Professional Documents
Culture Documents
No. 25
November 2012
CONTENTS
Editorial:
Inventories - What are they good for? 3
Dr. William P. Coffman: Celebrating 50
years of research on Chironomidae
Dear Sepp!
14
Current Research
Sharma, S. et al. Chironomidae (Diptera)
in the Himalayan Lakes - A study of subfossil assemblages in the sediments of
two high altitude lakes from Nepal 15
Krosch, M. et al. Non-destructive DNA
extraction from Chironomidae, including
fragile pupal exuviae, extends analysable
collections and enhances vouchering 22
Martin, J. Kiefferulus barbitarsis (Kieffer,
1911) and Kiefferulus tainanus (Kieffer,
1912) are distinct species
28
Short Communications
An easy to make and simple designed
rearing apparatus for Chironomidae 33
Some proposed emendations to larval
morphology terminology
35
Chironomids in Quaternary permafrost
deposits in the Siberian Arctic
39
New books, resources and
announcements
43
Finnish Chironomidae
47
Chironomini indet. (Paratendipes?) from La Selva Biological Station, Costa Rica. Photo by Carlos de la Rosa.
Front page layout: Chironomid in title from photograph by Steve Marshall, Graphic design by Kolbjrn Skarpnes,
NTNU Information Division.
Front page photo: Chironomini indet., possibly Paratendipes?
Photo by Carlos de la Rosa.
Editorial
Inventories What are they good for?
Do we have the same interpretation of species? Do we use the same literature? Can we trust any
chironomid species identification if it is not documented, either through description, reference
literature or reference material? - In my opinion the answers are no.
We have all experienced that re-identification of a midge sometimes results in a species or species group different from the one the specimen originally was identified as. It could be our own
original identification or it could be someone elses. The fact that many Chironomidae species are
difficult to identify perhaps makes us experience this more often in our group, I dont know, but
it is clearly a problem for static inventories and check-lists.
Moreover, many if not most identifications made in faunistic and ecological studies are unreferenced in publications. Thus, there is no possibility for the reader to know what literature was used,
if reference specimens were examined, or even if the material has been deposited in a publicly
accessible collection. How can the reader know if the author(s) used the most recent and geographically relevant taxonomic revisions and identification keys? It definitely makes a difference
in the interpretation of a species identification (e.g. Tanytarsus mendax) if it is clear that the latest clarification of its taxonomy (Spies 1998) is used and not an older revision (Reiss & Fittkau
1971). So apart from giving credit where it belongs, citing the taxonomic literature is scientifically important and should be as mandatory as any other methodological reference.
The increased use of molecular markers in species identification, discovery and delineation (i.e.
DNA barcoding) provides an additional set of objective characters for species in Chironomidae.
The Barcode of Life Data Systems database (BOLD) currently holds almost 16 000 sequences of
more than 1000 named species and offer a tool to unify the taxonomy and interpretation of chironomid species world-wide. Through our work with DNA barcoding of Chironomidae in Trondheim, we have discovered numerous cryptic species and molecular data has helped us understand
species boundaries as well as phylogenetic relationships. There is no doubt that this is part of the
future for chironomid taxonomy and thus also disciplines that rely on solid taxonomic foundations for identifications and analysis of species distributions and ecology.
To avoid the creation of useless inventories and erroneous check-lists we should all be careful
in how we use and document our identification procedures. It doesnt matter if they are based
on morphology or molecules. If the results are not made repeatable by proper description of the
methods, it is not science. Therefore, cite your methods and references, and deposit your specimens in scientific collections!
Torbjrn Ekrem
Museum of Natural History and Archaeology, Norwegian University of Science and Technology, NO-7491
Trondheim, Norway. E-mail: torbjorn.ekrem@ntnu.no
References
Reiss, F. and Fittkau, E.J. 1971. Taxonomie und kologie europisch verbreiteter Tanytarsus-Arten (Chironomidae, Diptera). - Archiv fr Hydrobiologie Supplement: 40: 75-200.
Spies, M. 1998. Resolution of Tanytarsus mendax Kieffer, 1925, T. dibranchius Kieffer in Zavrel, 1926, and
associated names (Diptera: Chironomidae). - Studia dipterologica 5: 195-210.
His work has delved into methodologies and defining characters, through ecology and biogeography, taxonomy and evolution. His insights
into the Chironomidae family species richness and
diversity, evolutionary patterns, extinction rates,
biogeographical puzzles and adaptation have been
profound and are likely to shape the directions in
this field of research in the future. His collection
of world-wide pupal exuviae is unparalleled and
extensive, with over 1,000 undescribed species,
many from the Neotropics. But perhaps his greatest contribution so far is a family of scientists and
researchers that have been touched, bitten perhaps,
by the bug of knowing and using this family to
resolve long-standing questions of evolution, biogeography, taxonomy and conservation.
ecosystems, supporting and maintaining the energy flow of most freshwater ecosystems. And their
diversity is barely known in much of the world.
Thousands of species remain to be discovered
and described, up to 20,000 estimated around the
world, including perhaps the most puzzling and
difficult fauna of all: terrestrial chironomids. In
spite of the challenges, the accumulated knowledge that a handful of specialists like Bill Coffman
have bequeathed us will continue to build through
the continued work on his and other collections.
It is only unfortunate that chironomid taxonomy,
as a field of study, has not been pursued or supported by most world universities or institutions.
Very few true specialists continue to work on the
family and the new generation is not coming up
in the needed numbers, at a time when the scientific world needs them the most. The current wave
of species extinction, mostly through habitat destruction and compounded by climate change, will
make the techniques and knowledge application of
this fauna more important than ever.
Sam Roback, the Dean of North American chironomid workers, and Ken Cummins and is wife
Peggy Wilzbach, who collected and shared many
specimens from their trips around the world. Scientists like Jim Sedell, Norman Anderson, Henri
Laville, Jan Sykora and former students like Rick
Jacobson, Lou Yurasits, Len Ferrington, Dave Wartinbee, Bill Walker, Roger Carrillo and I (among
others) benefited and made careers from our close
relationship with Bill, providing specimens, insights, writing papers and developing theories that
together represent the accumulated knowledge of a
productive career. Our gratitude cannot be expressed with simple words, but it is immortalized by
our joint accomplishments, the papers we wrote,
the courses we taught, the species we discovered
and the memories we shared. Thank you, Bill, for
all that you have done for all of us.
