075.1
AJ495533.1
EF606981.1
FJ867679.1
AJ495499.1
AJ495501.1
FJ867684.1
AJ850011.1
AY131111.1
AB501115.1
AB501118.1
AY327696.1
AY131140.1
AY131104.1
DQ156033.1
EF575495.1
AY131133.1
FJ867755.1
AJ496349.2
AJ496351.2
AJ850004.1
AF250096.2
AJ495585.1
FJ859938.1
FJ859941.1
EF450122.1
362
AY327661.1
EF577091.1
FJ867864.1
Species
Hylobius piceus
Hypera nigrirostris
Hypera postica
Hypera punctata
Hypera rumicus
Hypomeces squamosus
Lepyrus†
Limnobaris†
Liophloeus tessulatus
Listroderes†
Lixus†
Magdalis†
Metamasius hemipterus
Metialma†
Naupactus peregrinus
Naupactus xanthographus
Nedyus quadrimaculatus
Notaris†
Otiorhynchus armadillo
Otiorhynchus raucus
Pachylobius picivorus
Peritelus†
Phyllobius argentatus
Phyllobius oblongus
Phyllobius pyri
Phytobius†
Pissodes nemorensis
Pissodes schwarzi*
Pissodes strobi
Polydrusus inustus
Polydrusus mollis
Polydrusus sericeus
Polydrusus undatus
Procas†
Rhinoncus †
Rhynchaenus†
Rhynchites†
Rhynchophorus ferrugineus
Ruteria hypocrita
Scepticus griseus
Scepticus insularis
Sciaphilus asperatus
Sitona hispidulus
Sitona lineatus
Sitona suturalis
Sitophilus granarius
Sitophilus oryzae
Sitophilus zeamais
Sphenophorus†
Strophosomus capitatum
Strophosomus melanogrammum
Sympiezomias cribicollis
Tanymecus†
Tanysphyrus lemnae
Telephae†
Trachyphloeus†
Tropiphorus carinatus
Tychius†
Zacladus geranii
Eudiagogus†
Dermestidae
Anthrenus†
Attagenus†
Dermestes maculatus
Dermestes peruvianus
Orphilus subnitidus
Trogoderma glabrum
Trogoderma variabile
Elateridae
Adelocera†
Agriotes lineatus
Agriotes obscurus
Agriotes sputator
Agrypnus†
Athous†
Cardiophorus†
Chalcolepidius†
Conoderus dimidiatus
Ctenicera mediana
Elater†
Lacon profusa
Limonius†
Melanotus†
Erotylidae
Triplax thoracica
Histeridae
28s
HQ883576.1
16s
U16967.1
U16968.1
18s
COI
HQ883665.1
AJ850005.1
FJ867764.1
ArgK
HQ883894.1
Wng
Elf1a
FJ859943.1
DQ155880.1
EU522679.1
AJ495586.1
AJ495594.1
AJ495497.1
HQ883562.1
FJ867691.1
AY131079.1
HQ883536.1
AJ495518.1
AJ495593.1
FJ867695.1
AJ495560.1
FJ000469.1
AJ496375.1
AJ496378.1
AJ496348.1
AF389057.1
AF250098.2
AJ496377.1
AJ850017.1
AF250083.2
FJ867775.1
AJ850048.1
FJ000545.1
AJ495480.1
AJ495574.1
HQ883623.1
FJ859947.1
AY131108.1
HQ883617.1
HQ452972.1
EU111007.1
EF517599.1
HQ883849.1
FJ859952.1
HQ883708.1
FJ867873.1
AY131137.1
HQ883703.1
FJ867878.1
EU111038.1
AY165668.1
AF250103.2
AJ495485.1
DQ156044.1
AJ495492.3
AJ495491.1
FJ867702.1
AJ850014.1
AF250094.2
AJ850007.1
DQ156035.1
FJ859957.1
U77981.1
U77977.1
AY472043.1
HQ223014.1
HQ223010.1
HQ223023.1
AF250086.2
AJ495496.1
HQ883539.1
AJ495561.1
AJ495549.1
AJ495451.2
EU286370.1
AB663245.1
AB663244.1
EU286287.1
AJ850025.1
AF389061.1
AJ849982.1
EF125057.1
HQ883620.1
DQ156022.1
HQ883852.1
EU780007.1
EU286452.1
AJ495502.1
FJ867710.1
AJ495508.1
AJ495510.1
AY131072.1
AY131070.1
AY131071.1
FJ867714.1
EU131469.1
AJ495504.1
AJ495505.1
FJ867719.1
AY131069.1
AJ495507.1
AJ495596.1
AJ495516.1
AJ495488.1
AJ495544.1
AF250087.2
AJ496354.1
AF250070.2
AJ878601.1
AJ850021.1
FJ867793.1
AJ496352.1
AJ496353.1
AF250088.2
AJ850023.1
AJ850016.1
AJ496365.1
HQ883540.1
DQ155755.1
AY131101.1
AY131099.1
FJ859963.1
FJ859965.1
AY196875.1
EU522680.1
FJ867822.1
AY131098.1
FJ859970.1
DQ155813.1
AY837616.1
HQ883853.1
AY131130.1
AY131128.1
AY131129.1
FJ867894.1
HQ883706.1
AF250081.2
EF213924.1
DQ202647.1
EF213915.1
EF213862.1
DQ202544.1
HM051209.1
DQ202548.1
EF213848.1
EF213894.1
AY748110.1
AY748111.1
EF213882.1
HM243241.1
AB231216
DQ198733.1
DQ198734.1
DQ198735.1
DQ198738.1
HQ333885.1
HQ333847.1
HQ333842.1
HQ333861.1
HQ333884.1
AB231252.1
AB231253.1
AY310698.1
DQ198643.1
DQ198644.1
EU285480.1
DQ198645.1
DQ198648.1
HQ333701.1
HQ333671.1
HQ333666.1
EF213948.1
DQ221954.1
FJ763718.1
FJ589737.
HM243385.1
DQ155913.1
AF451943.1
AF451944.1
HQ333790.1
HQ333752.1
HQ333747.1
DQ155945.1
DQ198567.1
DQ198570.1
HQ333972.1
HQ333937.1
HQ333683.1
HQ333766.1
HQ333789.1
AY165675.1
HQ333949.1
HQ333971.1
HQ333673.1
AB298868.1
EF424482.1
AY310637.1
363
EU414679.2
Species
28s
Hister†
Saprinus†
Hydrophilidae
Berosus†
Helophorus aquaticus
Hydrobius fuscipes
Sternolophus rufipes
Tropisternus†
Laemophloeidae
Cryptolestes†
Lampyridae
Aspisoma aegrotum*
Bicellonycha†
Ellychnia californica
Ellychnia corrusca
Lucidota†
Luciola cruciata
Luciola lateralis
Photinus australis
Photinus pyralis
Photuris congener
Photuris pennsylvanica
Pyractomena angulata
Pyractomena borealis
Lucanidae
Dorcus parallelipipedus
Lucanus†
Prosopocoilus†
Lycidae
Calopteron†
Metriorrhynchus†
Melandryidae
Melandryidae†
Tetratomidae
Penthe†
Meloidae
Epicauta†
Mylabris†
Pryrota decorata
Tetraonyx†
Zonitis†
Melyridae
Melyridae†
Nitidulidae
Nitidula†
Passalidae
Aulacocyclus†
Odontotaenius disjunctus
Paxillus†
Phalacridae
Phalacridae†
Scarabaeidae
Adoretus lasiopygus
Adoretus versutus
Amphimallon solstitialis
Anomala bengalensis
Anomala dorsalis
Anomala polita
Anomala varicolor
Anoplotrupes stercorosus
Aphodius depressus
Aphodius luridus
Aphodius ater
Aphodius constans
Aphodius rufus
Aphodius lineolatus
Aphodius paykulli
Aphodius sticticus
Aphodius erraticus
Aphodius consputus
Aphodius prodromus
Aphodius sphacelatus
Aphodius contaminatus
Aphodius obliteratus
Aphodius haemorrhoidalis
Aphodius borealis
Aphodius vittatus mundus
Aphodius fossor
Aphodius conjugatus
Aphodius distinctus
Aphodius elevatus
Aphodius fimetarius
Aphodius foetens
Aphodius foetidus
Aphodius moestus
Aphodius pedellus
AJ810756.1
AJ810749.1
AJ810755.1
FJ818242.1
EU797397.1
18s
AY028360.1
AY028341.1
COI
JF794585.1
16s
AM287065.1
AM287056.1
AM287070.1
AJ810721.1
AJ810714.1
AJ810720.1
AM287087.1
AM287078.1
AM287092.1
FJ819808.1
EU797419.1
ArgK
Wng
Elf1a
EU797334.1
EF362981.1
AB232651.1
EU009285.2
EU009265.1
EU009255.2
EU009262.1
EU009256.1
AB436492.1
AB436502.1
EU009261.1
EU009248.1
EU009228.1
EU009218.1
EU009225.1
EU009219.1
AB298851.1
EU009224.1
EU009322.1
EU009302.1
EU009292.1
EU009299.1
EU009293.1
AB608760.1
AF360873.1
EU009298.1
DQ007905.1
EU301845.1
EU009270.1
EU009259.1
AB178302.1
AB178297.1
DQ181127.1
DQ181114.1
U65129.1
EU009233.1
EU009222.1
AY165656.1
EU009307.1
EU009296.1
AY745569.1
EF487586.1
DQ156023.1
AB110733.1
AB110728.1
AB298856.1
DQ181040.1
DQ181201.1
DQ181188.1
DQ202620.1
EF490139.1
EF209898.1
EF490167.1
DQ202639.1
EF517586.1
EF209892.1
DQ222035.1
DQ202670.1
AM712128.1
AJ633652.1
AJ633668.1
AM712135.1
AM712134.1
FJ000495.1
DQ222031.1
FJ462782.2
FJ000492.1
AY165674.1
DQ202556.1
AY748176.1
DQ155899.1
EF487938.1
EF487588.1
AY745573.1
DQ534686.1
DQ028959.1
EF209792.1
FM877920.1
EF487668.1
DQ524384.1
DQ524388.1
DQ295268.1
DQ524510.1
DQ524512.1
FJ000414.1
DQ202667.1
AF098384.1
DQ202623.1
DQ202524.1
DQ524609.1
DQ524614.1
EU084161.1
DQ524741.1
DQ524743.1
DQ524742.1
DQ524628.1
DQ680893.1
DQ680864.1
EF487948.1
DQ680966.1
AY132447.1
GU187736.1
DQ524518.1
EF487857
EF487992.1
EF487579.1
EF487597.1
AY132509.1
AY132510.1
AY132415.1
AY132514.1
AY132516.1
EF487610.1
EF487619.1
EF487813.1
EF487632.1
EF487991.1
EF487600.1
364
FJ606577.1
FJ606563.1
AY132556.1
AY132558.1
AY132561.1
AY132581.1
DQ155727.1
AY132583.1
AY132591.1
AY132593.1
AY132596.1
AY132607.1
AY132608.1
AY132615.1
AY132525.1
AY132553.1
AY132523.1
AY132527.1
AY132529.1
AY132528.1
AY132588.1
AY132532.1
Species
Apogonia†
Ataenius†
Bolbites onitoides
Bolboceras†
Bubas bison
Bubas bubalus
Caccobius schreberi
Canthidium†
Canthon indigaceus
Cetonia aurata
Cheironitis furcifer
Chiron digitatus
Copris lugubris
Copris lunaris
Copris sinicus
Coprophanaeus †
Cremastocheilus†
Cyclocephala†
Deltochilum aff amazonicum
Diabroctis mimas
Dichotomius bos
Dichotomius geminatus
Dichotomius nisus
Dichotomius semisquamosus
Dichotomius sericeus
Digitonthophagus gazella
Dynastes†
Eucranium arachnoides
Eurysternus caribaeus
Geotrupes spiniger
Glaresis†
Glyphoderus sterquilinus
Gromphas†
Gymnopleurus†
Haplidia†
Heliocopris†
Hoplia†
Hybosorus†
Lichnanthe rathvoni
Malagoniella†
Melolontha†
Microcopris†
Mimela†
Oniticellus fulvus
Onitis†
Ontherus†
Onthophagus fracticornis
Onthophagus hecate
Onthophagus marginicollis
Onthophagus nuchicornis*
Onthophagus pennsylvanicus
Onthophagus similis
Onthophagus taurus
Orphnus†
Oruscatus davus
Oryctes nasicornis
Oxysternon†
Oxythyrea†
Paracopris†
Paragymnopleurus sinuatus
Pentodon†
Phanaeus chalcomelas
Phanaeus igneus
Phanaeus vindex
Phyllognathus dionysius
Phyllopertha campestris
Phyllophaga delata*
Phyllophaga drakii
Phyllophaga fusca
Pleocoma†
Popillia japonica
Protaetia cuprea bancoi
Psammodius oregonesis
Scarabaeus†
Schizonycha†
Serica†
Sisyphus†
Sulcophanaeus†
Tiniocellus spinipes
Trichius zonatus
Tropinota†
Silphidae
Nicrophorus†
Nicrophorus vespilloides
Phosphuga atrata
Silpha perforata
28s
AY132499.1
AY132454.1
DQ430857.1
AY131779.1
AY131780.1
AY131747.1
EU162481.1
AY131634.1
AJ810779.1
AY131781.1
AY132462.1
AY131684.1
AY131686.1
AY131788.1
AY131642.1
AY131791.1
AY131750.1
EU658919.1
AY131722.1
AY131725.1
16s
EF487800.1
EF487831.1
18s
EF521876.1
EF487638.1
EF570408.1
AY131595.1
AY131596.1
AY131560.1
EU162530.1
AY131443.1
AY131597.1
EF487792.1
AY131493.1
AJ810744.1
DQ012283.1
AY745576.1
AY821529.1
AY131495.1
AY131604.1
EF487979.1
AY131452.1
AY131607.1
HQ824537.1
HQ824533.1
HQ824538.1
HQ824535.1
HQ824534.1
AY131563.1
AF499692.1
AY131536.1
COI
AY132406.1
AY132377.1
ArgK
Wng
EU677503.1
EU677655.1
Elf1a
AY131938.1
AY131939.1
AY131916.1
EU162436.1
AY131814.1
EU084041.1
AY131940.1
AY132381.1
AY131860.1
X88946.1
AY131862.1
AY131945.1
AY821538.1
EF487656.1
AY131822.1
GQ443313.1
AF002809.1
AY821527.1
HQ824543.1
HQ824539.1
HQ824544.1
HQ824541.1
HQ824540.1
AY131918.1
AF499752.1
AY131893.1
AY745561.1
AY745566.1
AY131723.1
AY131706.1
AY131731.1
EU084190.1
AY131707.1
AY131534.1
AY131517.1
AY131543.1
DQ202690.1
DQ202616.1
EF487785.1
DQ430874.1
EU084231.1
EF188112.1
DQ524724.1
AY131741.1
AY131783.1
DQ430853.1
AY131518.1
EF487850.1
AY131554.1
AY131599.1
AY131521.1
DQ430887.1
EU162503.1
EU162506.1
EU162508.1
AY131774.1
DQ430885.1
GU226574.1
DQ430858.1
EU162556.1
EU162558.1
AY131590.1
EU162573.1
EU432271.1
EU084149.1
AY131608.1
EF487962.1
AY131733.1
AY131545.1
EF487818.1
DQ012277.1
EF487686.1
EF487584.1
EF487583.1
DQ430830.1
EF487702.1
EF188027.1
EF487721.1
AY821522.1
DQ430835.1
EU162548.1
DQ012282.1
EF487658.1
EU432226.1
DQ430855.1
EU432264.1
EU084152.1
EU084291.1
AY521744.1
EF487659.1
DQ430828.1
EF487944.1
EF487973.1
FJ000407.1
GU226581.1
EF656727.1
AY131799.1
DQ524754.1
EU084263.1
AY131807.1
EU432269.1
AY131743.1
EF487653.1
EF188042.1
EF487661.1
EF487722.1
AY521832.1
AY131891.1
AY131877.1
AY131900.1
EU084066.1
AY131878.1
EF487755.1
DQ222020.1
HM120756.1
DQ524498.1
AY131910.1
AY131942.1
AY131881.1
JF340440.1
EU162454.1
EU162457.1
EU162460.1
EU162462.1
AY131933.1
EU162476.1
AY847568.1
EF487735.1
EU477362.1
EF487733.1
EF188219.1
AY131902.1
EU084045.1
EU477298.1
EU477345.1
EU477347.1
EF487737.1
DQ295285.1
GQ457144.1
GQ457147.1
EU156615.1
AY745574.1
HQ630652.1
EF656685.1
AF499702.1
DQ681012.1
EF487872.1
AY131626.1
AY821540.1
EF487712.1
DQ012286.1
AY131556.1
EF188046.1
EF487655.1
AB285555.1
EF213789.1
DQ295300.1
EF656776.1
AY131957.1
DQ524521.1
EF487776.1
AY131965.1
EU477356.1
EF188226.1
EF487734.1
HQ599135.1
EU147356.1
EU147362.1
AB285573.1
AB285541.1
AB285534.1
365
EU147476.1
EU147490.1
AB376111.1
AB376168.1
AB285636.1
AB285629.1
Species
Thanatophilus†
Silvanidae
Oryzaephilus surinamensis
Staphylinidae
Aleochara†
Creophilus maxillosis villosus
Nudobius cephalus
Ontholestes†
Philonthus†
Staphylinus†
Tachinus†
Tenebrionidae
Alleculinae†
Arthromacra aenea
Blaps†
Diaperis boleti
Elenophorus†
Erodius orientalis
Gnatocerus cornutus
Gonocephalum†
Hegeter amaroides
Hegeter brevicollis
Hegeter costipennis
Hegeter fernandezi
Hegeter grancanariensis
Hegeter lateralis
Hegeter politus
Hegeter tenuipunctatus
Hegeter transversus
Latheticus oryzae
Opatroides†
Palorus†
Phaleria†
Pimelia ascendens
Pimelia atlantis atlantis
Pimelia atlantis frigioides
Pimelia baetica
Pimelia boyeri
Pimelia canariensis
Pimelia costata
Pimelia cribra
Pimelia echidna
Pimelia elevata
Pimelia estevezi
Pimelia fernandezlopezi
Pimelia granulicollis
Pimelia integra
Pimelia interjecta
Pimelia laevigata costipennis
Pimelia laevigata laevigata
Pimelia laevigata validipes
Pimelia lutaria
Pimelia maura
Pimelia mauritanica
Pimelia monticola
Pimelia radula oromii
Pimelia radula radula
Pimelia rugosa
Pimelia scabrosa
Pimelia sparsa albohumeralis
Pimelia sparsa serrimargo
Pimelia sparsa sparsa
Pimelia variolosa
Probaticus†
Stenosis†
Tenebrio molitor
Tentyria rotunda
Tentyria schaumi
Tribolium anaphe
Tribolium audax
Tribolium brevicornis
Tribolium castaneum
Tribolium confusum
Tribolium destructor
Tribolium freemani
Tribolium madens
Uloma†
Zophosis†
Trogidae
Trox borrei
Trogossitidae
Tenebroides†
Outgroup Taxa
Raphidioptera sp.
Sialis sp.