Coffman, W.P., and C.L. de la Rosa, 1998. Taxonomic composition and temporal organization of
tropical and temperate species assemblages of
lotic Chironomidae . - J. Kansas Entomol. Soc.
71: 388-406.
Coffman, W.P., 1983. Thoracic chaetotaxy of chironomid pupae. - Mem. Amer. Ent. Soc. 34:6170.
Swegman, B. and W.P. Coffman, 1980. Stenopsyche kodaikanalensis: a new species of Stenopsyche from South India. - Aquatic Insects 2:
73-79.
Coffman, W.P., 1979. Neglected characters of pupae in chironomid taxonomy and phylogeny. Ent. Scand. Suppl. 19: 37-46.
Coffman, W.P., 1978. Chironomids. In: Methods
for the Assessment and Prediction of Mineral
Mining Impacts on Aquatic Communities: A
Review and Analysis. W. T. mason, Jr. (ed.)
U.S. Dept. of Interior. Fish and Wildlife Service, Biological Services Program.
Cofmman, W.P., K.W. Cummins and J.C. Wuycheck, 1971. Energy flow in a woodland stream
ecosystem: I. Tissue support tropic structure of
the autumnal community. - Arch. Hydrobiol.
68: 292-276.
Dear Sepp!
Wolfgang Wlker:
In 1968, you and I were together at the International Congress of Entomology in Moscow. Both
of us were selected to take part in a post-congress
excursion to Borok on the river Volga. Our Russian colleagues still acted with some reservation
towards the ugly Germans in Moscow, but in Borok they were thawing remarkably and we had nice
evenings with various local families. We both were
directed to stay in a small house together with a
nice young lady as our supervisor (most probably
the daughter of a mighty Russian). In the beginning, she wanted to take a bath and was not sure
whether or not this was ok in the presence of two
wild Germans in the same house. I never forget
you saying we are very shy persons which indeed we were - and so the bathing could go on!
Elisabeth Stur:
Sepp became my Doktorvater in 1993, right after
he retired as director of the ZSM. He had been persuaded by Ulrike Nolte to take one last PhD-student to work with chironomids in Brazil. Well, as
it turned out, I was not his last PhD student, he
could not say no to supervise the remaining last
students, my Brazilian friends and colleagues Angela, Sofia and Maria. Sepp and his beloved wife
Elise has been part of my life since then, and I am
very grateful for all their support.
Elise and Sepp at the train station in Icking, Germany, July 2011. Photo: Angela Sanseverino.
Joachim Illies and Ernst Josef Fittkau on their way to collect at the Wasserkuppe, 1948. Photo: Family archive.
I have many fond memories of Sepp. As supervisor he was very patient in explaining morphological structures and teaching how to draw, and he
was always encouraging in the writing process.
However, his care for students and colleagues beyond the professional activity was perhaps even
more remarkable no matter what background the
person might have.
One memory I have is from a warm sunny February-day in Munich when I could convince him
to leave the office for the benefit of some hours in
the nearby Biergarten. It was such a fine day and
we had a very joyful conversation of bygone days;
including stories from his time with Thienemann,
Brundin and many other chironomologists whom
I only knew from literature. I still remember this
day with affection.
Angela Sanseverino:
The lines are too straight; your drawing needs
more movement and life. It was a spring morning
in 2000, and with those words Sepp introduced me
the art of taxonomy. Immediately he took his pen
and ink, and showed me how to give to a seta a
10
Sofia Wiedenbrug:
I was lucky to be Sepps PhD student and there
are many stories that I could tell. He impressed me
quite a lot right from the beginning! I had brought
many samples from Brazil to Germany, but in Munich I realized that this material was insufficient
to work on my PhD and I was so desolated about
that! So, Sepp told me, he was invited to the I
Encontro Brasileiro sobre Chironomidae in Rio
and that I should attend this meeting, too. He offered himself to go with me to south Brazil in order
to do some sampling after the Congress. At that
Maria C Messias:
I was introduced to Sepp by my adviser Sebastio
de Oliveira, in Instituto Oswaldo Cruz , Rio de
Janeiro, 1995. Both had met after a period of 20
years, when Sepp and Oliveira celebrated this occasion for the peer-reviewing of my Ph.D. thesis
proposal to Instituto Oswaldo Cruz and Zoologis-
11
Roland Gerstmeier:
che Staatssammlung, Germany. Later, several times I heard Sepp encourage me to advance the research work telling me go on, Maria, the work is
going ok, a big master does not fall from heaven.
What I learned from him was not only on Chironomidae, but personal values such as happiness of
living, to share and celebrate friendship. A master
might not fall from heaven, but a master like Sepp
stimulated us to grow with him, to dream and work
hard, so that we can achieve our dreams, heaven
and perfection.
Martin Spies:
In my early 1980s undergraduate years long before I succumbed to the siren song of the midges
(you never ordered or coaxed me into this group)
on the way to a field course you told me some
things about your work on chironomids. Suddenly
you burst into a desperate cry (like I never heard
from you again): That [personal name omitted
here] is destroying my [taxon name omitted here]
systematics! In the years to follow, you proceeded
to collaborate closely with that same person to produce several standard references for everyone studying the taxon in question. You tried hard not to
let differences in professional opinion affect personal relations, and in the interests of the latter you
were ready to make compromises in the former.
Claus Orendt:
I remember Sepp as a master of very dry humour
or stories, which seemed unbelieveable to students
of today. One example of each:
When we were on our first field campaign for my
diploma thesis, Sepp accompanied me on our boat
at Lake Chiemsee hauling some corers for sediment
samples on a beautiful and warm day of spring.
Everything was great: the wheather, the sampling
site and landscape, and in the background the gorgious scenery of the nearer mountains of the Alps.
We all were really impressed by this paradise-like
surrounding, and when we thankfully realized and
expressed this with words at the same moment, he
concluded: Really, you should pay an extra charge to be allowed to do this work here.
12
References
13
Spies, M. 2002. Professor Ernst Josef Fittkau 75 years, 50 years for chironomid research.
- Chironomus Newsletter on Chironomidae Research 15: 2-13.
Sanseverino, A., Stur, E., Wagensonner, I., Gerber, J., Messias, M.C., Spies, M., Reiff, N.,
Ospina Torres, R., Wiedenbrug, S., Ekrem, T.,
Riss, W. 2007. Sepp has turned 80. - Chironomus Newsletter on Chironomidae Research 20:
7-8.
Marta MargreiterKownacka was born on 14 September 1939 in Krakw into the family of the renowned Polish archaeologist, Professor Stanislaus
Buratyski. After graduating from high school in
1956 Marta was accepted into the Faculty of Biology and Earth Sciences at the Jagiellonian University in Krakow to study biology. In October
1961 Marta was employed by the Department of
Water Biology, Polish Academy of Sciences and
embarked on a decade of important research. She
joined the team that was conducting a comprehensive study of high-mountain streams, lakes and
small reservoirs, mainly of the Tatra mountains,
and comparing the results with other mountain
14
Current Research
Chironomidae (Diptera) in the Himalayan Lakes A study of sub-fossil
assemblages in the sediments of two high altitude lakes from Nepal
Subodh Sharma1, Susan E. Gresens2 and Berthold Janecek3
Aquatic Ecology Centre, School of Science, Kathmandu University, Nepal
E-mail: subodh.sharma@ku.edu.np
2
Department of Biological Sciences, Towson University, MD, USA
3
University of Natural Resources and Applied Life Sciences (BOKU), Vienna, Austria
1
Himalayan glaciers in the face of climate warming (Bolch et al. 2012). Shrinkage of many of
these glaciers affects the discharge of rivers and
lakes in the region. Palaeolimnological techniques
present the possibility to infer change in environmental conditions over time, such as lake water
temperature (Walker et al. 1991) using sub-fossil
chironomid head capsules preserved in lake sediments. However, climate reconstruction modeling
requires a calibration dataset based on samples of
chironomid head capsules from a large number of
lakes spanning a gradient of average temperature.
The strong association between lake temperature
and lentic chironomid distribution has been wellestablished by numerous calibration datasets from
Western Europe and the Nearctic (Eggermont and
Heiri 2011). A calibration dataset for Nepalese
lakes has not yet been constructed: this study represents a necessary first step towards establishing
the relationship between chironomid assemblages
and temperature for Himalayan lakes by sampling two remote very high altitude lakes. There
is hardly any information available on chironomid
assemblages and temperature for Himalayan lakes
and this study is therefore a necessary first step,
based on which several other high altitude lakes
are planned to be studied in the future.
Abstract
Chironomid head capsules were identified from
sediment cores taken from Lake Gokyo (4750 m)
and Lake Gosaikunda (4350 m) in the east-central
Himalaya, to determine taxonomic composition of
chironomid assemblages over time. The sedimentation rate of Lake Gosaikunda was estimated at
0.05 cm/yr and that of Gokyo was estimated at 0.07
cm/yr by 210Pb and 137Cs dating. Micropsectra sp.
was dominant in the sediments of both lakes. Lake
Gokyo sediments also contained Pseudodiamesa
sp., Eukiefferiella/Tvetenia sp., Orthocladius/Cricotopus sp. and Rheocricotopus sp. The concentration of headcapsules was nearly an order of magnitude greater in the Lake Gosaikunda core, which
contained mostly Micropsectra sp. and Pseudodiamesa sp. These taxa are typical of cold oligotrophic lakes. Differences in lake depth, dissolved
minerals, plus epi- and hypolimnetic temperature
and dissolved oxygen suggest that stratification
and temperature-induced increases in primary
production may affect chironomid assemblages
in these pristine lakes. Palaeolimnological studies
of Himalayan lakes should include replicate cores
within lake, to increase headcapsule sample sizes
given potentially high rates of sedimentation from
glacial runoff.
Introduction
High-altitude lakes are considered to be very sensitive to climate change (Parry et al. 2007). Climate change is an especially pressing environmental issue for a country like Nepal with its location
in the Central Himalaya. The normal seasonal pattern of monsoon rainfall appears to be changing,
but there is little data on variation in the form and
amount of precipitation across the Himalaya, and
consequently much uncertainty about the fate of
15
ible impact of nutrient enrichment and eutrophication. In contrast, the water chemistry of high-elevation lakes is mainly determined by weathering
of rocks (Tartari et al. 1998); similar high altitude
lakes in Sikkim and Kashmir (Zutshi1991; Khan
and Zutshi 1980; Sharma and Pant 1979) exhibited
very low concentrations of dissolved nutrients and
minerals and also low phytoplankton abundance.
Study sites
Two lakes in the main Himalayan range were
studied: Gokyo (Third Lake) which lies in the
eastern zone of Everest National Park, and Lake
Gosaikunda, lying in the central zone of Langtang
National Park (Figure 1, Table 1). Lake Gokyo
is fed by drainage from Ngozumba glacier and
springs from Renjo La Pass to the north-west.
Although there are four other major lakes in the
headwaters of Kosi River system, they have no
surface-water connection to Lake Gokyo. Lake
Gosaikunda is fed by snowmelt and by a perennial
spring source to the north-east, considered a holy
place by Hindus and Buddhists. There are many
lakes in the series connecting Lake Gosaikunda
with the River Trisuli in the Gandaki river system.
The geology surrounding the lake consists mostly
of exposed bedrock and glacier debris (Bortolami
1998). The lake bottom is composed of silt-clay
sediment in the deeper water and sand-silt in the
littoral zone. The average minimum temperature in
the Everest region of these high-altitude lakes is
-7.7oC in January, and the average maximum temperature is 16.2oC in August (Tartari et al. 1998).
Vegetation in the catchment is sparse but consists
of juniper, rhododendron and herbs.
Bathymetric maps of the lakes were compiled using GIS software from sounding data collected us-
SedimentcoreLakeGokyo
SedimentcoreLakeGosaikunda
Figure 1. Location map and photos of the study area (not in scale). Lake Gokyo (Mt. Everest National Park) is shown
tothe right and Lake Gosaikunda (Langtang National Park) to the left.
16
Name of lake
Gokyo
Gosaikunda
Coordinates
8641N
285N
2757' E
8525 E
Elevation
4750 m
4426 m
Surface area
42.9 ha
13.8 ha
Max. depth
42.0 m
24.0 m
Temp (top/bottom)
8.0/4.5 C
9.2/7.5 C
DO (top/bottom)
8.3/6.3 mg/L
6.1/3.6 mg/L
Na
0.37 mg/L
1.93 mg/L
0.26 mg/L
1.80 mg/L
Fe
0.78 mg/L
0.80 mg/L
Figure 2. Bathymetric map of Lake Gokyo (left) and Lake Gosaikunda (right). Stars indicate sites where the sediment
cores were taken.