28s
AB285580.1
16s
AB285548.1
18s
EF213790.1
COI
DQ155789.1
ArgK
Wng
AB285643.1
Elf1a
FM877921.1
EF213812.1
EF213791.1
FJ763702.1
GU377335.1
GU377338.1
HM583885.1
EF213828.1
AY745632.1
GQ981067.1
AY663882.1
EU048306.1
EU048307.1
EU048290.1
AY663879.1
EF209961.1
EF209971.1
AY663861.1
EF209945.1
AF423769.1
AY663856.1
EU048291.1
AJ438152.1
AJ565942.1
AJ565956.1
AJ565953.1
AJ565961.1
AJ565954.1
AJ565943.1
AJ565964.1
AJ565959.1
AJ565970.1
AJ565960.1
AJ565948.1
AJ565949.1
AJ565947.1
AJ565963.1
AJ565958.1
AJ565950.1
AJ565951.1
AJ565952.1
AJ565939.1
AJ565957.1
AJ565955.1
AJ565965.1
AJ566048.1
AJ566062.1
AJ566059.1
AJ566067.1
AJ566060.1
AJ566049.1
AJ566070.1
AJ566065.1
AJ566076.1
AJ566066.1
AJ566054.1
AJ566055.1
AJ566053.1
AJ566069.1
AJ566064.1
AJ566056.1
AJ566057.1
AJ566058.1
AJ566045.1
AJ566063.1
AJ566061.1
AJ566071.1
AJ565941.1
AJ565969.1
AJ565966.1
AJ565945.1
AJ565946.1
AJ565944.1
AJ565962.1
AJ566047.1
AJ566075.1
AJ566072.1
AJ566051.1
AJ566052.1
AJ566050.1
AJ566068.1
X90683.1
FN392098.1
AJ565974.1
EU048302.1
EU048303.1
AJ438153.1
FN392041.1
AJ566080.1
AJ438150.1
AJ438151.1
AJ438149.1
AJ438143.1
AJ438145.1
AJ438147.1
AJ438141.1
AJ438140.1
FN392101.1
FN392044.1
EU048305.1
EU048301.1
EU048304.1
GU270008.1
FJ763716.1
GU377381.1
GU377384.1
GU377387.1
DQ155985.1
DQ155695.1
GU377483.1
GU377486.1
GU377489.1
FN544306.1
FM877930.1
FN544335.1
EU048282.1
FN544389.1
Z71729.1
Z71733.1
Z71739.1
Z71747.1
Z71741.1
Z71731.1
Z71743.1
Z71732.1
Z71734.1
EU048283.1
FM876597.1
AF139907.1
FN544674.1
AJ842966.1
AJ248202.1
AJ248199.1
AJ248206.1
AJ248200.1
AJ842969.1
AJ248209.1
EU048298.1
EU048299.1
AJ248215.1
AJ248205.1
X97211.1
X97212.1
AJ248208.1
AJ248204.1
AJ843063.1
AJ843037.1
AJ843010.1
X97216.1
AJ248203.1
AJ248201.1
AJ248210.1
AJ842971.1
AJ248198.1
AJ248214.1
AJ248211.1
X07801.1
HM156711.1
AJ248207.1
FN544690.1
AM947780.1
HQ891143.1
FM876314.1
Z71748.1
AJ438061.1
AJ438064.1
EU048281.1
AJ438075.1
AJ438079.1
AJ438067.1
AF139902.1
EU048279.1
EU048297.1
EU048293.1
EU048296.1
AY819656.1
EU048294.1
EU048295.1
EF209955.1
FN544775.1
EF487978.1
AF423774.1
EF487783.1
DQ202661.1
DQ202553.1
EF209680.1
FM877907.1
HM543340.1
AY521793.1
EU734902.1
EU815252.1
X89497.1
EU839719.1
GU013624.1
366
EU677513.1
EU797281.1
AY620203.1
HM156721.1
Appendix C
Carabidae trait data
367
Diploid number is the mean of all data available for each taxon included in the analysis. If there were conflicting records
the mean of all available records was used. Values for winged and wingless indicate probabilities. In cases where a
species was not represented the values used are calculated based on all data for the genus.
Taxon
Diploid Number
Winged
Wingless
27.9
0.84
0.16
ADCACA_Calosoma_inquistor
28
0.58
0.42
ADCACA_Carabus_auronitens
28.5
0.06
0.94
ADCACA_Carabus_blaptoides
28
0.06
0.94
ADCACA_Carabus_caelatus
28
0.06
0.94
27.5
0
1
ADCACA_Carabus_creutzeri
28
0.06
0.94
ADCACA_Carabus_fruhstorferi
28
0.06
0.94
ADCACA_Carabus_granulatus
22.5
0
1
28
0.06
0.94
ADCABR_Brachinus_sp
ADCACA_Carabus_cancellatus
ADCACA_Carabus_hispanus
ADCACA_Carabus_hortensis
28
0.06
0.94
ADCACA_Carabus_lineatus
28
0.06
0.94
ADCACA_Carabus_melancholicus
28
0.06
0.94
ADCACA_Carabus_monticola
28
0.06
0.94
ADCACA_Carabus_nemoralis
28.5
0
1
28
0.06
0.94
28.5
0.06
0.94
29
0.06
0.94
ADCACA_Carabus_problematicus
ADCACA_Carabus_punctatoauratus
ADCACA_Carabus_rutilans
ADCACA_Carabus_solieri
27
0.06
0.94
ADCACA_Carabus_splendens
29
0.06
0.94
ADCACA_Carabus_violaceus
28
0.06
0.94
ADCACA_Cychrus_caraboides
23
0
1
ADCAHA_Abax_parallelepipedus
35
0
1
ADCAHA_Agonum_muelleri
37
1
0
33.7
0.85
0.15
ADCAHA_Amphasia_sericea
29
1
0
ADCAHA_Anisodactylus_hispanus
37
0.97
0.03
ADCAHA_Apenes_sp
36
0.83
0.17
ADCAHA_Calathus_abaxoides
55
0.36
0.64
ADCAHA_Calathus_ambiguus
37
0.36
0.64
ADCAHA_Calathus_ascendens
23.5
0.36
0.64
ADCAHA_Calathus_asturiensis
39
0.36
0.64
ADCAHA_Calathus_auctus
37
0.36
0.64
ADCAHA_Calathus_brevis
37
0.36
0.64
ADCAHA_Calathus_circumseptus
43
0.36
0.64
ADCAHA_Calathus_erratus
37
0.36
0.64
ADCAHA_Amara_sp
368
ADCAHA_Calathus_freyi
36
0.36
0.64
ADCAHA_Calathus_fuscipes
37
0.5
0.5
ADCAHA_Calathus_granatensis
37
0.36
0.64
37.5
0
1
ADCAHA_Calathus_micropterus
37
0.36
0.64
ADCAHA_Calathus_mollis
39
0.36
0.64
ADCAHA_Calathus_melanocephalus
ADCAHA_Calathus_rectus
37
0.36
0.64
ADCAHA_Calathus_rotundatus
39
0.36
0.64
36.5
0.93
0.07
18
1
0
ADCAHA_Cymindis_sp
35.4
0.45
0.55
ADCAHA_Diplocheila_sp
38.9
1
0
ADCAHA_Dromius_sp
25
1
0
ADCAHA_Galerita_sp
30
0.8
0.2
ADCAHA_Harpalus_aesculanus
37
0.83
0.17
ADCAHA_Harpalus_affinis
37
1
0
ADCAHA_Harpalus_anxius
37
0.83
0.17
ADCAHA_Harpalus_attenuatus
37
0.83
0.17
ADCAHA_Chlaenius_sp
ADCAHA_Colliuris_pennsylvanica
ADCAHA_Harpalus_contemptus
37
0.83
0.17
ADCAHA_Harpalus_decipiens
37
0.83
0.17
ADCAHA_Harpalus_dimidiatus
35
0.83
0.17
ADCAHA_Harpalus_distinguendus
37
0.83
0.17
ADCAHA_Harpalus_ebeninus
37
0.83
0.17
ADCAHA_Harpalus_honestus
37
0.83
0.17
ADCAHA_Harpalus_pennsylvanicus
37
1
0
ADCAHA_Harpalus_rubripes
37
1
0
ADCAHA_Harpalus_serripes
38
0.83
0.17
ADCAHA_Harpalus_wagneri
30
0.83
0.17
ADCAHA_Lebia_sp
32
1
0
ADCAHA_Licinus_sp
30
1
0
ADCAHA_Loxandrus_sp
34.33
0.97
0.03
ADCAHA_Microlestes_sp
31
0.67
0.33
ADCAHA_Notiobia_sp
38.33
1
0
ADCAHA_Oodes_sp
25.33
1
0
ADCAHA_Ophonus_azureus
37
1
0
ADCAHA_Ophonus_longicollis
37
1
0
ADCAHA_Ophonus_stictus
37
1
0
ADCAHA_Poecilus_chalcites
37
1
0
ADCAHA_Poecilus_cupreus
43.5
0.72
0.28
27
0.17
0.83
ADCAHA_Poecilus_lucublandus
369
ADCAHA_Poecilus_versicolor
37
0.72
0.28
ADCAHA_Pterostichus_alacer
37
0.09
0.91
ADCAHA_Pterostichus_angustus
37
0
1
ADCAHA_Pterostichus_anthracinus
37
0.09
0.91
ADCAHA_Pterostichus_aterrimus
37
0.09
0.91
ADCAHA_Pterostichus_flavofemoratus
37
0.09
0.91
ADCAHA_Pterostichus_globosus
37
0.09
0.91
ADCAHA_Pterostichus_herculeanus
37
0
1
ADCAHA_Pterostichus_interruptus
37
0.09
0.91
ADCAHA_Pterostichus_melanarius
37
0.5
0.5
ADCAHA_Pterostichus_nemoralis
37
0.09
0.91
ADCAHA_Pterostichus_niger
37
0.09
0.91
ADCAHA_Pterostichus_nigrita
37
0.09
0.91
ADCAHA_Pterostichus_oblongopunctatus
28
0.09
0.91
ADCAHA_Pterostichus_spinolae
37
0.09
0.91
ADCAHA_Pterostichus_ziegleri
34
0.09
0.91
ADCAHA_Stenolophus_lecontei
25
1
0
ADCAHA_Stenolophus_mixtus
36
0.9
0.1
ADCAHA_Stenolophus_teutonus
24
0.9
0.1
ADCAHA_Stomis_sp
37
0
1
ADCALO_Loricera_pilicornis
19
1
0
ADCANE_Nebria_brevicollis
30
1
0
ADCANE_Notiophilus_sp
24
0.5
0.5
ADCAPS_Olisthopus_sp
41
1
0
ADCASC_Clivina_sp
36
0.83
0.17
37.67
0.4
0.6
ADCASC_Scarites_buparius
ADCASC_Scarites_eurytus
45
0.33
0.67
ADCASC_Scarites_hespericus
53
0.33
0.67
ADCASC_Scarites_laevigatus
61
0.33
0.67
ADCASC_Scarites_occidentalis_M
41
0.33
0.67
ADCASC_Scarites_terricola
57
0.33
0.67
ADCATR_Bembidion_aenulum
24
1
0
ADCATR_Bembidion_americanum
24
1
0
ADCATR_Bembidion_antiquum
24
0.89
0.11
ADCATR_Bembidion_arenobilis
24
0.89
0.11
ADCATR_Bembidion_articulatum
26
0.89
0.11
ADCATR_Bembidion_balli
24
1
0
ADCATR_Bembidion_bellorum
24
0.89
0.11
ADCATR_Bembidion_bifossulatum
24
1
0
ADCATR_Bembidion_californicum
24
1
0
370
ADCATR_Bembidion_chalceum
26
1
0
ADCATR_Bembidion_concolor
24
1
0
ADCATR_Bembidion_confusum
24
1
0
ADCATR_Bembidion_coxendix
23
1
0
ADCATR_Bembidion_decorum
24
0.89
0.11
ADCATR_Bembidion_honestum
24
1
0
ADCATR_Bembidion_inaequale
22
1
0
ADCATR_Bembidion_insulatum
24
1
0
ADCATR_Bembidion_integrum
24
0.89
0.11
ADCATR_Bembidion_lampros
23
0.5
0.5
ADCATR_Bembidion_lapponicum
24
1
0
ADCATR_Bembidion_levettei
22
1
0
ADCATR_Bembidion_levettei_carrianum
22
0.89
0.11
ADCATR_Bembidion_levigatum
24
1
0
ADCATR_Bembidion_louisella
24
0.89
0.11
ADCATR_Bembidion_mexicanum
24
1
0
ADCATR_Bembidion_obtusum
24
0.5
0.5
ADCATR_Bembidion_planatum
24
1
0
ADCATR_Bembidion_properans
24
1
0
ADCATR_Bembidion_punctulatum
24
0.89
0.11
23.3
0.5
0.5
24
1
0
ADCATR_Bembidion_quadrimaculatum
ADCATR_Bembidion_rapidum
ADCATR_Bembidion_rothfelsi
34
0.89
0.11
ADCATR_Bembidion_ruficorne
26
0.89
0.11
ADCATR_Bembidion_rufotinctum
24
1
0
ADCATR_Bembidion_sejunctum
24
0.5
0.5
ADCATR_Bembidion_tairuense
24
0.89
0.11
ADCATR_Bembidion_tetracolum
24.3
0.5
0.5
ADCATR_Bembidion_tibiale
24
0.89
0.11
ADCATR_Bembidion_transversale
24
1
0
ADCATR_Bembidion_umbratum
24
1
0
ADCATR_Pogonus_chalceus
22
1
0
ADCATR_Trechus_obtusus
23
0.5
0.5
ADCATR_Trechus_quadristriatus
.
23
1
0
371
Appendix D
Acari trait data
372
Species
Abacarus hystrix
Acaropsellina docta
Acarus siro
Aceria ficus
Aceria guerreronis
Aceria oleae
Aceria sheldoni
Achipteria coleoptrata
Achipteria punctata
Achipteria species
Acotyledon formosani
Aculops cornutus
Aculops fockeui
Aculops lycopersici
Aculops pelekassi
Aculops tetanothrix
Aculus persicae
Aculus schlechtendali
Aegyptobia ephedrae
Aegyptobia sp.
Aegyptobia thujae
Aegyptobia vannus
Afronothrus giganticus
Afronothrus incisivus
Afronothrus neotropicus
Afronothrus russeolus
Afronothrus schuilingi
Agistemus camerounensis
Agistemus exsertus
Agistemus sanctiluciae
Agistemus tranatalensis
Allonothrus giganticus
Allonothrus neotropicus
Allonothrus russeolus
Allonothrus schuilingi
Allosuctobelba obtusa
Allothrombium fuliginosum
Amblygamasus septentrionalis
Amblyomma americanum
Amblyomma cajennense
Amblyomma darwini
Amblyomma dissimile
Amblyomma helvolum
Amblyomma inornotum
Amblyomma limbatum
Amblyomma maculatum
Amblyomma moreliae
Amblyomma sp.
Amblyomma testudinarium
Amblyomma triguttatum
Amblyomma tuberculatum
Amblyseius aberrans
Amblyseius agrestris
Amblyseius barkeri
Amblyseius bibens
Amblyseius brevipes
Amblyseius chiapensis
Amblyseius chilensis
Amblyseius cucumeris
Amblyseius deleoni
Amblyseius deleoni.2
Amblyseius herbarius
Amblyseius hibisci
Reproductive
Mode
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
373
Female Diploid
Number
4
Ploidy
haplodiploidy
haplodiploidy
diplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
18
diplodiploidy
10
18
4
4
4
4
4
4
4
4
Sex
Chromosomes
XO
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
4
6
4
6
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
24
12
22
22
20
22
22
22
22
22
22
22
22
20
22
8
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
8
8
8
8
8
8
8
haplodiploidy
PGE
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
8
haplodiploidy
XO
XO
XY
XO
XO
XO
XY
XO
XY
XO
XO
XO
XO
Species
Amblyseius judaicus
Amblyseius largoensis
Amblyseius masiaka
Amblyseius messor
Amblyseius parasundi
Amblyseius rotundus
Amblyseius rubini
Amblyseius salish
Amblyseius sp.
Amblyseius swirskii
Amblyseius vazimba
Amnemochthonius taeniophorus
Anachipteria species
Anatetranychus tephrosiae
Androlaelaps casalis
Annectacarus mucronatus
Anoetus laboratorium
Antennophorus grandis
Anystis baccarum
Anystis salicinus
Apionoseius infirmus
Aponomma concolor
Aponomma fimbriatum
Aponomma hydrosauri
Aponomma undatum
Archegozetes longisetosus
Archegozets magnus
Archoplophora laevis
Archoplophora villosa
Areolaspis bifoliatus
Areolaspis sp.
Argas brumpti
Argas cooleyi
Argas hermanni
Argas japonicus
Argas persicus
Argas radiatus
Argas reflexus
Argas sanchezi
Argas tridentatus
Argas vespertilionis
Argas zumpti
Arrenurus bicuspidator
Arrenurus caudatus
Arrenurus maculata
Arrenurus pustulator
Arrenurus sp.1
Arrenurus sp.2
Arrenurus sp.3
Arrenurus sp.4
Arrenurus sp.5
Artacris macrorhynchus
Asca aelhiopica
Asca afroaphidioides
Asca aphidioides
Asca cranela
Asca evansi
Asca garmani
Asca muma
Asca piloja
Asca quinqueselosa
Atopochthonius artiodactylus
Atropacarus striculus
Reproductive
Mode
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
374
Female Diploid
Number
8
8
8
8
Ploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
8
8
haplodiploidy
haplodiploidy
8
8
8
haplodiploidy
haplodiploidy
haplodiploidy
6
14
haplodiploidy
haplodiploidy
8
24
haplodiploidy
haplodiploidy
20
22
18
20
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
10
haplodiploidy
haplodiploidy
24
26
26
26
26
26
26
26
26
26
26
20
26
20
23
20
26
4
Sex
Chromosomes
XO
XO
XO
XO
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
XY
XY
XY
XY
XY
XY
XY
XY
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
XY
Species
Austronothrus clarki
Austronothrus curviseta
Austronothrus flagellatus
Balaustium sp.
Banksinoma ovata
Bdella capitosa
Bdella tropica
Blattisocius patagiorum
Boophilus annulatus
Boophilus microplus
Brachychthonius berlesei
Brachychthonius pius
Brevipalpus californicus
Brevipalpus obovatus
Brevipalpus phoenicis
Brevipalpus pulcher
Brevipalpus russulus
Brevipalpus spinosus
Bryobia graminum
Bryobia kissophila
Bryobia lagodechiana
Bryobia latens
Bryobia neopraetiosa
Bryobia praetiosa
Bryobia rubrioculus
Bryobia sarcothamni
Caloglyphus berlesei
Caloglyphus michaeli
Caloglyphus mycophagus
Camisia aff. lapponica
Camisia carrolli
Camisia horrida
Camisia invenusta
Camisia segnis
Camisia spinifer
Carabodes femoralis
Carabodes granulatus
Cenopalpus lanceolatisetae
Ceratoppia bipilis
Ceratoppia sp.
Ceratozetes cf.
Ceratozetes cuspidatus
Ceratozetes gracilis
Ceratozetes parvulus
Cercoleipus coelonotus
Chamobates borealis
Chamobates voigtsi
Cheiroseius sp.
Cheletogenes ornatus
Cheyletus eruditus
Cheyletus malaccensis
Chrysomelobia labidomerae
Cilliba cassidea
Cilliba erlangensis
Cilliba minor
Cilliba sopronensis
Cilliba sp. 1
Cilliba sp. 2
Claveupodes delicatus
Clavidromus jackmickleyi
Cosmolaelaps gurabensis
Cosmolaelaps sp.1
Cosmolaelaps sp.2
Reproductive
Mode
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
375
Female Diploid
Number
Ploidy
Sex
Chromosomes
6
22
22
haplodiploidy
diplodiploidy
diplodiploidy
XO
XO
2
2
2
4
4
4
haplodiploidy
haplodiploidy
haplodiploidy
8
8
18
16
16
haplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
4
haplodiploidy
26
diplodiploidy
4
4
4
haplodiploidy
8
12
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
XO
XO
XO
Species
Cosmolaelaps vacua
Crotonia brachyrostrum
Crotonia caudalis
Cryptacarus promecus
Cultroribula bicultrata
Cultroribula divergens
Cunaxa capreolus
Cyrthermalllzia guadeloupensis
Cyrthermalllzia sp.1
Czenspinskia transversostriata
Damaeobelba minutissima
Damaeus angustipes
Damaeus verticillipes
Demodex caprae
Dermacentor albipictus
Dermacentor andersoni
Dermacentor hunteri
Dermacentor nitens
Dermacentor occidentalis
Dermacentor parumapertus
Dermacentor silvarum
Dermacentor sp.
Dermacentor sp.2
Dermacentor variabilis
Dermanyssus gallinae
Dermanyssus prognephilus
Dermanyssus prognephilus.2
Dermatophagoides farinae
Dicrocheles phalaenodectes
Discourella baloghi
Discourella modesta
Dolichotetranychus summers
Elliptochthonius profundus
Eniochthonius minutissimus
Ensliniella kostylevi
Ensliniella parasitica
Eobrachychthonius latior
Eohypochthonius magnus
Eohypochthonius travei
Eonychus curtisetosus
Eonychus grewiae
Eotetranychus befandrianae
Eotetranychus carpini
Eotetranychus friedmanni
Eotetranychus grandis
Eotetranychus imerinae
Eotetranychus paracybelus
Eotetranychus ranoma fanae
Eotetranychus rinoreae
Eotetranychus roedereri
Eotetranychus sakalavensis
Eotetranychus tiliarium
Eotetranychus tulearensis
Epidamaeus species
Epilohmannia cylindrica
Epilohmannia pallida
Epilohmannia styriaca
Epilohmannoides jacoti
Epilohmannoides terrae
Epitrimerus pyri
Eremobelba gracilior
Eriophyes laevis
Eriophyes tiliae
Reproductive
Mode
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
376
Female Diploid
Number
Ploidy
22
haplodiploidy
18
4
22
22
22
22
22
22
22
20
22
22
6
10
6
6
diplodiploidy
haplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
diplodiploidy
PGE
4
haplodiploidy
haplodiploidy
4
4
4
8
6
6
6
6
10
6
6
4
8
4
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
4
haplodiploidy
haplodiploidy
Sex
Chromosomes
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
Species
Erythraeus sp.