Results
thousands of years, they are usually lacking delicate morphological features; yet they can be identified to genus level, or at least grouped into consistent morphotypes. Extraction of head capsules
from sediment samples followed the methods of
Walker (2001) for air dried sediment samples, but
with slight modifications. Samples were deflocculated in 10% KOH overnight (up to 18 hrs) without
heating. The sediment was then passed through a
100m mesh sieve and chironomid head capsules
were removed by a fine probe under a stereomicroscope at 40. After dehydration in 95% ethanol, head capsules were slide-mounted in Euparal.
Identifications were made at 400, based on the
keys of Wiederholm (1983), Roback and Coffmann (1987), Walker (2007), and Brooks et al.
(2007). Head capsules with a complete mentum
or two corresponding halves were counted as one
individual. Slides are retained at the Aquatic Ecology Centre of the School of Science, Kathmandu
University.
17
"
"
!
"
#
!
"
#
! " #
Figure 3. Head capsules count in two different cores from Lake Gokyo (left), and single core in Lake Gosaikunda
(right).
Figure 4. Chironomid stratiography of Lake Gokyo in core 1 (left), and core 2 (right). Chironomid taxa are given as %
abundance; sum represents the total number of head capsules.
Figure 5. Chironomid stratiography of Lake Gosaikunda. Taxa are given as % abundance and sum represents
the total number of head capsules.
18
Table 2. Relative abundance (in %) of chironomid taxa recorded in the lake sediments of both the lakes.
Taxon
Eukiefferiella/Tvetenia sp.
Lake Gokyo
Lake Gosaikunda
core 1 (6 cm)
(20 cm)
25
Micropsectra sp.
38
53
58
18
37
Orthocladius sp.
Pseudodiamesa sp.
14
41
Rheocricotopus sp.
The low and variable concentration of head capsules, especially in the Lake Gokyo cores, creates uncertainty in interpretation of the shifts in
chironomid composition observed. This variation may reflect changes in silt load from glacial
melt-water. In general, it is more difficult to infer
recent temperature change, because variation in
local environmental factors masks our ability to
detect small temperature changes of only a couple of degrees (Velle et al. 2010, Velle et al. 2012,
Eggermont & Heiri 2012). Lake morphometry
influences variation in sediment cores, due to eco-
19
Acknowledgements
20
Swar, D. B. 1980. Present status of limnological studies and research in Nepal. In: Mori, S
& I, Ikusima (Edited). Proc. First Workshop
on Promotion of Limnology in Developing
CountriesXXI SIL Congress, Kyoto, Japan.
Pp 43-47.
Velle, G., Brodersen, K.P., Birks, H.J.B., Willassen, E. 2010. Midges as quantitative temperature indicator species: Lessons for palaeoecology. - The Holocene 20: 989-1002.
Reiss, F. 1968. Neue Chironomiden-Arten (Diptera) aus Nepal. - Khumbu Himal 3 (1): 55-73.
Roback, S. S. and Coffmann W. P. 1987. Results
of the Nepal Alpine Zone Research Project,
Chironomidae (Diptera). - Proceedings of the
Acadademy of Natural Sciences Philadelphia
139: 87-158.
Wiederholm, Y., 1983. Chironomidae of the Holarctic region. Part 1, Larvae. - Entomologica
Scandinavica Supplement 19, 457 pp.
Sharma, A.P. and Pant M.C. 1979. Certain physico-chemical features, chlorophyll concentration and phytoplankton population in high altitude lake. - Tropical Ecology 20:101-113.
21
Introduction
An answer to this mismatch in demand and taxonomic expertise may lie with the actual and potential use of molecular data. In the past two decades
researchers have used data derived from DNA to
estimate phylogenies for their inherent interest
(e.g. Guryev et al. 2001), in supplementing morphological characters (Cranston et al. 2012), in estimation of support for biogeographic hypotheses
(e.g. Krosch et al. 2009; Cranston et al. 2010) and
22
proved in our protocols. Largely in response to requests from interested colleagues, we describe here
the simple technique that makes possible retrieval
of PCR-quality DNA from chironomids including
even from single pupal exuviae. Previous attempts
to extract DNA from exuviae using a modified salting out protocol (Miller et al. 1988) failed, and we
assumed such specimens lacked sufficient remaining tissue post-adult emergence for DNA extraction. However with careful handling of specimens
and more sensitive modern extraction techniques
described below, good DNA can be recovered,
thereby allowing access to greater numbers of immature stages, and avoiding the need for laborious
and often unrewarding rearing, making molecular
associations between life stages simpler.
Methods
Laboratory
Field
We extract total genomic DNA from specimens using a Qiagen DNeasy Blood and Tissue Kit (Qiagen, Hilden, Germany) following manufacturers
protocol. Modifications to this protocol were as
follows: (1) tissue was digested with proteinase K
overnight at 37oC; (2) after digestion, cuticles were
removed carefully using fine-tipped forceps and
transferred to 90% isopropanol before vouchering
as slides in Euparal; (3) the final elution volume
was 100 L to increase the final DNA concentration of the eluate; and (4) the incubation time for
elution was increased to 5 minutes. Furthermore,
digestions were never vortexed while cuticles remained in solution to avoid damage to diagnostic
morphological characters. At all stages, care with
handling is essential and we found using the solutions surface tension, rather than direct pressure
from forceps, to be sufficient for transferring specimens between micro-centrifuge tubes. We anticipate that extraction kits from other manufacturers
may produce similar results under the same modification, but this remains untested currently.
We use all traditional and some less usual sampling when collecting material in the field, according to the purpose. For population genetics studies
we sample intensively in a narrowly defined area
containing the appropriate microhabitat(s), seeking known target taxa. For phylogenetic studies
in which we seek also to maximise diversity, we
sample all visible microhabitats. At all lotic sites
we place a drift net downstream of other collecting activity for the duration of the visit, or longer,
including overnight if feasible, for pupal including exuvial sampling. In contrast to most studies,
we sort specimens from the substrate at or close to
the sampling location using dissector microscopes.
Live sorting is preferable with specimens being
picked with fine forceps and placed immediately
into isopropanol or molecular grade ethanol, and
never into formalin, methanol or ethanol that has in
any way been denatured. Only if a sample cannot
be sorted live in timely manner do we preserve
in alcohol, after removal of larger particulate matter and sieving for medium-sized organic removal.