Euandrolaelaps sp.
Eugamasus kraepelini
Eugamasus magnus
Eulaelaps shanghaiensis
Eulohmannia ribagai
Eupalopsellus brevipilus
Eupalopsellus olearius
Eupelops hirtus
Eupelops plicatus
Eupelops torulosus
Eupodes sigmoidensis
Eupodes sp.
Eurytetranychus buxi
Eurytetranychus madagascariensis
Euseius hibisci
Euseius quetzali
Eutegaeus curviseta
Eutetranychus banksi
Eutetranychus eliei
Eutetranychus grandidieri
Eutetranychus orientalis
Eutetranychus ranjatori
Eutetranychus sambiranensis
Euzetes globulus
Evadorhagidia oblikensis
Eylais mutila
Eylais rimosa
Eylais setosa
Eylais sp.1
Eylais sp.2
Fosseremus laciniatus auct
Frontipoda musculus
Fuscouropoda appendiculata
Fuscozetes species
Gaeolaelaps aculeifer
Gaeolaelaps sp.1
Gaeolaelaps sp.2
Galumna ithacensis
Galumna sp.
Gamasellodes bicolor
Gamasellodes rectiventris
Gamasellodes sp.
Gamasellodes vermivorax
Gamasellus vibrissae
Gamasolaelaps whartoni
Geckobiella texana
Geholaspis alpinus
Geholaspis berlesei
Geholaspis longispinosus
Geholaspis longulus
Geholaspis mandibularis
Geholaspis pauperior
Gehypochthonius rhadamanthus
Gehypochthonius urticinus
Gehypochthonius xarifae
Geolaelaps oreithyiae
Glycyphagus domesticus
Glyptholaspis americana
Glyptholaspis confusa
Glyptholaspis fimicola
Glyptholaspis pontina
Gozmanyina majesta
Reproductive
Mode
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
parthenogenetic
377
Female Diploid
Number
12
10
16
Ploidy
diplodiploidy
haplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
8
6
haplodiploidy
haplodiploidy
18
10
6
haplodiploidy
haplodiploidy
haplodiploidy
PGE
PGE
6
8
4
6
6
4
18
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
diplodiploidy
6
4
4
4
6
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
19
diplodiploidy
18
haplodiploidy
haplodiploidy
haplodiploidy
18
diplodiploidy
16
Sex
Chromosomes
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
18
diplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
XO
Species
Gozmanyina majestus
Graptoppia (Stenoppia)
Gymnodamaeus bicostatus
Haemaphysalis bancrofti
Haemaphysalis bispinosa
Haemaphysalis bremneri
Haemaphysalis campanulata
Haemaphysalis flava
Haemaphysalis formosensis
Haemaphysalis hystricis
Haemaphysalis japonica
Haemaphysalis kitaokai
Haemaphysalis lagrangei
Haemaphysalis leachi
Haemaphysalis leporispalustris
Haemaphysalis longicornis
Haemaphysalis megaspinosa
Haemaphysalis pentalagi
Haemogamasus centrocarpus
Haemogamasus longipes
Halotydeus destructor
Halotydeus destructor.2
Haplochthonius simplex
Harpyrhynchus brevis
Harpyrhynchus novoplumaris
Hawaiieupodes thermophilus
Heminothrus interlamellaris
Heminothrus longisetosus
Heminothrus ornatissimus
Heminothrus paolianus
Heminothrus targionii
Hemisarcoptes coccophagus
Hermannia gibba
Hermanniella species
Hexanoetus conoidalis
Histiostoma bakeri
Histiostoma feroniarum
Histiostoma formosana
Histiostoma humiditatis
Histiostoma julorum
Histiostoma murchei
Holonothrus sp.
Holostaspella sp.
Homeopronematus anconai
Hormosianoetus laboratorium
Humerobates rostrolamellatus
Hyalomma aegyptium
Hyalomma anatolicum
Hyalomma anatolicum excavatum
Hyalomma asiaticum
Hyalomma asiaticum excavatum
Hyalomma detritum
Hyalomma dromedarii
Hyalomma franchinii
Hyalomma impeltarum
Hyalomma marginatum
Hyalomma rhipicephaloides
Hyalomma rufipes
Hydrachna globosa
Hydrachna leegei
Hydrachna sp.1
Hydrachna sp.2
Hydrachna uniscutata
Reproductive
Mode
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
378
Female Diploid
Number
22
22
22
22
22
20
22
20
22
16
22
29
22
22
Ploidy
Sex
Chromosomes
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
XO
XO
XO
XO
XO
XO
diplodiploidy
diplodiploidy
XO
XO
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
haplodiploidy
diplodiploidy
XO
XO
XO
XO
4
4
haplodiploidy
haplodiploidy
14
16
diplodiploidy
diplodiploidy
haplodiploidy
14
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
8
16
22
22
22
22
22
22
22
22
22
22
22
22
12
20
12
12
12
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
XO
Species
Hydrodroma despiciens
Hydrodroma despiciens.2
Hydrozetes dimorphus
Hydrozetes lacustris
Hydrozetes parisiensis
Hydrozetes terrestris
Hydrozetes tridactylus
Hydryphantes bayeri
Hydryphantes clypeatus
Hydryphantes ruber
Hydryphantes sp.
Hydryphantes sp.1
Hydryphantes sp.2
Hygrobates calliger
Hypoaspis lubrica
Hypochthonius luteus
Hypochthonius rufulus
Imparipes histricinus
Indotritia acanthophora
Iphidinychus geieri
Iphiduropoda penicillata
Iphiseius degenerans
Iponemus confusus
Iponemus radiatae
Ixodes cornuatus
Ixodes hexagonus
Ixodes holocyclus
Ixodes kingi
Ixodes laysanensis
Ixodes nipponensis
Ixodes ricinus
Ixodes tasmani
Jacotella species
Kennethiella trisetosa
Kurosaia jiju
Labidostomma luteum
Laelaspis sp.
Lasioseius berlesi
Lasioseius denlalus
Lasioseius subterraneus
Lasioseius youcefi
Lebertia porosa
Lebertia sp.1
Lebertia sp.2
Lebertia stackelbergi
Leiodinychus orbicularis
Leipothrix dipsacivagus
Leitneria granulata
Lepidozetes singularis
Leptotrombidium akamushi
Leptotrombidium arenicola
Leptotrombidium deliense
Leptotrombidium fletcheri
Leptotrombidium scutellare
Liacarus coracinus
Liacarus subterraneus
Limnesia maculata
Limnesia sp.1
Limnesia sp.2
Limnesia undulata
Limnochares aquatica
Limnozetes amnicus
Limnozetes atmetos
Reproductive
Mode
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
379
Female Diploid
Number
6
16
Ploidy
diplodiploidy
diplodiploidy
10
6
12
10
10
10
14
14
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
18
diplodiploidy
haplodiploidy
8
haplodiploidy
haplodiploidy
haplodiploidy
24
26
24
26
28
28
28
24
Sex
Chromosomes
XO
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
XO
XO
XY
XY
diplodiploidy
XY
haplodiploidy
haplodiploidy
haplodiploidy
16
16
18
18
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
12
28
14
14
14
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
18
18
18
18
6
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
XY
XY
Species
Limnozetes borealis
Limnozetes guyi
Limnozetes latilamellata
Limnozetes lustrum
Limnozetes onondaga
Limnozetes palmerae
Limnozetes sphagni
Linopodes sp.
Liochthonius brevis
Liochthonius sellnicki
Liochthonius strenzkei
Lohmannia banksi
Lohmannia lanceolata
Macrocheles boudreauxi
Macrocheles glaber
Macrocheles insignitus
Macrocheles lerreus
Macrocheles mammifer
Macrocheles matrius
Macrocheles merdarius
Macrocheles muscadomesticae
Macrocheles parapisentii
Macrocheles penicilliger
Macrocheles peniculalus
Macrocheles peregrinus
Macrocheles perglaber
Macrocheles pisentii
Macrocheles robustulus
Macrocheles rodriguezi
Macrocheles schaeferi
Macrocheles scutatus
Macrocheles similis
Macrocheles sp.
Macrocheles subbadius
Macrocheles vernalis
Mainothrus badius
Malacoangelia remigera
Malaconothrus crassisetosa
Malaconothrus gracilis
Malaconothrus hauseri
Malaconothrus robustus
Malaconothrus robustus asiaticus
Malaconothrus sp.1
Malaconothrus sp.2
Masthermannia sp.1
Meristacarus sp.
Metaseiulus occidentalis
Microppia minus
Microtritia minima
Mucronothrus nasalis
Multioppia species
Myialges pari
Myianoetus sp.
Nanhermannia comitalis
Nanhermannia coronata
Nanhermannia dorsalis
Nanhermannia elegantula
Nanhermannia nana
Nehypochthonius porosus
Nenteria breviunguiculata
Nenteria stylifera
Neodiscopoma pulcherrima
Neodiscopoma splendida
Reproductive
Mode
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
sexual
380
Female Diploid
Number
Ploidy
18
haplodiploidy
10
haplodiploidy
haplodiploidy
haplodiploidy
10
10
10
6
16
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
PGE
haplodiploidy
haplodiploidy
Sex
Chromosomes
Species
Neognathus terrestris
Neophyllobius aesculi
Neophyllobius elegans
Neophyllobius piniphilus
Neoseiulus californicus
Neoseiulus setulus
Neotetranychus rubi
Neotydeus ardisannae
Neumania vemalis
Neumania vernalis
Nipponiella sp
Nodele simplex
Nothrus anauniensis
Nothrus borussicus
Nothrus macedi
Nothrus monticolus
Nothrus palustris
Nothrus pratensis
Nothrus quadripilis
Nothrus silvestris
Nothrus silvicus
Nothrus terminalis carolinae
Nothrus truncatus
Novonothrus flagellatus
Obuloides sp.
Oligomerismus oregonensis
Oligonychus andrei
Oligonychus bessardi
Oligonychus chazeaui
Oligonychus coffeae
Oligonychus gossypii
Oligonychus monsarrati
Oligonychus pratensis
Oligonychus quercinus
Oligonychus randriamasii
Oligonychus sylvestris
Oligonychus thelytokous
Oligonychus ununguis
Oligonychus virens
Olodiscus minimus
Oodinychus karawaiewi
Oodinychus obscurasimilis
Oodinychus ovalis
Oodinychus spatulifera
Ophionyssus natricis
Oplitis alophora
Oplitis franzi
Oplitis wasmanni
Oppia cf.
Oppia nitens
Oppia nodosa
Oppia sp.
Oppiella nova
Oppioid sp.
Oribatella calcarata
Oribatella quadricornuta
Oribatella sakamorii
Oribatella sp.1
Oribatella sp.2
Oribatula sakamorii
Oribatula tibialis
Ornithodoros alactogalis
Ornithodoros asperus
Reproductive
Mode
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
sexual
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
381
Female Diploid
Number
Ploidy
22
22
20
haplodiploidy
haplodiploidy
haplodiploidy
PGE
14
haplodiploidy
4
4
diplodiploidy
diplodiploidy
4
haplodiploidy
6
haplodiploidy
4
8
8
6
4
8
8
6
4
4
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
6
8
haplodiploidy
haplodiploidy
18
haplodiploidy
18
haplodiploidy
33
16
diplodiploidy
Sex
Chromosomes
XY
Species
Ornithodoros capensis
Ornithodoros gurneyi
Ornithodoros lahorensis
Ornithodoros macmillani
Ornithodoros moubata
Ornithodoros nereensis
Ornithodoros savignyi
Ornithodoros tartakovskyi
Ornithodoros tholozani
Ornithonyssus bacoti
Ornithonyssus sylviarum
Orthogalumna terebrantis
Otobius lagophilus
Otobius megnini
Palaeacarus hystricinus
Palaeacarus kamenskii
Panonychus ulmi
Paragignathus tamaricis
Paralycus lavoipierrei
Paralycus parvulus
Parathyas dirempta
Parhypochthonius aphidinus
Pediculaster mesembrinae
Pelelhiphis berlesei
Pelelhiphis insignis
Pelelhiphis rufeseens
Peloribates cf.
Penthaleus major
Pergalumna curva
Pergalumna emarginata
Pergalumna formiparthenogenesis
(obligate)ria
Pergamasus brevicornis
Petrobia harti
Petrobia latens
Phaulodiaspis borealis
Phaulodiaspis rackei
Phorytocarpais hyalinus
Phthiracarus compressus
Phthiracarus setosellus
Phyllochthonius aoutii
Phyllocoptruta oleivora
Phytonemus pallidus
Phytoptus tiliae
Phytoseiulus persimilis
Phytoseius amba
Phytoseius finitimus
Phytoseius sp.1
Phytoseius sp.2
Piona camea
Piona coccinea coccinea
Piona nodata
Piona sp.1
Piona sp.2
Piona sp.3
Piona sp.4
Piona uncata uncata
Plarynothrus altimontanus
Plarynothrus banksi
Plarynothrus biangulatus
Plarynothrus bicarinatus
Plarynothrus brevisetosus
Plarynothrus carinatus
Reproductive
Mode
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
382
Female Diploid
Number
20
12
26
16
20
25
20
21
16
16
18
18
20
20
Ploidy
diplodiploidy
diplodiploidy
Sex
Chromosomes
XY
XY
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
XY
XY
XY
XY
haplodiploidy
haplodiploidy
haplodiploidy
6
8
haplodiploidy
haplodiploidy
18
diplodiploidy
6
haplodiploidy
12
4
8
diplodiploidy
haplodiploidy
haplodiploidy
4
4
4
8
8
8
8
8
22
20
8
8
20
22
haplodiploidy
haplodiploidy
haplodiploidy
PGE
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
20
Species
Plarynothrus castaneus
Plarynothrus major
Plarynothrus microclava
Platynothrus peltifer
Platynothrus punctatus
Platynothrus septentrionalis
Platynothrus sibiricus
Platynothrus skottsbergii
Platynothrus thori
Platynothrus traversus
Podocinum pacificum
Podocinum sagax
Podopterotegaeus tectus
Podoribates pratensis
Poecilochthonius spiciger
Poecilophysis faeroensis
Poecilophysis pratensis
Polyaspinus cylindricus
Polyaspinus patavinus
Polyaspis sansonei
Polyphagotarsonemus latus
Pomerantzia benhami
Pomerantzia kethleyi
Pomerantzia prolata
Porcupinychus insularis
Poroliodes farinosus
Proctolaelaps krimsei
Proctolaelaps longipilis
Protogamasellus brevicornis
Protogamasellus hibernicus
Protogamasellus massula
Protogamasellus mica
Protogamasellus sp.
Protokalumma salicis
Protoribates capucinus
Protoribates lophotrichus
Pseudonothrus hirtus
Pseudoparasitus sp.
Pseudouropoda calcarata
Pseudouropoda structura
Pseudouropoda tuberosa
Pterochthonius angelus
Punctoribates insignis
Punctoribates punctum
Pyemotes herfsi
Pyemotes scolyti
Pyemotes tritici
Pyemotes ventricosus
Quadroppia circumita
Quadroppia quadricarinata
Quadroppia sp.1
Quadroppia sp.2
Quadroppia sp.3
Raoiella indica
Rhagidia diversicolor
Rhagidia gigas
Rhinoseius colwelli
Rhipicephalus bursa
Rhipicephalus evertsi
Rhipicephalus sanguineus
Rhipicephalus secundus
Rhizoglyphus echinopus
Rhizoglyphus robini
Reproductive
Mode
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
383
Female Diploid
Number
Ploidy
18
diplodiploidy
10
10
haplodiploidy
4
haplodiploidy
8
18
haplodiploidy
diplodiploidy
haplodiploidy
haplodiploidy
Sex
Chromosomes
haplodiploidy
6
6
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
4
haplodiploidy
24
22
22
22
10
haplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
XO
XO
XO
XY
Species
Rhodacarellus silesiacus
Rhodacarus denticulatus
Rhyncaphytoptus ficifoliae
Rhysotritia ardua
Rhysotritia duplicata
Riccardoella limacum
Robustocheles mucronata
Rostrozetes foveolatus
Sancassania berlesei
Sancassania michaeli
Sancassania mycophaga
Saniosulus nudus
Sapracarus sp.
Sapracarus tuberculalUs
Sarcoptes scabiei
Scheloribates laevigatus
Scheloribates lanceoliger
Scheloribates species
Schizonobia oudemansi
Schizonobia sycophanta
Schizotetranychus australis
Schizotetranychus schizopus
Schwiebea elongata
Scutacaridae flechtmanni
Seiulus isolrichus
Sellnickochthonius immaculatus
Sellnickochthonius lydiae
Sellnickochthonius suecia
Sellnickochthonius zelawaiensis
Sericothrombium schar/atinum
Sericothrombium sp.1
Sericothrombium sp.2
Sericothrombium sp.3
Shibaia longisensilla
Siteroptes graminum
Siteroptes reniformis
Steganacarus magnus
Stigmocheylus sp.1
Stigmocheylus sp.2
Stratiolaelaps miles
Suctobelbella falcata
Suctobelbella hamata
Suctobelbella hurshi
Suctobelbella laevis
Suctobelbella palustris
Suctobelbella similis
Suctobelbella sp.1
Suctobelbella sp.10
Suctobelbella sp.11
Suctobelbella sp.12
Suctobelbella sp.13
Suctobelbella sp.14
Suctobelbella sp.15
Suctobelbella sp.16
Suctobelbella sp.17
Suctobelbella sp.18
Suctobelbella sp.19
Suctobelbella sp.2
Suctobelbella sp.20
Suctobelbella sp.21
Suctobelbella sp.3
Suctobelbella sp.4
Suctobelbella sp.5
Reproductive
Mode
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
parthenogenetic
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
384
Female Diploid
Number
Ploidy
Sex
Chromosomes
haplodiploidy
10
haplodiploidy
18
16
16
6
diplodiploidy
diplodiploidy
diplodiploidy
haplodiploidy
XO
XO
XO
18
diplodiploidy
XO
8
8
12
6
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
6
8
haplodiploidy
haplodiploidy
18
26
22
18
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
6
6
haplodiploidy
haplodiploidy
14
haplodiploidy
Species
Suctobelbella sp.6
Suctobelbella sp.7
Suctobelbella sp.8
Suctobelbella sp.9
Suctobelbella subcornigera
Suctobelbella tuberculata
Suctobelbella vera
Synchthonius crenulatus
Syringophiloidus minor
Tarsonemus confusus
Tarsonemus confusus.2
Tarsonemus lobosus
Tarsonemus nodosus
Tarsonemus randsi
Tarsonemus schlechtendali
Tarsonemus sp.
Tarsonemus talpae
Tarsonemus virgineus
Tarsonemus waitei
Tectocepheus cuspidatus
Tectocepheus minor
Tectocepheus sarekensis
Tectocepheus velatus
Tegolophus hassani
Tenuipalpoides acaciae
Tenuipalpus inophylli
Tetranychus atlanticus
Tetranychus cinnabarinus
Tetranychus hydrangeae
Tetranychus kaliphorae
Tetranychus ludeni
Tetranychus neocalendonicus
Tetranychus pacificus
Tetranychus panici
Tetranychus roseus
Tetranychus tumidus
Tetranychus urticae
Tetranychus viennensis
Tetranycopsis horridus
Torpacarus omittens
Trachytes aegrota
Trachytes irenae
Trachytes lamda
Trachytes minima
Trachytes montana
Trachytes pauperior
Trachytes pi
Trachyuropoda coccinea
Trachyuropoda pulchella
Trachyuropoda pyriformis
Trematurella elegans
Trhypochthoniellus badius
Trhypochthoniellus crassus
Trhypochthoniellus excavatus
Trhypochthoniellus setosus
Trhypochthoniellus sp.1
Trhypochthoniellus sp.2
Trhypochthonius americanus
Trhypochthonius nigricans
Trhypochthonius silvestris
Trhypochthonius tectorum
Trichouropoda schweizeri
Trichouropoda sociata
Reproductive
Mode
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
parthenogenetic
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
385
Female Diploid
Number
Ploidy
6
4
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
6
haplodiploidy
haplodiploidy
6
6
6
6
6
6
6
8
8
12
6
6
4
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
18
Sex
Chromosomes
Species
Trimalaconothrus glaber
Trimalaconothrus novus
Trimalaconothrus saxosus
Trimalaconothrus simplex
Troglocoptes sp.1
Troglocoptes sp.2
Troglocoptes sp.3
Troglocoptes sp.4
Tydeus caudatus
Typhlodromus athiasae
Typhlodromus caudiglans
Typhlodromus chazeaui
Typhlodromus contiguus
Typhlodromus drori
Typhlodromus fallacis
Typhlodromus guatemalensis
Typhlodromus gutierrezi
Typhlodromus phialatus
Typhlodromus porathi
Typhlodromus pyri
Typhlodromus rhenanus
Typhlodromus sp.
Typhlodromus stemlichti
Typhlodromus transvaalensis
Tyrophagus casei
Tyrophagus neiswanderi
Tyrophagus palmarum
Tyrophagus putrescentiae
Unionicola crassipes
Urodiaspis pannonica
Urodiaspis stammeri
Urodiaspis tecta
Uroobovella advena
Uroobovella arcuatus
Uroobovella cordieri
Uroobovella fracta
Uroobovella inermis
Uroobovella obovata
Uroobovella perforatus
Uroobovella woelkei
Uroplitella conspicua
Uroplitella paradoxa
Uropoda hamulifera
Uropoda italica
Uropoda kargi
Uropoda minima
Uropoda misella
Uropoda orbicularis
Uropoda undulata
Uroseius hunzikeri
Urotrachytes formicarius
Varroa destructor
Varroa jacobsoni
Vasates aceriscrumena
Veigaia cerva
Veigaia exigua
Veigaia kochi
Veigaia nemorensis
Veigaia parlita
Veigaia planieola
Veigaia pusilla
Veigaia tranisalae
Veigaia uneata
Reproductive
Mode
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
sexual
sexual
parthenogenetic
sexual
parthenogenetic
parthenogenetic
parthenogenetic
sexual
sexual
sexual
sexual
sexual
sexual
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
parthenogenetic
386
Female Diploid
Number
Ploidy
4
8
8
8
8
8
8
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
haplodiploidy
8
8
8
8
8
8
haplodiploidy
haplodiploidy
haplodiploidy
PGE
haplodiploidy
haplodiploidy
haplodiploidy
10
12
16
16
18
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
diplodiploidy
14
PGE
PGE
haplodiploidy
Sex
Chromosomes
XY
XY
XO
XO
Species
Xenillus tegeocranus
Xylobates capucinus
Xylobates oblongus
Xylobates robuSlior
Reproductive
Mode
sexual
parthenogenetic
parthenogenetic
parthenogenetic
.
387
Female Diploid
Number
18
Ploidy
diplodiploidy
Sex
Chromosomes
Appendix E
Hymenoptera trait data
388
Species
Abia candens
Acanthomyrmex ferox
Acanthomyrmex notabilis
Acanthomyrmex sp1
Acanthomyrmex sp2
Acanthomyrmex sp3
Acanthomyrmex sp4
Acromyrmex ambiguus
Acromyrmex coronatus
Acromyrmex crassipinus
Acromyrmex echinatior
Acromyrmex heyeri
Acromyrmex hispidus
Acromyrmex insinuator
Acromyrmex landolti
Acromyrmex octospinosus
Acromyrmex subterraneus
Acromyrmex versicolor
Acropyga acutiventris
Acropyga keira
Acropyga smithii
Acropyga sp1
Acropyga sp2
Acropyga sp3
Acropyga stygia
Adelomyrmex biroi
Aenictus brevicornis
Aenictus currax
Aenictus laeviceps
Aenictus sp1
Aethercerus discolor
Aethercerus dispar
Aethercerus nitidus
Aethercerus ranini
Agapostemon splendens
Agapostemon virescens
Ageniaspis fuscicollis
Aglaostigma albicincta
Aglaostigma amoorensis
Aglaostigma aucupariae
Aglaostigma nebulosa
Aglaostigma occipitosa
Aglaostigma sapporonis
Aglaostigma sp
Agrothereutes extrematus
Allantus luctifer
Allantus meridionalis
Allantus nakabusensis
Allantus sp
Alphostromboceros konowi
Amblyopone australis
Amblyopone fortis
Amblyopone pallipes
Amblyopone pluto
Amblyopone reclinata
Amblyopone sp1
Ametastegia geranii
Ametastegia pallipes
Ammophila pictipennis
Anagyrus lopezi
Anastatus catalonicus
Ancistrocerus adiabatuscytainus
Ancistrocerus densepilloserus
Ancistrocerus simulator
Ancistrocerus spilogaster
Ancistrocerus tuberculicepssutterianus
Andrena duboisi
Andrena dunningi
Andrena sp
Andrena togashii
Andricus curvator
Andricus fecundator
Andricus kollari
Haploid
Number
16.0
12.0
11.0
11.0
11.0
19.0
19.0
18.0
19.0
19.0
19.0
15.0
16.0
15.0
12.0
11.0
15.0
11.0
12.0
11.0
11.0
17.0
10.0
15.0
16.0
12.0
10.0
17.0
9.0
22.0
10.0
8.0
9.0
10.0
8.0
7.0
24.0
22.0
19.0
9.0
6.0
10.0
5.0
6.0
6.0
7.0
6.0
10.0
3.0
10.0
3.0
10.0
10.0
10.0
Eusocial
Colony Size
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
57
40
10000
1000
1000
3000
1000
92
10
150000
97500
Parasite
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Polyandry
Polygyny
Low
Relatedness
0
0
1
1
1
2
2
0
1
1
1
2
1
1
0
1
0
1
0
1
1
0
1
0
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
78
96
22.5
36.5
82
89
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
389
0
0
0
0
0
0
Species
Andricus quercuscalicis
Aneugmenus japonicus
Aneugmenus kiotonis
Aneugmenus stramineipes
Aneuretus simoni
Anisopteromalus calandrae
Anochetus altisquamis
Anochetus bequaerti
Anochetus diegensis
Anochetus faurei
Anochetus graeffei
Anochetus graeffei2
Anochetus horridus
Anochetus katonae
Anochetus madaraszi
Anochetus modicus
Anochetus sp1
Anochetus sp2
Anochetus sp3
Anochetus sp4
Anochetus sp5
Anochetus sp6
Anochetus yerburyi
Anonychomyrma itinerans
Anonychomyrma scrutator
Anonychomyrma sp1
Anoplius concinnus
Anoplius viaticus
Anoplolepis custodiens
Anoplolepis gracilipes
Anoplonyx sp
Anthidiellum notatum rufimaculatum
Anthidium mormonum
Anthophora acervorumvillosula
Anthophora bomboides
Anthophora californica
Anthophora plumipes
Aoplus pulchricornis
Apanteles sp1
Aphaenogaster albisetosus
Aphaenogaster araneoides
Aphaenogaster ashmeadi
Aphaenogaster beccarii
Aphaenogaster beccarii2
Aphaenogaster carolinensis
Aphaenogaster cockerelli
Aphaenogaster depilis
Aphaenogaster dromedaria
Aphaenogaster dulcinaea
Aphaenogaster famelica
Aphaenogaster flemingi
Aphaenogaster floridana
Aphaenogaster fulva
Aphaenogaster gibbosa
Aphaenogaster iberica
Aphaenogaster lamellidens
Aphaenogaster longiceps
Aphaenogaster miamiana
Aphaenogaster osimensis
Aphaenogaster phalangium
Aphaenogaster rudis
Aphaenogaster sardoa
Aphaenogaster senilis
Aphaenogaster smythiesi
Aphaenogaster sp1
Aphaenogaster spinosa
Aphaenogaster subterranea
Aphaenogaster testaceopilosa
Aphaenogaster tipuna
Aphaenogaster treatae
Aphelinus abdominalis
Aphelinus mali
Aphidius colemani
Haploid
Number
10.0
7.0
7.0
8.0
7.0
15.0
15.0
19.0
23.0
14.0
15.0
12.0
19.0
15.0
15.0
17.0
17.0
15.0
8.0
8.0
14.0
14.0
17.0
8.0
16.0
9.0
18.0
19.0
9.0
13.0
11.0
15.0
23.0
17.0
17.0
18.0
14.7
17.0
19.0
23.0
18.0
16.0
19.4
17.0
16.0
14.0
15.0
11.0
17.0
17.0
21.0
5.0
Eusocial
Colony Size
0
0
0
0
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
40
1
84
13
434
84
1750
Parasite
Polyandry
Polygyny
Low
Relatedness
1
1
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
15000
1
1
1
1
1
1
1
1
120
300
6000
100
300
50
281
300
120
797
3906
881.33
1
390
0
Species
Aphidius ervi
Aphidius rhopalosiphi
Aphytis mytilaspidus
Apis andreniformis
Apis cerana
Apis ceranaindica
Apis ceranajaponica
Apis dorsata
Apis florea
Apis mellifera
Apis mellifera adansonii
Apis mellifera ligustica
Apis mellifera mellifera
Apomyrma stygia
Apterostigma angulatum
Apterostigma collare
Apterostigma dentigerum
Apterostigma GR
Apterostigma mayri
Apterostigma pilosum
Apterostigma robustum
Apterostigma SH
Apterostigma sp1
Apterostigma sp2
Apterostigma sp3
Apterostigma SQ
Aptesis puncticollis
Arge cyanocrocea
Arge gracilicornis
Arge melanochroa
Arge nigripes
Arge pagana
Arge pectoralis
Arge ustulata
Ashmeadiella sp
Asiemphytus albilabris
Athalia bicolor
Athalia cordata
Athalia japonica
Athalia kashmirensis
Athalia lugensinfumata
Athalia rosaerosae
Athalia rosaeruficornis
Atopomyrmex mocquerysi
Atta bisphaerica
Atta cephalotes
Atta columbica
Atta laevigata
Atta sexdens
Atta texana
Augochlora pura
Augochlorella michaelis
Augochlorella persimilis
Augochlorella pura
Augochlorella striata
Augochloropsis brachycephala
Augochloropsis metallica
Augochloropsis sparsilis
Aulacidea hieracii
Austroplebeia australis
Austroplebeia symei
Axestotrigona ferruginea
Baeosemus dentifer
Baranisobas ridibundus
Basiceros manni
Belyta depressa
Biorhiza pallida
Biosteres carbonarius
Birka carinifrons
Bombus affinis
Bombus atratus
Bombus auricomus
Bombus bifarius
Haploid
Number
7.0
5.0
16.0
16.0
16.0
16.0
16.0
16.0
16.0
16.0
16.0
12.0
10.0
12.0
16.0
8.0
11.0
8.0
10.0
13.0
8.0
8.0
8.0
16.0
15.0
6.0
6.0
8.0
6.0
8.0
8.0
8.0
11.0
22.0
11.0
11.0
11.0
11.0
16.0
8.0
10.0
18.0
11.0
11.0
8.0
10.0
14.0
7.0
18.0
20.0
Eusocial
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
0
1
1
1
1
0
0
0
0
1
1
1
0
0
1
0
0
0
0
1
1
1
1
Colony Size
Parasite
Polyandry
Polygyny
Low
Relatedness
6884
2
2
0
1
1
36630
6271
60,000
2
2
2
0
0
0
1
1
1
1
1
0
0
0
0
0
0
0
1
0
0
1
0
2
2
2
2
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
52.5
155
100
155
27
45
46
41
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
65000
1000000
1420000
5000000
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
3.5
3.5
3.5
1
1
1
2000
2000
1
1
34.5
1
1
1
1
176
391
0
Species
Bombus bimaculatus
Bombus californicus
Bombus citrinus
Bombus consobrinuswittenburghi
Bombus deuteronymusmaruhanabachi
Bombus diversus
Bombus honshuensis
Bombus hortorum
Bombus hypnorum
Bombus hypocrita
Bombus ignitus
Bombus impatiens
Bombus lucorum
Bombus medius
Bombus moderatus
Bombus morio
Bombus muscorum
Bombus nevadensis
Bombus pennsylvanicus
Bombus pseudobaicalensis
Bombus ruderatus
Bombus schrencki
Bombus terrestris
Bombus terricola
Bombus terricolaoccidentalis
Bombus ussurensis
Bothriomyrmex gibbus
Bothriomyrmex hispanicus
Bothriomyrmex meridionalis
Bothriomyrmex pusillus
Bothriomyrmex sp1
Brachygastra augusti
Brachygastra bilineolata
Brachygastra lecheguana
Brachygastra mellifica
Brachygastra moebiana
Brachygastra scutellaris
Brachymeria intermedia
Brachymeria lasus
Brachymeria ovata
Brachymyrmex depilis
Brachymyrmex heeri
Brachymyrmex pictusbalboae
Brachymyrmex sp1
Brachymyrmex sp2
Brachymyrmex spnov
Brachymyrmex spnrobscurior
Calameuta filiformis
Calomyrmex laevissimus
Calomyrmex spANIC-1
Camargoia nordestina
Camponotus abscisus
Camponotus aethiops
Camponotus albicoxis
Camponotus alii
Camponotus americanus
Camponotus atriceps
Camponotus balzani
Camponotus bonariensis
Camponotus brevis
Camponotus caryae
Camponotus castaneus
Camponotus chromaiodes
Camponotus cingulatus
Camponotus claviscapus
Camponotus compressus
Camponotus confusus
Camponotus consobrinus
Camponotus crassisquamis
Camponotus crassus
Camponotus cruentatus
Camponotus discolor
Camponotus dolendus
Haploid
Number
18.0
19.0
23.0
18.0
17.0
18.0
18.0
18.0
18.0
20.0
17.5
18.0
17.0
17.0
18.0
18.0
18.0
18.0
11.0
11.0
11.0
28.0
3.0
5.0
5.0
9.0
20.0
14.0
17.0
21.0
21.0
20.0
20.0
20.0
20.0
20.0
15.0
23.0
20.0
10.0
19.0
10.0
Eusocial
Colony Size
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Parasite
Polyandry
Polygyny
60
1
1
1
0
0
0
Low
Relatedness
1
1
1
100
28.5
0
1
0
0
0
1
200
2184
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
50
139
400
100
0
0
0
0
0
1,000
900
7951
400
600
1
1
1
110
125
200
200
1
250
3560
350
3560
200
70
392
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Species
Camponotus excisus
Camponotus femoratus
Camponotus ferrugeneus
Camponotus festinus
Camponotus floridanus
Camponotus foreli
Camponotus herculeanus
Camponotus impressus
Camponotus japonicus
Camponotus kiusiuensis
Camponotus laevigatus
Camponotus lateralis
Camponotus ligniperdus
Camponotus linnaei
Camponotus mitis
Camponotus modoc
Camponotus mucronatus
Camponotus mus
Camponotus nawai
Camponotus nearcticus
Camponotus nipponicus
Camponotus noveboracensis
Camponotus obscuripes
Camponotus ocreatus
Camponotus papua
Camponotus parius
Camponotus pennsylvanicus
Camponotus piceus
Camponotus pilicornis
Camponotus punctulatus
Camponotus rectangularis
Camponotus rufipes
Camponotus rufoglaucus
Camponotus salvini
Camponotus sanctaefidei
Camponotus sericeiventris
Camponotus sericeus
Camponotus sexguttatus
Camponotus socius
Camponotus solon
Camponotus sp-impressusgroup
Camponotus sp1
Camponotus sp2
Camponotus sp3
Camponotus sp4
Camponotus sp5
Camponotus sp6
Camponotus sp7
Camponotus sp8
Camponotus sp9
Camponotus spANIC-1
Camponotus spANIC-10
Camponotus spANIC-11
Camponotus spANIC-12
Camponotus spANIC-13
Camponotus spANIC-14
Camponotus spANIC-2
Camponotus spANIC-3
Camponotus spANIC-5
Camponotus spANIC-8
Camponotus spANIC-9
Camponotus spvariegatuscomplex
Camponotus striatus
Camponotus subbarbatus
Camponotus sylvaticus
Camponotus Taylori
Camponotus thraso
Camponotus truncatus
Camponotus universitatis
Camponotus vagus
Camponotus variegatus
Camponotus vicinus
Camponotus vitiosus
Haploid
Number
22.0
19.0
17.0
13.5
14.0
14.0
14.0
10.0
13.0
14.0
20.0
25.0
20.0
20.0
18.0
20.0
22.0
26.0
9.0
19.0
20.0
19.0
18.0
20.0
23.0
23.0
16.0
19.0
10.0
19.0
23.0
24.0
16.0
16.0
18.0
10.0
20.0
12.0
20.0
14.0
13.0
9.0
Eusocial
Colony Size
Parasite
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
300
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1777
10000
19688
250
1950
31845
30
70
10900
300
2800
350
3698
5500
31
80
3560
51000
393
Polyandry
Polygyny
Low
Relatedness
0
0
0
1
0
0
1
0
0
0
0
0
0
0
1
1
0
0
1
1
1
0
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
Species
Camponotus vitreus
Cardiocondyla batesii
Cardiocondyla elegans
Cardiocondyla emeryi
Cardiocondyla nuda
Cardiocondyla obscurior
Cardiocondyla paradoxa
Cardiocondyla sp-Myrmobrachys
Cardiocondyla sp1
Cardiocondyla sp10
Cardiocondyla sp11
Cardiocondyla sp12
Cardiocondyla sp13
Cardiocondyla sp14
Cardiocondyla sp15
Cardiocondyla sp16
Cardiocondyla sp17
Cardiocondyla sp18
Cardiocondyla sp19
Cardiocondyla sp20
Cardiocondyla sp21
Cardiocondyla sp22
Cardiocondyla sp7
Cardiocondyla sp8
Cardiocondyla sp9
Cardiocondyla thoracica
Cardiocondyla wroughtonii
Carebara asina
Carebara pygmaeus
Carebara sauteri
Carebara sp1
Carebara sp2
Carebara sp3
Carebara sp4
Carebara sp5
Carebara sp6
Carebara sp7
Carebara sp8
Carebara spANIC-6
Carebara urichi
Carebara vidua
Carinostigmus filippovi
Cataglyphis bicolor
Cataglyphis bombycinus
Cataglyphis cursor
Cataglyphis hannae
Cataglyphis ibericus
Cataglyphis livida
Cataglyphis piliscapus
Cataglyphis sabulosa
Cataglyphis setipes
Catolaccus grandis
Celetrigona longicornis
Centromyrmex alfaroi
Centromyrmex feae
Centromyrmex sellaris
Cephalotrigona capitata
Cephus cinctus
Cerapachys biroi
Cerapachys brevis
Cerapachys cfcribrinodis
Cerapachys edentata
Cerapachys neotropicus
Cerapachys opaca
Cerapachys polynikes
Cerapachys sp1
Cerapachys sp2
Ceratina calcarata
Ceratina dupla dupla
Cerceris antipodes
Chalepoxenus kutteri
Chalepoxenus muellerianus
Chalicodoma sculpturalis
Haploid
Number
14.0
20.0
20.0
20.0
13.5
26.0
9.0
9.0
16.0
17.0
10.0
17.0
19.0
20.0
10.0
19.0
22.0
20.0
19.0
22.0
18.0
18.0
22.0
16.0
13.0
17.0
21.0
18.0
22.0
19.0
14.0
26.0
26.0
27.0
15.0
22.0
17.0
9.0
14.0
23.0
25.0
25.0
12.0
12.0
16.0
Eusocial
Colony Size
Parasite
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
0
0
0
1
1
0
4000
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
50
50
50
70
50
20000
568
Polyandry
Polygyny
Low
Relatedness
1
2
0
0
1
1
1
1
1
1
1
1
1
1
1
0
1
2
0
0
0
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
1
1
2658
333
369
408
1250
1
600
235
100
20
0
0
0
0
0
0
0
0
0
0
0
0
1
1
394
Species
Chalicodoma spissula
Chalybion californicum
Chalybion japonicum
Charmon cruentatus
Chasmias motatorius
Cimbex femorata
Cinetus lanceolatus
Cirrospilus dialllus
Cladius morio
Cladius pectinicornis
Cleptotrigona cubiceps
Coelichneumon cyaniventris
Coelichneumon sugillatorius
Coelioxys sp
Coelopisthia extenta
Colobostruma alinodis
Colobostruma sp1
Colobostruma spANIC-1
Colpoclypeus florus
Colpognathus celerator
Copidosoma buyssoni
Copidosoma floridanum
Copidosoma gelechiae
Copidosoma truncatellum
Corymbas fujisana
Corymbas nipponica
Cotesia glomeratus
Cratichneumon viator
Crematogaster arcuata
Crematogaster artifex
Crematogaster ashmeadi
Crematogaster atkinsoni
Crematogaster BB
Crematogaster biroi
Crematogaster brasiliensis
Crematogaster brunnea
Crematogaster bryophilia
Crematogaster carinata
Crematogaster crinosa
Crematogaster curvispinosa
Crematogaster distans
Crematogaster dohrniartifex
Crematogaster dohrnirogenhoferi
Crematogaster elegans
Crematogaster erecta
Crematogaster hespera
Crematogaster irritabilissubtilis
Crematogaster jardinero
Crematogaster KA
Crematogaster laboriosa
Crematogaster laevis
Crematogaster larreae
Crematogaster limata
Crematogaster lineolata
Crematogaster LO
Crematogaster longispina