Isopropanol, although more expensive, is widely
available for use in molecular labs, preserves DNA
very well, is less volatile and less flammable (important if flights are involved in field work) and
is not hygroscopic your 100% alcohol remains
absolute. Most failures to extract DNA from
specimens are associated with the age and storage
of older material (which should be kept cold and
dark) or unwitting use of certain denatured ethanols as supplied to many wet labs. We suspect
that some total DNA destruction is caused by customs irradiation of international mailed material
but cannot verify this.
23
A. Pharate adult pupa of Stictocladius lacuniferus Freeman, from New Zealand. Note the leg sheaths are all extended
and none recurved under wing sheath. B. Pupal exuviae: abdomen of Stictocladius uniserialis Freeman, from Australia. The genus Stictocladius, following morphological review by Cranston & Sther (2010) and Sther & Cranston
(2012) is now the subject of molecular study by Matt Krosch and Peter Cranston. The pupal stage not only provides
good morphological features, but has provided valuable DNA even from exuviae, as outlined in the article.
Vouchering specimens
In the early days, the few people working with
chironomid DNA had a rather casual approach to
vouchering material needed to verify that sequenc-
24
Results
This technique, which already has worked well
across other life stages from many subfamilies of
Chironomidae, was trialed on exuvial specimens
across a range of Orthocladiinae taxa. Of 58 exuvial specimens derived from separate field collections made at various sites across Australia during
2011 and 2012, extracted as described above, 27
have been successfully amplified and sequenced
for COI and 17 for 28S. Sequence quality always
was acceptable under standard QC metrics (e.g.
signal strength) applied by the sequencing software
and sequencing success was repeatable via both
multiple attempts using the same purified PCR
product and multiple initial PCR reactions. Moreover, vouchers made from cuticles post-digestion
showed that diagnostic characters remained intact
and morphology - and molecular-based identifications converged for all specimens. A number of
specimens, in addition to those successfully sequenced for either locus, amplified successfully
(as assessed using 1.5% agarose check gels) but
did not produce clean or useable sequence. This
suggests that further optimisation of protocols may
be necessary, especially with regard to the ratio of
DNA template to primer in sequencing reactions.
Discussion
The addition of PCR-quality DNA sequences from
wild caught pupal exuviae described here expands the number of sequenceable specimens for
chironomid researchers who collect in aquatic environments. The greatest benefit of this technique
will likely be in aiding the development of molecular associations between life stages of species
by negating the need for rearing of larvae through
to adults. This technique also has potential importance for those investigating population genetic
structure, gene flow and dispersal where sample
sizes must be maximised from each study site. We
encourage other researchers to retain, and attempt
to extract DNA from exuviae and not consider
these important specimens valueless for molecular
purposes.
However the addition of a new life history
stage, the pupal exuviae, to the repertoire of
those engaged in molecular studies of Chironomidae should not obscure that even this suite of
techniques do not provide the perfect answer to
all questions. We note that there are sporadic and
difficult to diagnose difficulties with extraction
for some specimens or collection conditions in
addition to the unknown age and history of collected exuviae, or interceptions of larvae, pupae
and adults by netting. There are sporadic problems
with amplification of DNA from mermithids rather
than the midge host, apparently restricted to 18S
rRNA and probably due to primer degeneracy.
25
(Diptera) species diversity using DNA barcoding. - Organismal Diversity and Evolution 10:
397-408.
Guryev, V., Makarevitch, I., Blinov, A. and Martin,
J. 2001. Phylogeny of the genus Chironomus
(Diptera) inferred from DNA sequences of
mitochondrial Cytochrome b and Cytochrome
oxidase I. - Molecular Phylogenetics and Evolution 19: 9-21.
Kaiser, T.S., Neumann, D., Heckel, D.G. and Berendonk, T.U. 2010. Strong genetic differentiation
and postglacial origin of populations in the
marine midge Clunio marinus (Chironomidae,
Diptera). - Molecular Ecology 19: 2845-2857.
Krosch, M.N. 2011. Phylogeography of Echinocladius martini Cranston (Diptera: Chironomidae) in closed forest streams of eastern Australia. - Australian Journal of Entomology 50:
258-268.
Acknowledgments
Our molecular studies of Chironomidae have
been funded mainly by an endowment by Evert
Schlinger to the University of California at Davis
(2000-2010). An NSF grant MIDGEPEET: A Collaborative Effort to Increase Taxonomic Expertise
in Understudied Families of Nematocerous Diptera award 0933218 to PI John (Kevin) Moulton
funded us to study Stictocladius, including to test
these techniques on extraction from pupal exuviae.
We thank Jon Martin for review, helpful comments
and permission to cite from his experiences.
References
Carew, M.E., Pettigrove, V. and Hoffmann, A.A.
2005. The utility of DNA markers in classical
taxonomy: using cytochrome oxidase I markers to differentiate Australian Cladopelma
(Diptera: Chironomidae) midges. - Annals of
the Entomological Society of America 98: 587594.
Krosch M.N., Baker A.M., Mather P.B. and Cranston, P.S. 2011b. Systematics and biogeography of the Gondwanan Orthocladiinae
(Diptera: Chironomidae). - Molecular Phylogenetics and Evolution 59: 458-468.
Martin J., Guryev V., Blinov A. and Edward D.H.D.
2002. A molecular assessment of the extent
of variation and dispersal between Australian
populations of the genus Archaeochlus Brundin (Diptera: Chironomidae). - Invertebrate
Systematics 16: 599-603.
Carew, M.E., Pettigrove, V., Cox, R.L. and Hoffmann, A.A. 2007. DNA identification of urban
Tanytarsini chironomids (Diptera: Chironomidae). - Journal of the North American Benthological Society 26: 587-600.
Cranston, P.S., Edward, D.H.D. and Cook, L.G.
2002. New status, distribution records and phylogeny for Australian mandibulate Chironomidae (Diptera). - Australian Journal of Entomology 41: 357-366.
Ekrem, T., Willassen, E. and Stur, E. 2007. A comprehensive DNA sequence library is essential
for identification with DNA barcodes. - Molecular Phylogenetics and Evolution 43: 530-542.
Ekrem, T., Stur, E. and Hebert, P.D.N. 2010. Females do count: Documenting Chironomidae
26
Wiederholm, T. 1986. Chironomidae of the Holarctic region. Keys and diagnoses. Part 2. Pupae. - Entomologica scandinavica, Supplement
28: 1-482.