Crematogaster minutissima
Crematogaster minutus
Crematogaster monteverdensis
Crematogaster montezumia
Crematogaster nigropilosa
Crematogaster raptor
Crematogaster RE
Crematogaster rochai
Crematogaster rogenhoferi
Crematogaster rothneyi
Crematogaster scutellaris
Crematogaster smithi
Crematogaster sotobosque
Crematogaster sp1
Crematogaster sp10
Crematogaster sp11
Crematogaster sp12
Haploid
Number
16.0
24.0
5.0
17.0
8.0
10.0
6.0
6.0
6.0
18.0
13.0
13.0
16.0
5.0
11.0
10.0
11.0
6.0
11.0
12.0
9.5
11.0
10.0
10.0
10.0
12.0
14.0
12.0
18.0
13.0
25.0
20.0
13.0
12.0
28.5
Eusocial
Colony Size
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Parasite
Polyandry
Polygyny
Low
Relatedness
0
1
0
0
1
0
0
0
0
1
1
0
0
1
1
0
1
1
0
0
1
0
1
0
1
1
1
1
1
0
1
0
0
0
1
1
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
1
1
1
1
1
1
52275
25000
500
99
56947
5690
300
5000
39
1100
789
200
221
200
150
125
5128
165
395
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Species
Crematogaster sp13
Crematogaster sp4
Crematogaster sp5
Crematogaster sp6
Crematogaster sp7
Crematogaster sp8
Crematogaster sp9
Crematogaster spANIC-1
Crematogaster spANIC-2
Crematogaster subnuda
Crematogaster subtilis
Crematogaster sumichrasti
Crematogaster torosa
Croesus japonicus
Croesus septentrionalis
Croesus varus
Cryptopone motschulskyi
Cryptopone rotundiceps
Cryptopone sauteri
Cryptopone testacea
Cullumanobombus rufocincus
Cylindromyrmex brasiliensis
Cynips divisa
Cynips erinacea
Cyphomyrmex cornutus
Cyphomyrmex costatus
Cyphomyrmex longiscapus
Cyphomyrmex minutus
Cyphomyrmex morschi
Cyphomyrmex muelleri
Cyphomyrmex rimosus
Dacetinops cibdelus
Dacetinops concinnus
Daceton armigerum
Dacnusa sibirica
Dacnusa sp1
Dactylurina staudingeri
Diacamma australe
Diacamma ceylonense
Diacamma cyaneiventre
Diacamma rugosum
Diacamma sp1
Diacamma sp2
Diacamma sp3
Diacamma sp4
Diacamma sp5
Diacamma sp6
Diacamma sp7
Diacamma spfromMalaysia
Diacamma spJapan
Diadasia enavata
Diadromus prosopius
Diadromus pulchellus
Diadromus subtilicornis
Diadromus troglodytes
Diadromus varicolor
Diaeretiella rapae
Dianthidium heterulkeiheterulkei
Diastrophus nebulosus
Dibrachys sp1
Dicaelotus pumilis
Dicaelotus spnrParvulus
Diglyphus isaea
Dinoponera australis
Dinoponera gigantea
Dinoponera lucida
Dinoponera quadriceps
Diphyus latebricola
Diphyus raptorius
Diplolepis elganteria
Diplolepis nervosum
Diplolepis rosae
Diplolepis spinosissimae
Haploid
Number
13.0
13.0
13.0
18.0
18.0
12.0
13.0
12.0
14.5
18.0
8.0
8.0
8.0
6.0
14.0
9.0
19.0
17.0
10.0
12.0
11.0
10.0
16.0
8.0
17.0
17.0
7.0
18.0
22.0
15.0
33.0
29.0
15.0
11.0
11.0
11.0
11.0
11.0
6.0
15.0
10.0
5.0
11.0
11.0
41.0
57.0
46.0
12.0
12.0
9.0
9.0
10.5
9.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
1
1
0
0
1
1
1
1
1
1
1
1
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
Colony Size
5000
1000
1
1
1
20
Parasite
Polyandry
Polygyny
Low
Relatedness
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
2072.5
222
152.67
89.5
252
109
200.5
10
5000
0
0
0
0
0
0
0
0
0
0
1
129
332.33
214
40
247
86
118
1
1
1
1
1
1
1
1
1
1
1
1
21.5
74
90
82
1
1
1
1
1
1
396
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Species
Diprion nipponicus
Diprion pini
Diprion similis
Dirhinus himalayanus
Dirophanes callopus
Dirophanes fulvitarsis
Dirophanes invisor
Discoelius japonicus
Discothyrea sp1
Doleromyrma sp1
Doleromyrma sp2
Dolerus aeneus
Dolerus ephippiatus
Dolerus gessneri
Dolerus hematodes
Dolerus lewisii
Dolerus niger
Dolerus nigratus
Dolerus similisjaponicus
Dolerus subfasciatus
Dolerus varispinus
Dolerus yokohamensis
Dolichoderus bispinosus
Dolichoderus imitator
Dolichoderus lamellosus
Dolichoderus lutosus
Dolichoderus mariae
Dolichoderus plagiatus
Dolichoderus pustulatus
Dolichoderus quadripunctatus
Dolichoderus scabridus
Dolichoderus sp1
Dolichoderus taschenbergi
Dolichoderus thoracicus
Dolichoderus validus
Dolichovespula arenaria
Dolichovespula maculata
Dolichovespula media
Dolichovespula norwegica
Dolichovespula saxonica
Dolichovespula sylvestris
Dorylus laevigatus
Dorylus molestus
Dorylus wilverthi
Dorymyrmex bicolor
Dorymyrmex bicolor
Dorymyrmex bossutus
Dorymyrmex bureni
Dorymyrmex elegans
Dorymyrmex flavus
Dorymyrmex grandulus
Dorymyrmex insanus
Dorymyrmex pulchellus
Dorymyrmex pyramicus
Dorymyrmex reginicula
Dorymyrmex thoracicus
Dryocosmus kuriphilus
Dulophanes morio
Dyspetes arrogator
Echinopla sp1
Eciton burchelli
Eciton dulciumcrassinode
Eciton hamatum
Eciton lucanoides
Eciton mexicanum
Eciton vagansangustatum
Ectatomma brunneum
Ectatomma edentatum
Ectatomma muticum
Ectatomma opaciventre
Ectatomma parasiticum
Ectatomma permagnum
Ectatomma ruidum
Haploid
Number
14.0
10.5
10.5
5.0
9.0
10.0
10.0
12.0
15.0
7.0
6.0
8.0
8.0
8.0
8.0
14.0
8.0
8.0
9.0
9.0
8.0
8.0
14.0
14.0
9.0
15.0
13.0
13.0
9.0
9.0
9.0
10.0
6.0
10.0
12.0
22.0
23.0
20.0
23.0
Eusocial
Colony Size
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Parasite
Polyandry
Polygyny
Low
Relatedness
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
1
1
1
1
2
0
0
0
1
0
0
0
0
2
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
25
80
7000
181
74
44
69
76
325000
18500000
1000
1000
1000
1000
1000
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
762500
400000
104
159
160
397
0
0
0
0
0
0
0
0
0
0
0
1
0
0
Species
Ectatomma tuberculatum
Ectemnius continuus
Ectemnius rubicolanipponis
Elinora koehleri
Empria sp
Empronus obsoletus
Encarsia berlesei
Encarsia formosa
Encarsia pergandiella
Ephedrus sp1
Ephialtes manifestator
Epyris nigir
Eretmocerus eremicus
Eretmocerus mundus
Eriocampa mitsukurii
Eudecatoma biguttata
Eufriesea violacea
Euglossa cyanaspis
Euglossa hyacinthina
Euglossa sp
Eumenes fraternus
Eumenes smithii
Euodynerus foraminatus scutellaris
Euodynerus hidalgo
Euodynerus quadrifasciatus
Eupelmus vesicularis
Eurhopalothrix biroi
Eurhopalothrix floridanus
Eurhopalothrix procera
Eurhopalothrix sp1
Eurylabus torvus
Eurytoma californica
Eutomostethus juncivorus
Exomalopsis aureopilosa
Exomalopsis sp
Fervidobombus atratus
Fervidobombus californicus
Fervidobombus fervidus
Fervidobombus morio
Fervidobombus pennsylvanicus
Forelius foetida
Forelius mccooki
Forelius pruinosus
Forelius pruinosus
Formica 3spp-fuscagr
Formica 4spp
Formica aquilonia
Formica archboldi
Formica argentea
Formica bradleyi
Formica candida
Formica cinerea
Formica corsica
Formica cunicularia
Formica dakotensis
Formica dirksi
Formica dolosa
Formica exsecta
Formica exsectoides
Formica foreli
Formica frontalis
Formica fusca
Formica gagates
Formica gerardi
Formica grouvellei
Formica incerta
Formica japonica
Formica lemani
Formica lugubris
Formica montana
Formica neorufibarbis
Formica obscuripes
Formica opaciventris
Haploid
Number
18.0
14.0
10.0
15.0
10.0
5.0
6.0
17.0
15.0
14.0
9.0
9.0
15.0
21.0
20.0
21.0
8.0
5.0
5.0
9.0
10.0
10.0
6.0
9.0
8.0
20.0
19.0
18.0
20.0
18.0
16.0
16.0
27.0
26.0
26.0
26.0
27.0
27.0
26.0
26.0
26.0
27.0
27.0
27.0
27.0
27.0
26.0
27.0
26.0
Eusocial
Colony Size
Parasite
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
428
0
Polyandry
Polygyny
1
Low
Relatedness
1
0
0
1
1
1
1
1
0
1
1
1
0
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
50
100
0
0
0
0
1
1
1
1
1
87.5
10000
400000
500
630
56089.5
139938
500
1191
3567
4000
650
398
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
Species
Formica pallidefulva
Formica paralugubris
Formica pergandei
Formica podzolica
Formica polyctena
Formica pratensis
Formica pressilabris
Formica reflexa
Formica rufa
Formica rufibarbis
Formica sanguinea
Formica schaufussidolosa
Formica selysi
Formica subintegra
Formica subrufa
Formica subsericea
Formica talbotae
Formica transkaucasica
Formica truncorum
Formica ulkei
Formica uralensis
Formica yessensis
Formicoxenus chamberlini
Formicoxenus hirticolis
Formicoxenus nitidulus
Formicoxenus provancheri
Formicoxenus quebecensis
Frieseomelitta doederleini
Frieseomelitta ghiliani
Frieseomelitta languida
Frieseomelitta varia
Gelis sp1
Geotrigona mombuca
Gigantiops destructor
Gilpinia abieticola
Gilpinia frutetorum
Gilpinia hercyniae
Gilpinia pallida
Gilpinia polytoma
Glypta lapponica
Gnamptogenys annulata
Gnamptogenys bicolor
Gnamptogenys binghamii
Gnamptogenys hartmani
Gnamptogenys horni
Gnamptogenys ingeborgae
Gnamptogenys macretes
Gnamptogenys menadensis
Gnamptogenys regularis
Gnamptogenys sp2
Gnamptogenys sp3
Gnamptogenys striatula
Gnamptogenys strigata
Gnamptogenys tornata
Gorytes atricornis
Habrobracon hebetor
Habrobracon juglandis
Habrobracon pectinophorae
Habrobracon serinopae
Halictus aerarius
Halictus hesperus
Halictus ligatus
Halictus lutescens
Halictus marginatus
Halictus poeyi
Harpagoxenus canadensis
Harpagoxenus sublaevis
Harpegnathos saltator
Hemibeleses nigriceps
Hemichroa alni
Heptamelus(Heptamelus) ochroleucus
Heptamelus(Pseudoheptamelus) runari
Heterischnus nigricollis
Haploid
Number
26.0
26.0
26.0
26.0
26.0
26.0
27.0
26.0
26.0
26.0
27.0
26.0
26.0
26.0
14.0
16.0
15.0
11.0
14.0
15.0
15.0
15.0
15.0
13.0
15.0
39.0
7.0
7.0
7.0
7.0
6.0
9.0
34.0
11.0
21.0
23.0
17.0
10.0
10.5
10.0
10.0
14.0
18.0
20.0
16.0
9.0
10.0
7.0
11.0
Eusocial
Colony Size
Parasite
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
1
1
1
1
1
1
1
1
1
0
0
0
0
0
2556.5
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
3280
5500000
100000
600
40000
8916
55000
307000000
1
2500
133
1
1
1
1
1
1
89
317
28
15
40
204.5
24
Polyandry
Polygyny
1
0
1
Low
Relatedness
0
1
1
1
1
1
0
1
1
1
1
1
1
0
1
0
0
1
1
1
1
0
0
1
1
1
1
1
0
1
0
1
1
0
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
1
1
1
1
149
589
1
295
1
1
1
1
1
399
1
1
0
0
0
0
1
0
0
1
0
0
1
Species
Heterischnus truncator
Heteroponera dolo
Heteroponera relicta
Heterospilus prosopidis
Homotherus locutor
Hoplitis robusta
Hylaeus affinis
Hylaeus ellipticus
Hylaeus nippon
Hylaeus sp1
Hylaeus sp2
Hylaeus stevensi
Hylaeus transversali
Hypoponera confinis
Hypoponera eduardi
Hypoponera gleadowi
Hypoponera inexorata
Hypoponera opaciceps
Hypoponera opacior
Hypoponera pruinosa
Hypoponera sp1
Hypoponera sp2
Hypoponera sp3
Hypoponera sp4
Hypoponera sp5
Hypoponera sp6
Hypoponera spANIC-1
Hypoponera spANIC-2
Hypotrigona araujoi
Hypotrigona braunsi
Hypotrigona gribodoi
Ichneumon albiger
Ichneumon amphibolus
Ichneumon bucculentus
Ichneumon confusor
Ichneumon crassifemur
Ichneumon croceipes
Ichneumon extensorius
Ichneumon formosus
Ichneumon gracilentus
Ichneumon gracilicornis
Ichneumon ingratus
Ichneumon inquinatus
Ichneumon insidiosus
Ichneumon lugens
Ichneumon melanotis
Ichneumon minutorius
Ichneumon molitorius
Ichneumon nereni
Ichneumon sarcitorius
Ichneumon stramentarius
Ichneumon submarginatus
Ichneumon suspiciosus
Ichneumon validicornis
Ichneumonidae sp1
Iridomyrmex anceps
Iridomyrmex anceps2
Iridomyrmex gracilis
Iridomyrmex humilis
Iridomyrmex mattiroloi
Iridomyrmex purpureus
Iridomyrmex scrutator
Iridomyrmex sp4
Iridomyrmex spANIC-11
Iridomyrmex spANIC-12
Iridomyrmex spANIC-13
Iridomyrmex spANIC-14
Iridomyrmex spANIC-15
Iridomyrmex spANIC-16
Iridomyrmex spANIC-17
Iridomyrmex spANIC-5
Iridomyrmex spANIC-6
Isepeolus viperinus
Haploid
Number
11.0
12.0
11.0
17.0
11.0
16.0
11.0
8.0
18.0
28.0
14.0
8.0
19.0
12.0
19.0
19.0
18.0
19.0
19.0
14.0
14.0
12.0
12.0
12.0
12.0
12.0
12.0
12.0
11.0
12.0
11.0
12.0
13.0
12.0
12.0
12.0
12.0
12.0
11.0
12.0
10.0
12.0
12.0
12.0
15.0
18.0
24.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
7.0
9.0
16.0
Eusocial
Colony Size
0
1
1
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
Parasite
Polyandry
Polygyny
Low
Relatedness
1
1
1
1
1
1
0
0
1
1
1
1
1
1
1
1
1
32
50
50
50
34
24
21
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2500
425
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
37500
3000
1
400
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Species
Janus integer
Labidus coecus
Labidus praedator
Labidus spininodis
Laelius utilis
Lagidina irritans
Lagidina platycerus
Lariophagus distinguesdus
Larra bicolor
Lasioglossum allodalum
Lasioglossum cooleyi
Lasioglossum duplex
Lasioglossum imitatum
Lasioglossum laevissumum
Lasioglossum lineatulus
Lasioglossum lineatulus
Lasioglossum malachurum
Lasioglossum marginatum
Lasioglossum rhytidophorum
Lasioglossum rowheri
Lasioglossum taeniolellum
Lasioglossum versatum
Lasioglossum zephyrus
Lasius alienus
Lasius brunneus
Lasius carniolicus
Lasius cinereus
Lasius emarginatus
Lasius flavus
Lasius fuliginosus
Lasius grandis
Lasius lasioides
Lasius latipes
Lasius minutus
Lasius mixtus
Lasius myops
Lasius nearcticus
Lasius neglectus
Lasius niger
Lasius pallitarsus
Lasius pliferus
Lasius sabularum
Lasius sakagamii
Lasius talpa
Lasius umbratus
Lepisiota capensis
Lepisiota sp1
Lepisiota sp2
Leptanilla japonica
Leptogenys attenuata
Leptogenys bituberculata
Leptogenys borneensis
Leptogenys castanea
Leptogenys chinensis
Leptogenys diminuta
Leptogenys distinguenda
Leptogenys hysterica
Leptogenys iridescens
Leptogenys kraepelini
Leptogenys minchinii
Leptogenys myops
Leptogenys nitida
Leptogenys ocellifera
Leptogenys peugueti
Leptogenys processionalis
Leptogenys purpurea
Leptogenys schwabi
Leptogenys sp1
Leptogenys sp2
Leptogenys sp3
Leptogenys sp4
Leptogenys sp5
Leptomastix dactylopii
Haploid
Number
24.0
10.0
18.0
18.0
5.0
20.0
18.0
9.0
12.0
12.0
6.0
16.0
13.0
14.5
15.0
15.0
15.0
14.0
15.0
15.0
14.0
15.0
15.0
15.0
9.0
9.0
9.0
23.0
17.5
13.0
23.0
13.0
26.0
24.0
27.0
23.0
24.0
15.0
27.0
Eusocial
Colony Size
0
1
1
1
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
2000000
Parasite
Polyandry
Polygyny
Low
Relatedness
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
1
1
1
1
5.5
13.5
2.5
20.5
270
6
1
32.5
24.5
3000
171
4885
10000000
39756.5
3000
175
90
300
289
367
361.33
40000
21
595
24000
1250
184
53
782
401
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
1
0
0
0
0
1
0
0
0
1
1
0
0
1
1
0
0
1
0
1
0
1
0
0
0
0
1
1
1
0
0
0
1
1
1
0
0
0
Species
Leptomyrmex erytrocephalus
Leptomyrmex fragilis
Leptothorax acervorum
Leptothorax albipennis
Leptothorax ambiguus
Leptothorax curvispinosus
Leptothorax goesswaldi
Leptothorax gredleri
Leptothorax kutteri
Leptothorax longispinosus
Leptothorax muscorum
Leptothorax nylanderi
Leptothorax pacis
Leptothorax pergandei
Leptothorax pocahontas
Leptothorax retractus
Leptothorax sp1
Leptothorax sp10
Leptothorax sp2
Leptothorax sp3
Leptothorax sp4
Leptothorax sp5
Leptothorax sp6
Leptothorax sp7
Leptothorax sp8
Leptothorax sp9
Leptothorax sphagnicola
Leptothorax unifasciatus
Lestrimelitta limao
Leucospis affinis
Leurotrigona muelleri
Leurotrigona pusilla
Linepithema humile
Linepithema pilifer
Linepithema sp1
Liometopum occidentale
Liostenogaster flavolineata
Liostenogaster sp1
Liostenogaster vechti
Lissonota sp
Loderus eversmanniobscurus
Loderus genucinctusinsulicola
Lophomyrmex bedoti
Lophomyrmex sp1
Lordomyrma sp1
Lordomyrma sp2
Macrocentrus grandii
Macrocentrus thoracicus
Macrodiprion nemoralis
Macrophya albipuncta
Macrophya annulitibia
Macrophya apicalis
Macrophya carbonaria
Macrophya coxalis
Macrophya esakiiexilis
Macrophya falsifica
Macrophya fascipennis
Macrophya imitator
Macrophya infumata
Macrophya malaisei
Macrophya montana
Macrophya punctumalbum
Macrophya ribis
Macrophya rohweri
Macrophya rufipes
Macrophya sp
Macrophya timida
Manica rubida
Mastrus smithii
Mayriella abstinens
Megachile albitarsis
Megachile rotundata
Megachile ainu
Haploid
Number
12.0
13.0
8.0
28.0
11.0
24.0
19.2
26.0
18.0
17.5
16.0
17.0
21.0
17.0
12.0
12.0
13.0
14.0
22.0
15.5
13.0
14.0
6.0
8.0
15.0
8.0
9.0
9.0
7.0
11.0
14.0
11.0
19.0
19.0
11.0
13.0
7.0
8.0
10.0
8.0
10.0
10.0
8.0
10.0
12.0
8.0
9.0
10.0
8.0
10.0
12.0
10.0
10.0
10.0
10.0
22.0
13.0
9.0
16.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
Colony Size
350
45.07
38
69
36.5
37
116
Parasite
0
0
0
0
0
0
1
0
1
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
Polyandry
Polygyny
Low
Relatedness
0
1
1
1
1
1
1
1
0
1
0
0
0
1
1
0
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
1
1
0
0
0
1
1
1
1000000
0
0
0
0
27
34
1
1
1
12.5
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
0
1
402
Species
Megachile pseudomonticola
Megachile relativa
Megachile rotundata
Meliplebeia beccari
Meliplebeia nebulata komiensis
Melipona anthidioides
Melipona asilvae
Melipona beecheii
Melipona bicolor bicolor
Melipona capixaba
Melipona compressipes
Melipona crinita
Melipona fasciata melanopleura
Melipona favosa
Melipona interrupta fasciculata
Melipona mandacaia
Melipona marginata
Melipona marginata carrikeri
Melipona marginata marginata
Melipona mondury
Melipona nigra
Melipona panamica
Melipona quadrifasciata
Melipona quinquefasciata
Melipona rufiventris
Melipona scutellaris
Melipona subnitida
Meliponula becarii
Meliponula bocandei
Melissodes desponsa
Melissodes illata
Melittobia chalybii
Meranoplus bicolor
Meranoplus minor
Meranoplus sp1
Meranoplus sp2
Meranoplus sp3
Meranoplus sp4
Meranoplus spinosus
Messor aciculatus
Messor andrei
Messor barbarus
Messor bouvieri
Messor capitatus
Messor ebeninus
Messor minor
Messor sp1
Messor structor
Messor wasmanni
Metapolybia sp
Meteorus gyrator
Meteorus pallipes
Meteorus versicolor
Microbembex monodonta
Microstigmus arlei
Microstigmus brasiliensis
Microstigmus comes
Microstigmus cooperi
Microstigmus crucifex
Microstigmus luederwaldti
Mirax sp1
Mischocyttarus cassununga
Mischocyttarus sp
Miscophus slossonae
Monoctenus nipponicus
Monoctenus suffusus
Monodontomerus clemellti
Monodontomerus montivagus
Monodontomerus obscurus
Monodontomerus saltuosus
Monomorium destructor
Monomorium dichroum
Monomorium floricola
Haploid
Number
16.0
16.0
16.0
17.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
9.0
14.0
9.0
9.0
9.0
17.0
18.0
5.0
8.0
11.0
10.0
11.0
11.0
11.0
22.0
20.0
21.0
20.0
19.0
10.0
10.0
8.0
3.0
5.0
3.0
4.0
3.0
10.0
32.0
34.0
15.0
7.0
6.0
6.0
5.0
5.0
8.0
Eusocial
Colony Size
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
1
1
0
1
1
0
0
0
0
0
0
0
1
1
1
1
1
1
Parasite
Polyandry
Polygyny
Low
Relatedness
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
1
0
0
0
1
1
1
1
1097.5
894
2000
384.33
210
191
483.33
666.67
1
150
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
3.5
3.5
6.5
0
3.5
1
1
1
1
1
1
1
0
0
0
403
Species
Monomorium glabrum
Monomorium indicum
Monomorium latinode
Monomorium minimum
Monomorium orientale
Monomorium pharaonis
Monomorium rothsteini
Monomorium salomonis
Monomorium scabriceps
Monomorium sp1
Monomorium sp10
Monomorium sp11
Monomorium sp12
Monomorium sp2
Monomorium sp3
Monomorium sp4
Monomorium sp5
Monomorium sp6
Monomorium sp7
Monomorium sp8
Monomorium sp9
Monomorium subopacum
Monomorium viride
Monomorium whitei
Monosoma pulveratum
Mourella caerulea
Muscidifurax zaraptor
Mycocepurus goeldii
Mycocepurus smithii
Mycocepurus sp1
Myopias sp1
Myopias sp2
Myopias sp3
Myrmecia banksi
Myrmecia brevinoda
Myrmecia cephalotes
Myrmecia chasei
Myrmecia croslandi
Myrmecia desertorum
Myrmecia dispar
Myrmecia forficata
Myrmecia froggatti
Myrmecia fulvipes
Myrmecia gulosa
Myrmecia haskinsorum
Myrmecia imaii
Myrmecia mandibularis
Myrmecia michaelseni
Myrmecia nigrocincta
Myrmecia occidentalis
Myrmecia pavida
Myrmecia picta
Myrmecia piliventris
Myrmecia pilosula
Myrmecia pyriformis
Myrmecia simillima
Myrmecia spcfarnoldi
Myrmecia spcffulvipes
Myrmecia tarsata
Myrmecia tepperi
Myrmecia varians
Myrmecia vindex
Myrmecina americana
Myrmecina graminicola
Myrmecina nipponica
Myrmecina sp1
Myrmecina sp2
Myrmecina spA
Myrmecina transversa
Myrmica americana
Myrmica brevispinosa
Myrmica fracticornis
Myrmica hirsuta
Haploid
Number
19.0
11.0
35.0
11.0
10.0
11.0
11.0
19.0
11.0
19.0
11.0
11.0
21.0
16.0
11.0
11.0
11.0
11.0
11.0
17.0
17.0
11.0
12.0
8.0
17.0
5.0
8.0
8.0
9.5
42.0
33.0
23.0
1.5
26.0
21.3
19.0
9.0
3.5
28.5
27.0
11.0
32.0
22.0
17.8
10.5
41.0
35.0
29.3
6.0
35.0
37.5
14.0
14.0
33.0
34.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Colony Size
3000
47250
10000
20225
1
1
1352
163
60
55
30
2794
29.76
62.66
211
38
108.56
908.31
402
18
821
6.91
165.5
707.5
1361.5
162.4
1126
117
145
190.5
37
27
130
100
445
627
404
Parasite
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
Polyandry
Polygyny
Low
Relatedness
0
1
1
0
1
1
1
1
1
1
1
1
1
0
0
1
0
1
0
0
0
1
0
1
0
0
0
0
0
1
1
0
0
0
1
0
0
0
0
1
0
0
0
1
0
1
1
1
1
1
Species
Myrmica incompleta
Myrmica kotokui
Myrmica latifrons
Myrmica lobicornis
Myrmica lobifrons
Myrmica pinetorum
Myrmica punctiventris
Myrmica rubra
Myrmica ruginodis
Myrmica sabuleti
Myrmica scabrinodis
Myrmica schencki
Myrmica schenckioides
Myrmica spatulata
Myrmica spinosior
Myrmica sulcinodis
Myrmica tahoensis
Myrmicaria brunnea
Myrmicaria eumenoides
Myrmicaria opaciventris
Myrmicaria sp1
Myrmicaria sp2
Myrmicaria sp3
Myrmicaria sp4
Myrmicaria sp5
Myrmicocrypta ednaella
Myrmicocrypta sp1
Myrmoteras barbouri
Myrmoteras toro
Myrmoxenus adlerzi
Myrmoxenus algeriana
Myrmoxenus bernardi
Myrmoxenus corsica
Myrmoxenus goesswaldi
Myrmoxenus gordiagini
Myrmoxenus kraussei
Myrmoxenus ravouxi
Myrmoxenus stumperi
Mystrium camillae
Mystrium mysticum
Mystrium oberthueri
Mystrium rogeri
Mystrium sp1
Nannotrigona perilampoides
Nannotrigona sp
Nannotrigona sp1
Nannotrigona testaceicornis
Nasonia vitripennis
Neivamyrmex alfaroi
Neivamyrmex asper
Neivamyrmex californicus
Neivamyrmex carolinensis
Neivamyrmex gibbatus
Neivamyrmex humilis
Neivamyrmex impudens
Neivamyrmex leonardi
Neivamyrmex mexicanus
Neivamyrmex nigrescens
Neivamyrmex nyensis
Neivamyrmex opacithorax
Neivamyrmex pilosus
Neivamyrmex punctaticeps
Neivamyrmex swainsonii
Neivamyrmex texanus
Nematinus fuscipennis
Nematinus luteus
Nematinus willigkiae
Nematus alaskaensis
Nematus dimmockii
Nematus erichsonii
Nematus geniculatus
Nematus leucotrochus
Nematus lucidus
Haploid
Number
12.0
23.5
24.0
23.0
22.0
23.0
26.0
22.0
22.0
22.0
22.0
22.0
23.0
15.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
16.0
17.0
17.0
17.0
5.5
18.0
8.0
8.0
8.0
8.0
8.0
7.0
8.0
9.0
8.0
Eusocial
Colony Size
Parasite
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
918
280
255
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
1
1
0
0
0
0
0
0
1243
396
74.9
5500
2035.5
408
296
143.5
175
19898.5
147230
145
11
23
40
58
43
35
156
29
Polyandry
0
0
1
1
Polygyny
1
Low
Relatedness
1
1
1
1
0
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
2500
1
30000
125000
30000
30000
1
1
1
1
1
1
1
1
1
405
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
2
Species
Nematus melanaspis
Nematus nigricornis
Nematus obductus
Nematus olfaciens
Nematus pallipes
Nematus pavidus
Nematus ribesii
Nematus ruficornis
Nematus rufipes
Nematus sp
Nematus sp1
Nematus sp2
Nematus sp3
Nematus viminalis
Nematus viridescens
Neodiprion abietis
Neodiprion compar
Neodiprion dubiosus
Neodiprion lecontei
Neodiprion maurus
Neodiprion nanulus
Neodiprion nigroscutum
Neodiprion pinetum
Neodiprion prattibanksianae
Neodiprion sertifer
Neodiprion sp1
Neodiprion sp2
Neodiprion swainei
Neodiprion taedaetaedae
Neodiprion tsugae
Neodiprion virginiana
Neostromboceros itoi
Neostromboceros nipponicus
Neostromboceros okinawaensis
Neostromboceros sinanensis
Neotheronia bicincta
Neuroterus laeviusculus
Neuroterus numismalis
Neuroterus quescusbaccarum
Nogueirapis mirandula
Nomia nevadensisangelesia
Nothomyrmecia macrops
Notoncus ectatommoides
Ochetellus glaber
Odontomachus bauri
Odontomachus brunneus
Odontomachus chelifer
Odontomachus clarus
Odontomachus coquereli
Odontomachus haematodus
Odontomachus hastatus
Odontomachus latidens
Odontomachus meinerti
Odontomachus panamensis
Odontomachus relictus
Odontomachus rixosus
Odontomachus ruginodus
Odontomachus scalptus
Odontomachus simillimus
Odontomachus sp1
Odontomachus spANIC-1
Odontoponera transversa
Oecophylla longinoda
Oecophylla smaragdina
Omalus djozanushondonis
Onychomyrmex hedleyi
Opisthopsis rufithorax
Orectognathus clarki
Orectognathus darlingtoni
Orectognathus versicolor
Ormyrus sp1
Oronotus binotatus
Orthocentrus sp1
Haploid
Number
8.0
8.0
8.0
9.0
8.0
8.0
9.0
8.0
8.0
9.0
7.0
8.0
8.0
8.0
8.0
8.0
7.0
7.0
7.0
7.0
7.0
7.0
7.0
7.0
7.0
7.0
8.0
7.5
7.0
7.0
7.0
6.0
7.0
7.0
7.0
10.0
10.0
10.0
21.0
47.0
22.0
14.0
22.0
22.0
15.5
22.0
15.0
22.0
22.0
22.0
22.0
22.0
12.0
8.0
19.0
25.0
15.0
11.0
11.0
6.0
11.0
14.0
Eusocial
Colony Size
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
3178.5
1
69.67
416
172.5
38
300
200
375
200
500000
1
1073
104
1
1
1
406
Parasite
Polyandry
Polygyny
Low
Relatedness
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
1
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
Species
Orthopelma mediator
Osmia cornifrons
Osmia cornuta
Osmia glauca
Osmia lignariapropinque
Osmia nigrifrons
Osmia pentstemonis
Osmia taurus
Oxytrigona spcfflaveola
Oxytrigona tataira
Pachycondyla aenescens
Pachycondyla analis
Pachycondyla apicalis
Pachycondyla arhuaca
Pachycondyla astuta
Pachycondyla caffraria
Pachycondyla carinulata
Pachycondyla chinensis
Pachycondyla concava
Pachycondyla constricta
Pachycondyla crassinoda
Pachycondyla crenata
Pachycondyla curvinodis
Pachycondyla dismarginata
Pachycondyla gilberti
Pachycondyla goeldii
Pachycondyla harpax
Pachycondyla havilandi
Pachycondyla impressa
Pachycondyla insignis
Pachycondyla inversa
Pachycondyla krugeri
Pachycondyla leeuwenhoeki
Pachycondyla lutea
Pachycondyla luteipes
Pachycondyla marginata
Pachycondyla mesonotalis
Pachycondyla metanotalis
Pachycondyla moesta
Pachycondyla pergandei
Pachycondyla purpurascens
Pachycondyla rubiginosa
Pachycondyla rubra
Pachycondyla rufipes
Pachycondyla sp1
Pachycondyla sp11
Pachycondyla sp2
Pachycondyla sp3
Pachycondyla sp4
Pachycondyla sp5
Pachycondyla sp6
Pachycondyla sp7
Pachycondyla sp8
Pachycondyla sp9
Pachycondyla stigma
Pachycondyla striata
Pachycondyla striatula
Pachycondyla sublaevis
Pachycondyla subversa
Pachycondyla succedanea
Pachycondyla tarsatus
Pachycondyla tridentata
Pachycondyla unidentata
Pachycondyla venusta
Pachycondyla verenae
Pachycondyla villosa
Pachyprotasis asteris
Pachyprotasis caerulescenskashmirica
Pachyprotasis erratica
Pachyprotasis fukii
Pachyprotasis hayasuensis
Pachyprotasis hiensis
Pachyprotasis hiyodorii
Haploid
Number
11.0
16.0
16.0
16.0
16.0
16.0
16.0
16.0
17.0
17.0
24.0
12.0
10.0
12.0
11.0
27.0
15.0
31.0
13.0
13.5
15.0
12.0
48.0
47.0
15.0
8.0
8.0
11.0
23.0
13.0
35.0
13.0
38.0
16.0
24.0
11.0
26.0
19.0
22.0
30.0
11.0
14.0
11.0
18.0
18.0
6.0
52.0
13.5
7.0
14.0
6.0
24.0
28.4
17.0
10.0
9.0
10.0
10.0
10.0
10.0
10.0
Eusocial
Colony Size
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
583
127
276
50
30
500
28
20
52.5
1198.5
19
9
15
9.6
1214.67
1
1
1
1
1
1
1
407
Parasite
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Polyandry
Polygyny
Low
Relatedness
1
0
1
0
0
1
1
1
1
0
1
1
0
1
1
1
1
0
0
0
1
0
0
0
1
0
1
0
1
1
1
Species
Pachyprotasis iwatai
Pachyprotasis longicornis
Pachyprotasis Malaise
Pachyprotasis malaisei
Pachyprotasis nigrinotata
Pachyprotasis nogusai
Pachyprotasis okutanii
Pachyprotasis pallidiventris
Pachyprotasis rapae
Pachyprotasis sasabensis
Pachyprotasis sawadai
Pachyprotasis sengaminensis
Pachyprotasis serii
Pachyprotasis sp1
Pachyprotasis sp10
Pachyprotasis sp2
Pachyprotasis sp3
Pachyprotasis sp4
Pachyprotasis sp5
Pachyprotasis sp6
Pachyprotasis sp7
Pachyprotasis sp8
Pachyprotasis sp9
Pachyprotasis tanakai
Pachyprotasis volatilis
Pachyprotasis yamahakkai
Pachyprotasis zukaensis
Papyrius nitidus
Paracharactus leucopodus
Parachartergus colobopterus
Parachartergus smithii
Paraponera clavata
Paratrechina arenivaga
Paratrechina concinna
Paratrechina faisonensis
Paratrechina flavipes
Paratrechina ha
Paratrechina indica
Paratrechina longicornis
Paratrechina pallida
Paratrechina parvula
Paratrechina phantasma
Paratrechina pubens
Paratrechina sp1
Paratrechina sp10
Paratrechina sp11
Paratrechina sp2
Paratrechina sp3
Paratrechina sp4
Paratrechina sp5
Paratrechina sp6
Paratrechina sp7
Paratrechina sp8
Paratrechina sp9
Paratrechina spANIC-1
Paratrechina steinheili
Paratrechina vividula
Paratrechina wojciki
Paratrigona subnuda
Partamona ailyae
Partamona cupira
Partamona helleri
Partamona mulata
Partamona pearsoni
Partamona peckolti
Partamona peckoltimusarum
Partamona seridoensis
Partamona spn
Partamona vicina
Patrocloides chalybeatus
Pemphredon diervillae
Pemphredon krombeini
Pemphredon lethifer
Haploid
Number
10.0
10.0
11.0
11.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
11.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
8.0
10.0
27.0
15.0
8.0
7.0
8.0
15.0
13.0
14.0
8.0
14.0
15.0
8.0
15.0
15.0
15.0
17.0
17.0
17.0
17.0
17.0
17.0
17.0
17.0
17.0
17.0
17.0
8.0
4.0
8.0
8.0
Eusocial
Colony Size
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Parasite
Polyandry
Polygyny
Low
Relatedness
0
1
1
0
0
1
1
0
0
0
0
0
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
1
393
150
1031.5
100
100
50.13
890
351
2000
500
100
100
300000
200
150
60
168
100
3012.5
1
1
1
1
408
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Species
Perga sp
Perineura esakii
Perineura japonica
Perineura okutanii
Perineura pictipennis
Perineura sp
Peristenus digoneutis
Peristenus stygicus
Phaenocarpa persimilis
Phaeogenes melanogonos
Phaeogenes nigridens
Phaeogenes semivulpinus
Phaeogenes spiniger
Pheidole acutidens
Pheidole adrianoi
Pheidole argentina
Pheidole bic
Pheidole bicarinata
Pheidole biconstricta
Pheidole bicornis
Pheidole bilimeki
Pheidole binghamii
Pheidole BSH
Pheidole capellinii
Pheidole CCC
Pheidole CCR
Pheidole cerebrosior
Pheidole cho
Pheidole cramptoni
Pheidole dentata
Pheidole dentigula
Pheidole desertorum
Pheidole DSS
Pheidole elecebra
Pheidole embolopyx
Pheidole eye
Pheidole fallax
Pheidole fervens
Pheidole fervida
Pheidole fiorii
Pheidole flavens
Pheidole floridana
Pheidole gilvescens
Pheidole GLO
Pheidole hfb
Pheidole hortensis
Pheidole hyatti
Pheidole indica
Pheidole inquilina
Pheidole int
Pheidole lanuginosa
Pheidole latinoda
Pheidole megacephala
Pheidole metallescens
Pheidole MGA
Pheidole MGS
Pheidole MIC
Pheidole microgyna
Pheidole MIN
Pheidole minutula
Pheidole moerens
Pheidole morrisi
Pheidole multispina
Pheidole mus
Pheidole NEB
Pheidole neokohli
Pheidole NGA
Pheidole nigricula
Pheidole NIT
Pheidole nitidula
Pheidole noda
Pheidole obscurithorax
Pheidole pallidula
Haploid
Number
8.0
17.0
17.0
17.0
17.0
17.0
17.0
11.0
11.0
9.0
11.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
21.0
10.0
6.0
10.0
18.7
11.0
Eusocial
Colony Size
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Parasite
Polyandry
Polygyny
Low
Relatedness
0
0
1
1
0
0
1
0
0
1
0
0
0
1
0
1
0
1
0
0
0
1
0
0
1
0
1
0
1
0
1
0
1
0
1
0
0
0
0
1
0
1
1
1
1
1
1
350
275
200
150
600
206
172
375
134
12
88
873
300
24814
22
36
300
7317.5
1500
1000
500
44
205
126
1000
152
159
62
80
1500
600
9250
95
61
62
61
547
10000
3300
409
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
Species
Pheidole parasitica
Pheidole PCHO
Pheidole PEB
Pheidole pilifera
Pheidole plagiaria
Pheidole PNGA-nigricul
Pheidole pnig
Pheidole pnigb
Pheidole poc
Pheidole porcula
Pheidole PPOC
Pheidole praeses
Pheidole PRUG-L
Pheidole PRUG~B
Pheidole renb
Pheidole rpc
Pheidole rugulosa
Pheidole ruida
Pheidole sbi
Pheidole soritis
Pheidole sp1
Pheidole sp10
Pheidole sp11
Pheidole sp12
Pheidole sp13
Pheidole sp14
Pheidole sp15
Pheidole sp16
Pheidole sp17
Pheidole sp18
Pheidole sp19
Pheidole sp2
Pheidole sp20
Pheidole sp21
Pheidole sp22
Pheidole sp23
Pheidole sp24
Pheidole sp25
Pheidole sp26
Pheidole sp27
Pheidole sp28
Pheidole sp29
Pheidole sp3
Pheidole sp30
Pheidole sp31
Pheidole sp32
Pheidole sp33
Pheidole sp34