Willassen, E. 1999. Take 25mg of insect.. simmering signals of phylogeny from chironomid
mitochondria. - Chironomus Newsletter on
Chironomidae Reseach 12: 5-10.
Willassen, E. 2005. New species of Diamesa (Diptera: Chironomidae) from Tibet: conspecific
males and females associated with mitochondrial DNA. - Zootaxa 1049: 19-32.
Wiederholm, T. 1983. Chironomidae of the Holarctic region. Keys and diagnoses. Part 1. Larvae. - Entomologica scandinavica, Supplement
19: 1-457.
27
Introduction
In his paper on chironomids of Thailand, Cranston
(2007) noted that the identity of Kiefferulus species of South East Asia was not clear. One group
for which uncertainty has existed in the past, is
three species described by Kieffer: K. barbatitarsis (Kieffer 1911), K. tainanus (Kieffer 1912), and
K. biroi (Kieffer 1918). Kieffer originally placed
K. barbatitarsis and K. biroi in Chironomus, but
he described K. tainanus as a Tendipes. Kiefferulus barbatitarsis, described from India, remained
in Chironomus, although Sublette and Sublette
(1973) classed it as unknown Chironomini. Chaudhuri and Ghosh (1986) re-examined the types in
the Indian Museum, along with rearings, and redescribed the species as Kiefferulus.
Material Examined
While the conclusions here are largely based on
a comparison of previously published work, some
additional specimens were examined.
Kiefferulus barbatitarsis:
28
Figure 1. Male hypopygium (left) and superior volsella (right) of K. barbatitarsis (top), K. tainanus (middle), and
Australian K. tainanus (below). Photos at top and middle courtesy of P.S. Cranston.
29
Nilodorum (e.g. Cranston and Martin 1989). Another obvious difference is in the shagreen pattern
of the pupa. While both species have an anterior
and posterior row of spines on tergite II, spines
on other tergites of K. barbatitarsis are relatively
small (Chaudhuri & Ghosh 1986), while those of
K. tainanus are larger and more extensive (Fig. 2).
The tergal spines of K. tainanus were well illustrated by Sasa (1979). It might be noted that material identified as K. tainanus in Australia (Saxena
et al. 1985, as N. biroi; Cranston et al. 1990, Bugledich et al. 1999) is probably a distinct but closely
related species. The most obvious difference is that
the male superior volsella is longer and narrower
Figure 2. Spinose patches on posterior two thirds of tergite V of pupa of Kiefferulus tainanus. Note the shorter
median spines and longer posterior spines.
Possibly the most obvious is the relatively shorter palps of K. tainanus, as can be seen in Table
1, which had led to it being placed in the genus
Table 1. Listing of most obvious differences between Kiefferulus barbatitarsis and Kiefferulus tainanus. Abbreviations
here and in text as in Sther (1980).
Character
Kiefferulus barbatitarsis
Kiefferulus tainanus
H-type
Y-type
Palp ratios
4: 3: 7: 11: 15
4: 3: 5: 8: 10
IV (Fig. 1)
relatively small
S1 setae
deeply feathered
palmate
Adult male
Pupa
Larva
30
Acknowledgements
I am grateful to Peter Cranston for helpful discussion and for making available material of K. tainanus and K. barbatitarsis from Thailand for DNA
extraction, and for providing the photographs of
the hypopygia of both species. Also to Kimio Hirabayashi for providing larvae of K. tainanus from
Japan for DNA sequencing. Sumitra Saxena provided specimens of K. tainanus from India, while
material from Australia was collected in the course
of fieldwork funded by the Australian Research
Grants Committee and the Office of the Supervising Scientist of the Alligator Rivers Region, with
further funding supplied by The University of
Melbourne.
Finally, the DNA barcode sequence of the mitochondrial COI gene of K. tainanus from Japan has
been published (Martin et al. 2007), and a further
sequence was obtained from an adult from Thailand. These can be compared with the equivalent
data for K. barbatitarsis (Fig. 3). As previously
noted, the K. barbatitarsis sequence came from
the same specimen as the hypopygium in Figure 1.
This comparison shows that, while there are 14
polymorphic sites between K. tainanus from Japan and India (2.3%), there are 42 (6.8%) and
39 (6.3%) respectively between the Japanese and
Thai sequences of K. tainanus and the sequence of
K. barbatitarsis. While the difference between the
Figure 3. Polymorphic sites in 621 bases of the mitochondrial COI sequences of two populations of K.
tainanus, and K. barbatitarsis
References
two K. tainanus samples is well within the arbitrary five percent limit for intraspecific variation of
this sequence, the difference of the K. barbatitarsis sequence falls outside that limit. In a Neighborjoining tree of Kiefferulus species (not shown), the
two species do not cluster together.
It therefore must be concluded that the gross similarity of some adult characters does not indicate
any particularly close relationship. Differences exist for larvae and pupae, as well as for the adults.
Indeed the two species were previously consid-
31
Goetghebuer, M. 1937-1954. Tendipedidae (Chironomidae). b. Subfamilie Tendipedinae (Chironomidae) A. Die Imagines. In Lindner, E.
(Ed.) Die Fliegen der palaearktischen Region
13c: 1-138.
32
Short Communications
An easy to make and simple designed rearing apparatus for Chironomidae
Armin Namayandeh1 and David V. Beresford2
Environmental & Life Sciences, Environmental Science Building Room A 105 Trent University, Peterborough, Ontario, Canada, K9J 7B8, Tel: (705)748-1011 X 7891
E-mail: arminnamayand@trentu.ca
1
Biology Department, Trent University, LHS D21, K9J 7B8, (705)748-1011 X 7540 Peterborough, Ontario, Canada
E-mail: davidberesford@trentu.ca
Introduction
Identification of immature Chironomidae can be impossible, as most species are identified based on adult
males, for which the associated pupae or larvae are not known or described. To overcome this, Chironomidae researchers rear larvae and pupae in aquaria to obtain the adults. This is usually done on fourth or
final instar larvae collected from aquatic habitats, and mass reared (e.g. Biever 1965; Hein and Mahadeva
1993). An advantage of mass rearing is that a large number of specimens can be obtained for identification. However, with mass rearing, matching individual larvae with pupal and adult stages is not possible.