Pheidole sp35
Pheidole sp36
Pheidole sp37
Pheidole sp38
Pheidole sp39
Pheidole sp4
Pheidole sp40
Pheidole sp41
Pheidole sp42
Pheidole sp43
Pheidole sp44
Pheidole sp45
Pheidole sp46
Pheidole sp47
Pheidole sp48
Pheidole sp49
Pheidole sp5
Pheidole sp50
Pheidole sp51
Pheidole sp6
Pheidole sp7
Pheidole sp8
Pheidole sp9
Pheidole spe
Pheidole specularis
Haploid
Number
10.0
10.0
9.0
9.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
16.5
10.0
8.0
19.0
9.0
10.0
11.0
10.0
19.0
10.0
10.0
10.0
9.0
10.0
10.0
10.0
18.0
21.0
10.0
10.0
15.0
10.0
10.0
10.0
10.0
14.0
9.0
9.0
9.0
16.0
10.0
9.0
10.0
9.0
9.0
10.0
9.0
10.0
10.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Colony Size
200
300
100
113
105
58
200
200
200
170
60
60
300
124
42
56
75
410
Parasite
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Polyandry
Polygyny
Low
Relatedness
0
0
1
0
1
0
0
0
0
0
Species
Pheidole spininodis
Pheidole subarmata
Pheidole symbiotica
Pheidole teneriffana
Pheidole tepicana
Pheidole tysoni
Pheidole umphreyi
Pheidole vistana
Pheidole woodmasoni
Pheidole zeteki
Pheidologeton diversus
Pheidologeton sp1
Pheidologeton sp2
Philidris cordata
Philomastix sp
Phyllocolpa sp
Plagiolepis alluaudi
Plagiolepis ampeloni
Plagiolepis grassei
Plagiolepis pygmaea
Plagiolepis regis
Plagiolepis schmitzii
Plagiolepis sp1
Plagiolepis sp2
Plagiolepis taurica
Plagiolepis xene
Platythyrea conradti
Platythyrea lamellosa
Platythyrea parallela
Platythyrea punctata
Platythyrea quadridenta
Platythyrea schultzei
Platythyrea tricuspidata
Plebeia denoiti
Plebeia droryana
Plebeia emerina
Plebeia minima
Plebeia mosquito
Plebeia remota
Plebeia saiqui
Plebeia schrottkyi
Plebeia sp1
Plebeia subnuda
Plebeina denoiti
Podomyrma adelaidae
Pogonomyrmex anergismus
Pogonomyrmex apache
Pogonomyrmex badius
Pogonomyrmex barbatus
Pogonomyrmex brevispinosus
Pogonomyrmex californicus
Pogonomyrmex carbonarius
Pogonomyrmex coarctatus
Pogonomyrmex colei
Pogonomyrmex comanche
Pogonomyrmex desertorum
Pogonomyrmex huachucanus
Pogonomyrmex imberbiculus
Pogonomyrmex laticeps
Pogonomyrmex longibarbis
Pogonomyrmex magnacanthus
Pogonomyrmex maricopa
Pogonomyrmex mayri
Pogonomyrmex montanus
Pogonomyrmex occidentalis
Pogonomyrmex pima
Pogonomyrmex rugosus
Pogonomyrmex subnitidus
Pogonomyrmex tenuispinus
Polistes annularis
Polistes apachus
Polistes bellicos
Polistes canadensis
Haploid
Number
10.0
10.0
9.0
9.0
21.0
21.0
8.0
8.0
9.0
9.0
9.0
9.0
9.0
9.0
46.5
18.0
17.5
18.0
18.0
18.0
17.0
18.0
18.0
25.5
16.0
16.0
16.0
16.0
16.0
16.0
16.0
18.0
15.0
16.0
16.0
16.0
16.0
16.0
16.0
23.5
16.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Colony Size
3000
81
3000
Parasite
Polyandry
Polygyny
Low
Relatedness
1
1
0
0
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
2
2
0
0
1
1
2
0
1
2
0
1
0
0
0
0
0
0
0
1
0
0
0
1
0
0
1
1
1
0
0
1
1
0
1
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
1
302.33
115
50
60
33.33
21
29
0
1
1
0
1
0
0
0
0
1
0
0
0
0
0
0
0
2700
1175
2900
1500
300
7268
12358
4533
500
500
41.5
45
250
194
750
603
1665
4103
326
6291.5
5934
3000
156
52.39
411
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
2
2
1
2
Species
Polistes carolinus
Polistes chinensis
Polistes chinensis antennalis
Polistes cinerascens
Polistes dominula
Polistes dorsalis
Polistes exclamans
Polistes fuscatus
Polistes gallicus
Polistes hebraeus
Polistes instabilis
Polistes jadwigae
Polistes mandarinus
Polistes metricus
Polistes nimpha
Polistes omissus
Polistes simillimus
Polistes snelleni
Polistes versicolor versicolor
Polybia bicyttarella
Polybia bistriata
Polybia catillifex
Polybia chrysothorax
Polybia diguetana
Polybia dimidiata
Polybia emaciata
Polybia erythrothorax
Polybia jurinei
Polybia micans
Polybia occidentalis
Polybia parvulina
Polybia paulista
Polybia platycephala
Polybia quadricincta
Polybia rejecta
Polybia ruficeps
Polybia rufitarsis
Polybia scrobalis
Polybia scutellaris
Polybia sericea
Polybia simillima
Polybia singularis
Polybia sp1
Polybia sp2
Polybia striata
Polybioides raphigastra
Polybioides tabidus
Polyergus breviceps
Polyergus lucidus
Polyergus rufescens
Polyergus samurai
Polyrhachis ammon
Polyrhachis bicolor
Polyrhachis debilis
Polyrhachis dives
Polyrhachis furcata
Polyrhachis gribodoi
Polyrhachis hector
Polyrhachis hippomanes
Polyrhachis hirsutula
Polyrhachis hodgsoni
Polyrhachis illaudata
Polyrhachis lacteipennis
Polyrhachis lamellidens
Polyrhachis limbata
Polyrhachis muelleri
Polyrhachis nigropilosa
Polyrhachis omyrmex
Polyrhachis proxima
Polyrhachis rastellata
Polyrhachis rufiventrus
Polyrhachis schellerichae
Polyrhachis sp1
Haploid
Number
19.0
23.0
23.0
27.0
33.0
26.0
21.0
9.0
31.0
26.0
26.0
22.0
14.0
28.0
30.0
31.0
17.0
17.0
17.0
27.0
16.0
17.0
27.0
21.0
21.0
24.0
21.0
20.0
16.0
21.0
21.0
21.0
21.0
Eusocial
Colony Size
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
102.25
244.5
Parasite
55.86
107.56
25
Polyandry
Polygyny
Low
Relatedness
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
1
1
1
0
0
0
1
0
1
0
1
0
0
0
1
0
1
0
0
0
60.02
76.26
80
60
18
30
800
1,250
141
300
500
100
261
900
3457
300
1,150
1,150
1,100
80
40
1,300
500
300
1,000
1,200
3,000
4000
3000
133.75
845
325
1000000
6418
150
9193
475
100
89
1093
60
1422
200
7203
412
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
Species
Polyrhachis sp2
Polyrhachis sp3
Polyrhachis spANIC-1
Polyrhachis spinifera
Polysphincta tuberosa
Pompilidae sp1
Pompilidae sp2
Ponera coarctata
Ponera exotica
Ponera japonica
Ponera pennsylvanica
Ponera scabra
Ponera sp1
Ponera sp2
Ponera testacea
Praon abjectum
Praon sp.
Prenolepis imparis
Prenolepis jerdoni
Prionopelta amabilis
Prionopelta modesta
Prionopelta opaca
Priophorus varipes
Pristomyrmex punctatus
Pristomyrmex pungens
Pristomyrmex sp1
Pristomyrmex sp2
Pristomyrmex sp3
Proatta butteli
Proatta sp1
Probolomyrmex boliviensis
Probolomyrmex dammermani
Probolomyrmex sp1
Proceratium crassicorne
Proceratium croceum
Proceratium goliath
Proceratium pergandei
Proceratium silaceum
Proceratium sp1
Proceratium sp2
Proceratium sp3
Procryptocerus batesi
Procryptocerus mayri
Procryptocerus scabriusculus
Prolasius spANIC-1
Prolasius spANIC-2
Propodea fentoni
Protomognathus americanus
Protonectarina sylveirae
Protopolybia acutiscutis
Protopolybia emortualis
Protopolybia exigua exigua
Protopolybia holoantha
Protopolybia minutissima
Protopolybia pumila
Protopolybia scutellaris
Psenulus carnifronsiwatai
Psenulus maculipes
Pseudaugochloropsis graminea
Pseudaugochloropsis sp
Pseudoamblyteles homocerus
Pseudohemitaxonus dryopteridis
Pseudolasius breviceps
Pseudolasius sp1
Pseudolasius sp2
Pseudolasius sp3
Pseudolasius sp4
Pseudolasius sp5
Pseudolasius xene
Pseudomyrmex apache
Pseudomyrmex concolor
Pseudomyrmex cubaensis
Pseudomyrmex ejectus
Haploid
Number
21.0
20.0
21.0
9.0
15.0
15.0
6.0
6.0
3.5
6.0
6.0
4.0
8.0
19.7
6.0
12.0
11.0
14.0
16.0
14.0
18.0
24.0
9.0
9.0
10.0
11.0
29.0
31.0
21.0
8.0
21.0
8.0
9.0
5.0
14.0
15.0
15.0
8.0
17.0
Eusocial
1
1
1
1
0
0
0
1
1
1
1
1
1
1
1
0
0
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
Colony Size
150
1
1
1
135
50
17.5
Parasite
Polyandry
Polygyny
Low
Relatedness
0
0
0
0
1
1
1
1
1
1
0
0
0
0
0
0
1
1
0
0
1
0
0
1
0
1
0
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
527.48
495.5
20
1
200
300000
100
5500
21
30
30
106
25
28
17
13
70
50
62
1
6.3
92500
6,000
50
155.5
40
200
400
1
1
1
1
1
1
350
1104
80
413
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
Species
Pseudomyrmex elongatus
Pseudomyrmex ferruginea
Pseudomyrmex gracilis
Pseudomyrmex holmgreni
Pseudomyrmex inquilinus
Pseudomyrmex nigrocinctus
Pseudomyrmex pallidus
Pseudomyrmex penetrator
Pseudomyrmex peperi
Pseudomyrmex schuppi
Pseudomyrmex simplex
Pseudomyrmex sp1
Pseudomyrmex sp2
Pseudomyrmex sp3
Pseudomyrmex viduus
Pseudopolybia compressa
Pseudopolybia difficilis
Pseudopolybia vespiceps
Psithyrus ashtoni
Psithyrus citrinus
Pteromalus puparum
Pteromalus venustus
Pteroptrix orientalis
Pterygophorus sp
Ptilothrix bombiformis
Ptilotrigona lurida
Pyramica alberti
Pyramica augustandrewi
Pyramica bauderi
Pyramica bunki
Pyramica clypeata
Pyramica creightoni
Pyramica deyrupi
Pyramica dietrichi
Pyramica dohertyi
Pyramica eggersi
Pyramica emmae
Pyramica gundlachi
Pyramica hexamera
Pyramica margaritae
Pyramica membranifera
Pyramica mutica
Pyramica ohioensis
Pyramica ornata
Pyramica pergandei
Pyramica pilinasis
Pyramica pulchella
Pyramica rostrata
Pyramica schulzi
Pyramica sp1
Pyramica sp2
Pyramica sp3
Pyramica subedentata
Pyramica talpa
Pyrobombus ardens
Pyrobombus edwardsii
Pyrobombus ephippiatus
Pyrobombus huntii
Pyrobombus impatiens
Pyrobombus melanopygus
Pyrobombus mixtus
Pyrobombus perplexus
Pyrobombus sitkensis
Pyrobombus ternarius
Pyrobombus vagans
Pyrobombus vosnesenskii
Recurvidris sp1
Rhopalum pygidiale
Rhopalum watanabei
Rhoptromyrmex melleus
Rhysipolis decorator
Rhytidoponera aciculata
Rhytidoponera araneoides
Haploid
Number
35.0
25.0
12.0
12.0
21.0
25.0
22.0
8.0
25.0
26.0
5.0
5.0
11.0
8.0
6.0
11.0
12.0
18.0
19.0
13.0
8.0
18.0
18.0
18.0
18.0
18.0
18.0
18.0
18.0
18.0
18.0
18.0
18.0
12.0
13.0
13.0
6.0
26.0
Eusocial
Colony Size
Parasite
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
1
0
1
1
80
461
93
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
30
Polyandry
Polygyny
Low
Relatedness
0
0
0
0
0
0
1
0
1
1
1
0
0
1
1
1
1
0
0
1
0
0
1
1
1
0
0
0
0
0
0
0
0
1
1
1
1
1
0
1
0
1
1
0
0
1
1
1
0
1
1
0
0
0
1
0
300
323
40
1
1
1
1
1
1
1
40
25.31
50
50
50
50
20
25
250
62
72
53
20
204
30
33
114
50
60
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
450
100
70
0
1
1
5000
1
50
414
0
0
0
Species
Rhytidoponera aurata
Rhytidoponera chalybaea
Rhytidoponera confusa
Rhytidoponera impressa
Rhytidoponera laciniosa
Rhytidoponera lamellinodis
Rhytidoponera maniae
Rhytidoponera mayri
Rhytidoponera mayri
Rhytidoponera metallica
Rhytidoponera punctata
Rhytidoponera purpurea
Rhytidoponera spANIC-10
Rhytidoponera spANIC-11
Rhytidoponera spANIC-13
Rhytidoponera spANIC-14
Rhytidoponera spANIC-15
Rhytidoponera spANIC-16
Rhytidoponera spANIC-9
Rhytidoponera tasmaniensis-1
Rhytidoponera tasmaniensis-2
Rhytidoponera victoriae
Rocalia japonica
Rocalia longipennis
Rocalia sp
Sapyga pumila
Scaptotrigona acantha
Scaptotrigona angustula
Scaptotrigona appendiculata
Scaptotrigona babai
Scaptotrigona barrocoloralensis
Scaptotrigona chanchamayoensis
Scaptotrigona clavipes
Scaptotrigona dentipes
Scaptotrigona depilis
Scaptotrigona esakii
Scaptotrigona fenestrata
Scaptotrigona fiebrigi
Scaptotrigona flavipes
Scaptotrigona fulgiduslongiplumosis
Scaptotrigona fulviventris
Scaptotrigona fuscipennis
Scaptotrigona japonica
Scaptotrigona latitarsis
Scaptotrigona megastigmata
Scaptotrigona minangkabou
Scaptotrigona muelleri
Scaptotrigona okinawana
Scaptotrigona pectoralis
barrocoloradensis
Scaptotrigona postica
Scaptotrigona quadripunctata
Scaptotrigona recursa
Scaptotrigona smaragdula
Scaptotrigona sp1
Scaptotrigona spinipes
Scaptotrigona subterranea
Scaptotrigona violacea
Scaptotrigona xanthotricha
Scaura latitarsis
Sceliphron caementarium
Separatobombus griseocollis
Sericomyrmex amabilis
Sericomyrmex urichi
Siobla ferox
Siobla metallica
Siobla ruficornis
Siobla sturmi
Siobla venustaapicalis
Sirex cyaneus
Sirex juvencus
Sirex noctilio
Solenopsis abdita
Haploid
Number
21.0
21.0
26.0
29.7
25.0
25.0
18.7
50.0
19.0
23.5
25.0
26.0
11.0
25.0
26.0
24.0
15.0
23.0
21.0
11.0
9.0
8.0
25.5
17.0
17.0
16.0
16.0
17.0
17.0
17.0
17.0
17.0
16.0
16.0
17.0
17.0
13.0
16.0
17.0
17.0
17.0
17.0
20.0
8.0
17.0
17.0
17.0
17.0
17.0
14.0
17.0
17.0
17.0
16.0
17.0
19.0
25.0
9.0
9.0
9.0
9.0
9.0
8.0
8.0
8.0
Eusocial
Colony Size
Parasite
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
361
271
387.5
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
0
1
1
1
0
0
0
0
0
0
0
0
1
27875
1650
125
577
555
Polyandry
Polygyny
Low
Relatedness
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
223.5
393
778.67
945.5
1
1
1
1
1
1
1
1
200
415
0
0
0
Species
Solenopsis aurea
Solenopsis carolinensis
Solenopsis CO
Solenopsis COR
Solenopsis fugax
Solenopsis GA
Solenopsis geminata
Solenopsis globularia
Solenopsis invicta
Solenopsis lou
Solenopsis MEK-005
Solenopsis MI
Solenopsis molesta
Solenopsis nickersoni
Solenopsis OR
Solenopsis papuana
Solenopsis pergandei
Solenopsis picta
Solenopsis PL
Solenopsis PO
Solenopsis richteri
Solenopsis saevissima
Solenopsis SO
Solenopsis sp1
Solenopsis sp2
Solenopsis spnrabdita
Solenopsis spnrcarolinensis
Solenopsis SS
Solenopsis tennesseensis
Solenopsis texana
Solenopsis tonsa
Solenopsis xyloni
Sphinctomyrmex cfsteinheili
Sphinctomyrmex steinheili
Stelis chlorocyanea
Stelopolybia areata
Stelopolybia multipicta
Stelopolybia pallipes_pallipes
Stelopolybia vicina
Stenamma alas
Stenamma brevicorne
Stenamma debile
Stenamma diecki
Stenamma expolitum
Stenamma felixi
Stenamma impar
Stenamma meridionale
Stenamma schmitti
Stenamma westwoodii
Stenichneumon culpator
Stenodynerus frauenfeldi
Stictiella serrata
Stigmacros spANIC-1
Stigmacros spANIC-3
Streblognathus aethiopicus
Streblognathus peetersi
Stromboceros koebelei
Strongylogaster blechni
Strongylogaster filicis
Strongylogaster lineata
Strongylogaster macula
Strongylogaster mixta
Strongylogaster moiwana
Strongylogaster onocleae
Strongylogaster osmundae
Strongylogaster ruber
Strongylogaster secunda
Strongylogaster tambensis
Strongylognathus afer
Strongylognathus huberi
Strongylognathus karawajewi
Strongylognathus minutus
Strongylognathus pisarskii
Haploid
Number
16.0
11.0
16.0
16.0
11.0
16.0
16.0
19.0
11.0
16.0
23.0
17.0
32.0
32.0
4.0
20.0
14.0
10.0
19.0
10.0
7.0
9.0
8.0
8.0
7.0
9.0
9.0
8.0
7.0
9.0
9.0
7.0
14.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
Colony Size
200
42
47
19
28000
50
152039.67
70
16
80
200
140
1000
10000
200
54
185
300000
555
200
200
184
200
200
240
Parasite
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Polyandry
Polygyny
Low
Relatedness
1
1
1
1
0
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
1
0
1
6875
5000
1000000
200
70
110
72
200
54
15
121
0
0
0
0
0
0
0
0
0
0
1
1
43.5
95
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
1
416
0
0
Species
Strongylognathus potanini
Strongylognathus testaceus
Strongylognathus tylonum
Strumigenys bajarii
Strumigenys br
Strumigenys doriae
Strumigenys elongata
Strumigenys emmae
Strumigenys friedae
Strumigenys frivaldszkyi
Strumigenys gl
Strumigenys godeffroyi
Strumigenys gundlach
Strumigenys lj
Strumigenys ljls
Strumigenys loriae
Strumigenys louisianae
Strumigenys mayri
Strumigenys mi
Strumigenys nf
Strumigenys rogeri
Strumigenys sp1
Strumigenys wk
Subterraneobombus appositus
Subterraneobombus borealis
Svastra obliquaexpurgata
Sycaonia sicaria
Syspasis alboguttata
Syspasis scutellator
Tapinoma erraticum
Tapinoma indicum
Tapinoma litorale
Tapinoma madeirense
Tapinoma melanocephalum
Tapinoma minutum
Tapinoma nigerrimum
Tapinoma ramulorum
Tapinoma sessile
Tapinoma simrothi
Tapinoma sp1
Taxonus alboscutellatus
Technomyrmex albipes
Technomyrmex sp1
Technomyrmex sp2
Technomyrmex sp2bicolor
Technomyrmex sp3
Telenomus fariai
Temnothorax acervorum
Temnothorax affinis
Temnothorax albipennis
Temnothorax allardycei
Temnothorax ambiguus
Temnothorax andrei
Temnothorax angustulus
Temnothorax bradleyi
Temnothorax cfinterrupus
Temnothorax cflichtensteini
Temnothorax congruus
Temnothorax corticalis
Temnothorax crassipilis
Temnothorax crassispinus
Temnothorax curvispinosus
Temnothorax duloticus
Temnothorax exilis
Temnothorax faberi
Temnothorax flavicornis
Temnothorax gredosi
Temnothorax interruptus
Temnothorax kraussei
Temnothorax lichtensteini
Temnothorax longispinosus
Temnothorax melas
Temnothorax minutissimus
Haploid
Number
11.0
12.0
21.0
16.0
16.0
21.0
11.0
11.0
11.0
8.0
6.0
5.0
9.0