To overcome this, larvae can be grown individually, a potentially costly undertaking. Even though most
chironomids can withstand the room temperatures, most cannot tolerate low oxygen levels that are created
in the vials (Mendes 2002).
In this paper we describe an inexpensive and rapid method of constructing an apparatus so that all three stages of individual specimen can be obtained, and that maintains sufficient oxygen during the rearing process.
Materials and Methods
Rearing vials were placed in a modified base constructed from centrifuge tube stands made of styrofoam,
which are readily available in chemistry or biology labs. A standard foam is 20cm long, 16cm wide and 3cm
indeep, and has 25 holes for individual centrifuge tubes. To make the base we cut a foam stand in half to
produce a 10cm by 16 cm stand, and glued this to a second stand to create a stepped structure (Figure 1i).
The rearing chamber was constructed from two centrifuge tubes to create an angled chamber with a screw
cap on each end. The bottom end with screw cap was made water tight using plumbers tape and further
sealed from the outside of the screw cap using aquarium glue. The top cap was left removable to allow access, and removal of emergent adults.
To modify the tubes, the bottom tube was cut perpendicularly at the 7.5 cm height (Figure 1ii). The bottom
section had two drilled holes, one sized to receive an air tube that feds the centrifuge tube from an air pump,
and the second used to feed the larvae. We found that the second hole could be dispensed with, and reared
4th instar larvae successfully in using stream water with no feeding access port. When the second hole was
installed, it was outfitted with a small plastic tube, 1.5 cm in length to allow feeding (Figure 1iii).The top
section was cut diagonally at the tip at a 45 angle (Figure 1ii). The bottom vial was then inserted into the
top at a 135o angle (Figure 1iii) and the joint of these two sealed using aquarium silicon. Finally, both the
feeding tube and the air tube were made watertight using aquarium cement.
Using this apparatus, stream water and individual larvae were added to the rearing chamber from the top,
and the chamber secured to the base using tape. The bottom fitted the stand holes and was secure.
Because the apparatus is small, any air pressure entering needed to be low to prevent water from splashing
in the chamber. We adjusted the air pressure using hose clamps, and found that 1-2 air bubbles every second
provided sufficient aeration.
33
Figure 1: i) Apparatus stand made from centrifuge tubes stands ii) Centrifuge tubes, bottom and top parts, the position
of holes and cuts, and assembling the top part to the bottom part iii) Apparatus on stand, position of larvae and adults
iv) The apparatus assembled.
The larvae were placed at the base of chamber, and adults emerged into the top section, and remained dry
due to the lip. Pupae exuviae and larva head capsule were obtained from the bottom part once the adults
were removed (Figure 1iii). Using this apparatus we have been able to acquire all three stages of Chironomidae for identification.
References
Biever, K.D., 1965. A Rearing Technique for the Colonization of Chironomid Midges. - Annals of the Entomological Society of America 58: 135-136.
Hein, J. and Mahadeva, M.N., 1992. Rearing and Maintaining Midge Cultures (Chironomus tentans) for
Laboratory Studies. - The American Biology Teacher 54: 236-240.
Mendes, H.F. 2002. Rearing Tanypodinae, Telmatogetoninae and Orthocladiinae in Brazil an empirical
approach. - Chironomus Newsletter on Chironomidae Research 15: 29-32.
34
35
Figure 1. Dorsal larval heads of Chironomini. A. Dicrotendipes septemmaculatus (Becker); B. Glyptotendipes paripes
Edwards); C. Glyptotendipes sp. E Epler 2001; D. Kiefferulus longilobus (Kieffer); E. Kiefferulus calligaster Kieffer; F. Kiefferulus barbatitarsis Kieffer; G. Endotribelos sp.; H. Goeldichironomus carus (Townes); I. Goeldichironomus sp.
This unit comprising the frons plus clypeus plus labrum is split off as a single structure at ecdysis it is
completely bounded by weak ecdysial lines. The genae are not part of this, and there are no genal elements
on the dorsal head. This is the basic pattern against which variants can be defined. Thus fusion of the clypeus with frons, with S3-5 on a single unit is a frontoclypeus. Fusion of frontoclypeus with labrum is
frontoclypeolabrum, bearing all S setae (1-5). Rarely (uniquely) there is a fusion of clypeus and labrum
to form a clypeolabrum (Kribiodorum). The anterior margin of the frons or frontoclypeus may be straight,
concave or convex. The frons, whether separate from the clypeus or fused, can have a fenestra (interiorly
thinned cuticle). We do not know if pits are homologous to oval fenestra, but certainly pits do not cooccur with fenestra and are not pits (depressions in the external cuticle).
The test of this hypothesis is its applicability in all Chironominae, and hopefully beyond. An issue is that
in taxonomic studies dorsal heads are not always illustrated. In many cases this seems to be because of
apparent uniformity, for example in Tanypodinae, the Harnischia cpx and many Tanytarsiini. However, all
36
37
38
Figure 1. Location of the Bolshoj Ljakhobskiy Island and coastal sector of Oyogos Yar in north-eastern Siberia.
39
Strait connects the Laptev and East Siberian seas, which are the widest and shallowest shelves worldwide.
The mean temperature of the warmest month at the coast is about 4C, the mean temperature of the coldest
month is about -30C (Station Cape Svyatoy Nos, 72 53 N, 140, 45 E in Rivas-Martnez, 1996-2009).
From a total of 50 chironomid head capsules, recovered from Eemian deposits, 16 taxa were identified (Table 1). Eight of these taxa belong to the subfamily Orthocladiinae and eight to the subfamily Chironominae
(five Chironomini, three Tanytarsini). The dominant taxa were Limnophyes, Smittia foliacea-type and the
Psectrocladius sordidellus-type. The majority of the taxa are indicators of temperate shallow lakes or littoral conditions that are associated with macrophytes. Limnophyes and the subdominant taxa, Metriocnemus eurynotus-type and Parametriocnemus/ Paraphaenocladius are also frequently associated with macrophytes (Cranston et al., 1983; Brodin, 1986). Furthermore, these taxa are typically indicative of lake level
fluctuations (Massaferro & Brooks, 2002). Smittia foliacea-type could be indicative of erosional processes
or unstable lake level conditions (Cranston et al., 1983). The subdominant taxa Chironomus anthracinustype and Cricotopus laricomalis-type are among the most frequently occurring taxa in present-day Yakutian lakes and can be found in a broad range of ecological conditions (Nazarova et al., 2005, 2008, 2011).