8.0
9.0
5.0
9.0
8.5
15.0
14.0
14.0
15.0
10.0
9.0
11.0
10.0
23.0
12.0
14.0
9.0
10.5
17.5
23.0
24.0
11.0
15.5
11.0
17.0
12.0
14.0
14.0
12.0
13.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
0
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Colony Size
400
15
100
15
18
16
48
400
90
100
48
26
100
80
26
Parasite
Polyandry
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Polygyny
Low
Relatedness
1
1
0
0
0
0
0
0
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
0
1
0
0
0
1
0
1
1
1
1
1
20
500
300
65952
1
2912000
1
323
412
23
46
50
226
94.5
417
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
Species
Temnothorax nadigi
Temnothorax neomexicanus
Temnothorax niger
Temnothorax nigriceps
Temnothorax nitens
Temnothorax nylanderi
Temnothorax obliquicanthus
Temnothorax palustris
Temnothorax parvulus
Temnothorax pergandei
Temnothorax rabaudi
Temnothorax racovitzai
Temnothorax recedens
Temnothorax rottenbergii
Temnothorax rugatulus
Temnothorax salvini
Temnothorax schaumii
Temnothorax smithi
Temnothorax sordidulus
Temnothorax specularis
Temnothorax spinosior
Temnothorax spinosius
Temnothorax ssptuberumgroup
Temnothorax subditivus
Temnothorax texanus
Temnothorax tristis
Temnothorax tuberointerruptus
Temnothorax tuberum
Temnothorax unifasciatus
Tenthredo abdominalis
Tenthredo amoena
Tenthredo angustiannulata
Tenthredo arcuata
Tenthredo atra
Tenthredo basizonata
Tenthredo bipunctula
Tenthredo brevicornis
Tenthredo colon
Tenthredo contusa
Tenthredo convergenata
Tenthredo crassa
Tenthredo cylindrica
Tenthredo decens
Tenthredo dentina
Tenthredo fagi
Tenthredo ferruginea
Tenthredo finschiseguro
Tenthredo flavomandibulata
Tenthredo fortunei
Tenthredo fukaii
Tenthredo fulvaadusta
Tenthredo fuscoterminata
Tenthredo gifui
Tenthredo hilaris
Tenthredo hokkaidonis
Tenthredo japonica
Tenthredo jozana
Tenthredo latifasciata
Tenthredo limbata
Tenthredo livida
Tenthredo mandibularis
Tenthredo marginella
Tenthredo matsumurai
Tenthredo melanogastra
Tenthredo mesomelas
Tenthredo mortivaga
Tenthredo nigropicta
Tenthredo nitidiceps
Tenthredo notha
Tenthredo obsoleta
Tenthredo olivacea
Tenthredo omissa
Tenthredo opaciceps
Haploid
Number
18.0
9.0
11.0
14.0
9.0
21.0
12.0
11.0
13.5
9.0
11.0
17.0
12.0
16.0
12.0
21.0
9.0
8.5
10.0
18.0
8.0
12.5
10.0
9.0
10.0
19.5
10.0
10.0
10.0
10.0
10.0
10.0
10.0
13.0
11.0
10.0
10.0
12.0
10.0
10.0
11.0
12.0
9.0
10.0
10.0
9.0
10.0
10.0
10.0
10.0
11.0
19.0
10.0
10.0
10.0
10.0
10.0
19.0
10.0
10.0
10.0
10.0
Eusocial
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Colony Size
50
50
100
121
35
50
220
63
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
418
Parasite
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Polyandry
Polygyny
1
0
0
0
0
1
0
Low
Relatedness
1
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
1
1
0
1
1
0
0
1
1
1
0
1
0
Species
Tenthredo opposita
Tenthredo ornatula
Tenthredo picticornis
Tenthredo platycera
Tenthredo procincta
Tenthredo providens
Tenthredo pseudolivacea
Tenthredo rubrocaudata
Tenthredo solitaria
Tenthredo sp
Tenthredo sp1
Tenthredo sp2
Tenthredo sp3
Tenthredo sp4
Tenthredo sp5
Tenthredo sp6
Tenthredo subolivacea
Tenthredo takeuchii
Tenthredo temula
Tenthredo ussuriensis
Tenthredo velox
Tenthredo viridatrix
Tenthredo viridis
Tenthredopsis carinata
Tenthredopsis litterata
Tenthredopsis nassata
Tenthredopsis sp1
Tenthredopsis sp2
Tenthredopsis sp3
Terataner alluaudi
Terataner foreli
Tetragona angustula
Tetragona buchwaldi
Tetragona clavipes
Tetragona iridipennis
Tetramorium adelphon
Tetramorium brevidentatum
Tetramorium caespitum
Tetramorium caldarium
Tetramorium eleates
Tetramorium forte
Tetramorium guineense
Tetramorium impurum
Tetramorium kheperra
Tetramorium lanuginosum
Tetramorium pacificum
Tetramorium pnyxis
Tetramorium semilaeve
Tetramorium seneb
Tetramorium simillimum
Tetramorium smithi
Tetramorium sp1
Tetramorium sp10
Tetramorium sp11
Tetramorium sp12
Tetramorium sp2
Tetramorium sp3
Tetramorium sp4
Tetramorium sp5
Tetramorium sp6
Tetramorium sp7
Tetramorium sp8
Tetramorium sp9
Tetramorium spinosum
Tetramorium tsushimae
Tetramorium walshi
Tetraponera allaborans
Tetraponera sp1
Tetraponera sp2
Tetraponera sp3
Tetrastichus gigas
Tetrastichus megachilidis
Thrinax athyrii
Haploid
Number
9.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
9.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
10.0
12.0
8.0
9.0
9.0
8.0
8.0
8.0
17.0
17.0
11.0
10.0
14.0
14.0
14.0
11.0
7.0
7.0
11.0
10.0
7.0
10.0
7.0
13.0
10.0
18.0
10.0
12.0
9.0
10.0
9.0
10.0
11.0
9.0
13.0
7.0
13.0
7.0
16.0
22.0
21.0
6.0
6.0
6.0
Eusocial
Colony Size
Parasite
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
78
22
0
0
2152.5
6500
2550
6272
7316.43
300
70
3000000
6953
1
1
1
419
Polyandry
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Polygyny
Low
Relatedness
1
1
0
0
1
1
0
1
0
1
0
0
1
1
1
1
1
1
0
0
Species
Thrinax japonicus
Thrinax melanogyne
Thrinax minomensis
Thrinax paucipunctatus
Thrinax sasayamensis
Thrinax struthiopteridis
Thrinax tokunagai
Torymus baccharidis
Torymus californicus
Torymus capillaceus
Torymus koebelei
Torymus occidentalis
Torymus tubicola
Torymus umbilicatus
Torymus vesiculi
Torymus warreni
Trachusa gummifera
Trachymyrmex cornetzisp1
Trachymyrmex isthmicus
Trachymyrmex jamaicensis
Trachymyrmex opulentus
Trachymyrmex ruthae
Trachymyrmex septentrionales
Trachymyrmex sp1
Trachymyrmex sp2
Trachymyrmex zeteki
Trichiosoma lucorum
Trichogramma brassicae
Trichogramma chilonis
Trichogramma deion
Trichogramma dendrolimi
Trichogramma evanescens
Trichogramma nubialale
Trichogramma platneri
Trichogramma pretiosum
Trichogramma sp1
Trigona carbonaria
Trigona clypearis
Trigona corvina
Trigona fuscipennis
Trigona hockingsi
Trigona jaty
Trigona julianii
Trigona mellipes
Trigona silvestri
Trigona spinipes
Trigona staudingeri
Trigona subterranea
Trigonaspis megaptera
Trigonisca atomaria
Trigonisca buyssoni
Trybliographa bochei
Trychofoenus sp1
Trypoxylon albitarse
Trypoxylon asuncicola
Trypoxylon fabricator
Trypoxylon nitidum
Trypoxylon obsonator
Trypoxylon petiolatum
Trypoxylon politum
Trypoxylon sp1
Trypoxylon sp2
Tycherus australogeminus
Tycherus bellicornis
Tycherus dilleri
Tycherus fuscicornis
Tycherus ischiontelinus
Tycherus ophthalmicus
Tycherus osculator
Tycherus suspicax
Typhlomyrmex meire
Typhlomyrmex rogenhoferi
Urocerus augur
Haploid
Number
8.5
6.0
8.0
6.0
6.0
6.0
7.0
6.0
6.0
6.0
5.0
6.0
6.0
5.0
6.0
6.0
16.0
10.0
6.0
9.0
8.0
5.0
5.0
5.0
5.0
5.0
5.0
5.0
17.0
17.0
17.0
17.0
10.0
10.0
14.0
16.0
16.0
16.0
13.7
14.0
14.0
12.5
13.0
11.0
10.5
11.0
11.0
9.0
11.0
11.0
11.0
10.0
18.0
18.0
Eusocial
Colony Size
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
300
236
100
331
966.67
348.5
1
Parasite
Polyandry
Polygyny
Low
Relatedness
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
1
1
10000
500
7200
6500
3500
963
2000
500
63500
1
500
136
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
1
420
Species
Urocerus gigas
Venturia canescens
Vespa crabro
Vespa ducalis
Vespa mandarinia
Vespa simillima xanthoptera
Vespula flaviceps
Vespula germanica
Vespula maculifrons
Vespula pensylvanica
Vespula rufa
Vespula squamosa
Vespula vulgaris
Virgichneumon digrammus
Virgichneumon faunus
Vollenhovia brachycera
Vollenhovia emeryii
Vollenhovia nipponica
Vollenhovia sp1
Vollenhovia sp2
Vollenhovia sp3
Vollenhovia sp4
Vollenhovia spANIC-3
Vulgichneumon saturatorius
Wasmannia auropunctata
Xenomyrmex floridanus
Xestophanes potentillae
Xylocopa virginica krombeini
Haploid
Number
13.0
11.0
25.0
25.0
25.0
25.0
25.0
17.0
11.0
18.0
11.0
25.0
17.0
18.0
20.0
9.0
10.0
Eusocial
Colony Size
0
0
1
1
1
1
1
1
1
1
1
1
1
0
0
1
1
1
1
1
1
1
1
0
1
1
0
0
1
1
550
Parasite
61640
11793
1594
1385
1
1
150
1
571.33
50
1
421
0
0
1
0
0
0
0
0
0
0
Polyandry
Polygyny
Low
Relatedness
1
0
1
1
0
0
1
0
1
1
2
2
0
0
1
1
1
2
2
0
1
0
1
1
1
1
1
1
0
1
0
Appendix F
R code for ancestral condition test
422
This function is dependent on the R phytools package version 0.4-31. This function is
part of the package evobiR and may be installed directly from github.
# Installation
library(devtools)
install_github('coleoguy/evobir')
library(evobiR)
# Code for the Ancestral Condition Test
function (trees, data, derived.state, iterations = 10)
{
dt.vec <- data[, 3]
names(dt.vec) <- data[, 1]
ct.data <- data[data[, 3] != derived.state, ]
ct.vec <- as.numeric(ct.data[, 2])
names(ct.vec) <- ct.data[, 1]
anc.states.cont.trait <- list()
if (class(trees) == "multiPhylo") {
for (leslie in 1:length(trees)) {
cat(paste("Performing ASR of continuous character on tree:",
leslie, "\n"))
anc.states.cont.trait[[leslie]] <- anc.ML(trees[[leslie]],
ct.vec, model = "BM")
}
}
if (class(trees) == "phylo") {
anc.states.cont.trait[[1]] <- anc.ML(trees, ct.vec, model = "BM")
}
anc.state.dt <- list()
if (class(trees) == "multiPhylo") {
for (abbi in 1:length(trees)) {
temp.anc <- make.simmap(trees[[abbi]], dt.vec, model = "ARD",
nsim = 1, pi = c(1, 0), message = F)
anc.state.dt[[abbi]] <- temp.anc
}
}
if (class(trees) == "phylo") {
temp.anc <- make.simmap(trees, dt.vec, model = "ARD",
nsim = 1, pi = c(1, 0), message = F)
anc.state.dt[[1]] <- temp.anc
}
producing.nodes.list <- list()
if (class(trees) == "multiPhylo") {
for (i in 1:length(trees)) {
anc.state.bi <- anc.state.dt[[i]]
ss_nodes <- anc.state.bi$mapped.edge[, 1] > 0 & anc.state.bi$mapped.edge[,
2] > 0
423
wanted_branches <- ss_nodes[ss_nodes == T]
wanted_nodes <- names(wanted_branches)
wanted_nodes <- gsub(",.*", "", wanted_nodes)
producing.nodes.list[[i]] <- unique(wanted_nodes)
}
}
if (class(trees) == "phylo") {
anc.state.bi <- anc.state.dt[[1]]
ss_nodes <- anc.state.bi$mapped.edge[, 1] > 0 & anc.state.bi$mapped.edge[,
2] > 0
wanted_branches <- ss_nodes[ss_nodes == T]
wanted_nodes <- names(wanted_branches)
wanted_nodes <- gsub(",.*", "", wanted_nodes)
producing.nodes.list[[1]] <- unique(wanted_nodes)
}
hd.nodes <- vector()
if (class(trees) == "multiPhylo") {
for (i in 1:length(trees)) {
anc.states <- anc.states.cont.trait[[i]]
producing.nodes <- producing.nodes.list[[i]]
hd.nodes[i] <- mean(anc.states$ace[names(anc.states$ace) %in%
producing.nodes])
}
}
if (class(trees) == "phylo") {
anc.states <- anc.states.cont.trait[[1]]
producing.nodes <- producing.nodes.list[[1]]
hd.nodes[1] <- mean(anc.states$ace[names(anc.states$ace) %in%
producing.nodes])
}
hd.nodes <- mean(hd.nodes)
null.anc.nodes <- vector()
counter <- 1
number.of.trans <- vector()
if (class(trees) == "multiPhylo") {
for (i in 1:length(trees)) {
producing.nodes <- producing.nodes.list[[i]]
number.of.trans[i] <- length(producing.nodes)
anc.dt <- anc.state.dt[[i]]
anc.ct <- anc.states.cont.trait[[i]]
node.states <- describe.simmap(anc.dt)$states
anc.cond.nodes <- anc.ct$ace[names(anc.ct$ace) %in%
names(node.states)[node.states != derived.state]]
for (j in 1:iterations) {
null.anc.nodes[counter] <- mean(sample(anc.cond.nodes,
length(producing.nodes)))
counter <- counter + 1
424
}
}
}
if (class(trees) == "phylo") {
producing.nodes <- producing.nodes.list[[1]]
number.of.trans[1] <- length(producing.nodes)
anc.dt <- anc.state.dt[[1]]
anc.ct <- anc.states.cont.trait[[1]]
node.states <- describe.simmap(anc.dt)$states
anc.cond.nodes <- anc.ct$ace[names(anc.ct$ace) %in%
names(node.states)[node.states !=
derived.state]]
for (j in 1:iterations) {
null.anc.nodes[counter] <- mean(sample(anc.cond.nodes,
length(producing.nodes)))
counter <- counter + 1
}
}
plot(density(null.anc.nodes), main = "", xlab = colnames(data)[2],
lwd = 2)
max.y <- range(density(null.anc.nodes)[2])[2]
lines(x = c(hd.nodes, hd.nodes), y = c(0, max.y), col = "red",
lty = 1, lwd = 2)
text(17, 0.4, label = paste("p-value", sum(null.anc.nodes <
hd.nodes)/length(null.anc.nodes)))
bigger <- (sum(null.anc.nodes >= hd.nodes)/length(null.anc.nodes)) *
100
smaller <- (sum(null.anc.nodes <= hd.nodes)/length(null.anc.nodes)) *
100
cat(paste("Derived State Mean Ancestral Cond:", hd.nodes,
"\n"))
cat(paste("Number of producing nodes:", mean(number.of.trans),
"\n"))
cat(paste("Mean of null dist:", mean(null.anc.nodes), "\n"))
cat(paste("SD of null dist:", sd(null.anc.nodes), "\n"))
cat(paste(bigger), "% of simulations had a derived state which arrose\n",
"with a mean continuous value larger than inferred for the observed derived
state\n")
cat(paste(smaller), "% of simulations had a derived state which arrose\n",
"with a mean continuous value smaller than inferred for the observed derived
state\n")
results <- list()
results[[1]] <- hd.nodes
results[[2]] <- number.of.trans
results[[3]] <- null.anc.nodes
results[[4]] <- bigger
results[[5]] <- smaller
425
names(results) <- c("OriginatingNodes", "NTrans", "NullDist",
"bigger", "smaller")
return(results)
}
# Code for the creation of simulated data
# Sim Functions
#simulate trees
library(geiger)
library(phytools)
make.trees <- function(taxa=200, iterations=100, b=1, d=.5){
trees <- list()
for(i in 1:iterations){
set.seed(i)
trees[[i]] <- drop.extinct(sim.bdtree(b=b, d=d, stop="taxa", n=taxa, extinct=F))
}
class(trees) <- "multiPhylo"
return(trees)
}
sim.cont <- function(trees, iterations=100, rate=.2){
cont.traits <- list()
for(i in 1:iterations){
set.seed(i)
foo <- sim.char(trees[[i]], par=rate)
foo2 <- as.vector(foo)
names(foo2) <- row.names(foo)
cont.traits[[i]] <- foo2
}
return(cont.traits)
}
stretch.trees <- function(trees, cont.trait, low.quar=1, high.quar=1, iterations = 100){
#now we will scale branches based on their
#reconstructed continuous trait value
stretched.trees <- list()
for(i in 1:iterations){
x <- cont.trait[[i]]
node.vals <- fastAnc(trees[[i]], x)
names(x) <- 1:length(x)
node.vals <- c(x, node.vals)
current.tree <- trees[[i]]
lookup <- current.tree[[1]]
scale.factor <- vector()
for(j in 1:nrow(lookup)){
scale.factor[j] <- mean(c(node.vals[lookup[j, 1]], node.vals[lookup[j, 2]]))
426
}
low <- summary(scale.factor)[[2]]
high <- summary(scale.factor)[[5]]
for(j in 1:length(scale.factor)){
if(scale.factor[j] >= high){
scale.factor[j] <- high.quar
}else if(scale.factor[j] <= low){
scale.factor[j] <- low.quar
}else{
scale.factor[j] <- 1
}
}
current.tree[[4]] <- current.tree[[4]] * scale.factor
stretched.trees[[i]] <- current.tree
}
return(stretched.trees)
}
sim.disc <- function(trees, q=list(rbind(c(-.02, .02), c(0, 0)))){
disc.cor <- list()
test <- F
while(test == F){
foo <- sim.char(trees, par=q, model="discrete", root=1)
if(sum(foo == 1) > sum(foo == 2) & sum(foo == 2) > 1){
test <- T
}
}
foo2 <- as.vector(foo)
names(foo2) <- row.names(foo)
disc.cor <- foo2
return(disc.cor)
}
.
427
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Biographical Information
Heath Blackmon has been an amateur naturalist and beetle collector since
childhood. He graduated summa cum laude with a Bachelors of Science degree in
Environmental Science with a concentration in fisheries and wildlife biology from Oregon
State University in 2010. He continued his education working with Dr. Jeffery P. Demuth
at the University of Texas at Arlington earning a PhD in Quantitative Biology in 2015.
Heath has been recognized for his research and teaching, receiving a NESCent graduate
fellowship, the Carrizo Oil and Gas doctoral fellowship, and the T.E. Kennerly Excellence
in Teaching award.
Heath’s research interests are centered on building an understanding of the
relationships between phenotypic traits and genomic traits and using comparative
approaches to bridge the gap between pattern and process. He will be leaving the
University of Texas at Arlington for a postdoctoral position in the lab of Dr. Emma
Goldberg at the University of Minnesota and plans on pursuing a career in academia.
447
Heath Blackmon