Tanytarsus pallidicornis-type 1 is characteristic of relatively warm and productive lakes (Brodin, 1986).
In contrast, Tanytarsus lugens-type and Parakiefferiella triquetra-type are cold stenotherms, occurring in
oligotrophic cold subarctic lakes (Walker & Mathewes, 1989; Brodin, 1986). Brillia and Endochironomus
albipennis-type often are associated with submerged wood and dead leaves (Cranston et al., 1983). Some
species of the group E. albipennis-type are leaf or stem miners (Pinder & Reiss, 1983).
The chironomid record indicates the presence of submerged coarse plant debris in the paleo-lake, most
probably the remains of trees and shrubs. Most of the identified chironomids indicate shallow water conditions and a pronounced macrophyte zone (Cranston et al., 1983; Brodin, 1986).
The Eemian was an interglacial period which began about 130,000 years ago and ended about 114,000
years ago. It was the second-to-latest interglacial period of the current Ice Age, the most recent being the
Holocene which extends to the present day. The prevailing Eemian climate is believed to have been similar
to that of the Holocene. Comparison of the Eemian fauna and modern chironomid assemblages from the
35 small permafrost lakes of Bolshoj Ljakhobskiy Island and Oyogos Yar mainland coastal area has shown
certain fuanistic differences (Table 1) with a higher proportion of cold tolerant taxa in modern chironomid
assemblages. Remains of terrestrial and freshwater organisms, including chironomids, preserved in frozen
deposits of a shallow Eemian paleo-lake indicate a boreal climate at todays Arctic during that time. The
use of transfer functions on the base of chironomid assemblages (Nazarova et al. 2011) brought a mean July
air temperature reconstruction of 12.90.9C and a water depth of 1.000.34 m for the time of deposition.
Table 1. Eemian chironomid fauna from Oygos Yar permafrost deposits and most frequent chironomid taxa from
Bolshoj Ljakhobskiy Island and Oyogos Yar mainland coastal area (Northern Yakutia, Russia. Taxa that are found in
modern and in Eemian sediments are given in bold.
%
22.4
17.0
13.9
10.1
5.4
2.9
2.7
2.7
2.2
1.9
1.3
1.2
1.2
1.2
1.2
1.1
40
Figure 2. Photos of chironomids from Eemian permafrost deposits: (a) Metriocnemus eurynotus-type; (b) Limnophyes;
(c) Smittia foliacea-type; (d) Microtendipes pedellus-type.
References
Brodin, Y. 1986. The postglacial history of Lake Flarken, interpreted from subfossil insect remains. - Internationale Revue der gesamten Hydrobiologie, 71: 371432.
Cranston, P.S., Oliver, D.R. and Saether, O.A. 1983. The larvae of Othocladiinae (Diptera: Chironomidae)
of the Holarctic region keys and diagnoses. In: Wiederholm (ed.): Chironomidae of the Holarctic region, Part 9. - Entomologica Scandinavica Supplement 19: 149-291.
Ilyashuk, B.P., Andreev, A.A., Bobrov, A.A., Tumskoy, V.E. and Ilyashuk, E.A. 2006. Interglacial history
of a palaeo-lake and regional environment: a multi-proxy study of a permafrost deposit from Bolshoy
Lyakhovsky Island, Arctic Siberia. - Journal of Paleolimnology. 35: 855872
Kienast, F., Wetterich, S., Kuzmina, S., Schirrmeister, L., Andreev, A., Tarasov, P., Nazarova, L., Kossler,
A., Frolova, L. and Kunitsky, V. 2011. Paleontological records prove boreal woodland under dry inland
climate at todays Arctic coast in Beringia during the last interglacial. - Quaternary Science Reviews 31
2134-2159.
Massaferro, J. and Brooks, S., 2002. The response of chironomids to Late Quaternary climate change in the
Taitao Peninsula, southern Chile. - Journal of Quaternary Science 17: 101-111.
Nazarova, L., Kumke, T., Pestryakova, L. and Hubberten, H.-W., 2005.
Chironomid fauna of Central Yakutian lakes (Northern Russia) in palaeoenvironmental investigations. - Chironomus Newsletter on Chironomidae Research 18: 25-27.
Nazarova, L.B., Pestryakova, L.A., Ushnitskaya, L.A. and Hubberten, H.-W. 2008. Chironomids (Diptera:
Chironomidae) in lakes of Central Yakutia and their indicative potential for paleoclimatic research. Sibirskii Ekologicheskii Zhurnal, 15 (3): 141150 (in Russian).
41
Nazarova, L., Herzschuh, U., Wetterich, S., Kumke, Th. and Pestjakova, L. 2011. Chironomid-based inference models for estimating mean July air temperature and water depth from lakes in Yakutia, northeastern Russia. - Journal of Paleolimnology. DOI: 10.1007/s10933-010-9479-4.
Pinder, L.C.V. and Reiss, F., 1983. The larvae of Chironominae (Diptera: Chironomidae) of the Holarctic
region. - Keys and diagnoses. In: Wiederholm (ed.): Chironomidae of the Holarctic region, Partb10. Entomologica Scandinavica Supplement 19: 293-435.
Rivas-Martnez, S. 1996-2009. Climate diagrams, Worldwide Bioclimatic Classification System. Internet
source, http://www.globalbioclimatics.org/plot/diagram.htm, Phytosociological Research Center, Spain.
Self, A.E., Brooks, S.J., Birks, H.J.B., Nazarova, L., Porinchu, D., Odland, A., Yang, H. and Jones, V.J.
2011. The distribution and abundance of chironomids in high-latitude Eurasian lakes with respect to
temperature and continentality: development and application of new chironomid-based climate-inference models in northern Russia. - Quaternary Science Reviews. 30 1122-1141
Smol, J.P., Wolfe, A.P., Birks, H.J.B., Douglas, M.S.V., Jones, V.J., Korhola, A., Pienitz, R., Rhland, K,
Sorvari, S, Antoniades, D., S.J. Brooks, S.J., Fallu, M.-A., Hughes, M, Keatley, B., Laing, T., Michelutti, N., Nazarova, L., Nyman, M., Paterson, A.M, Perren, B., Quinlan, R., Rautio, M., Saulnier-Talbot,
., Siitonen, S., Solovieva, N. and Weckstrm, J. 2005. Climate-Driven Regime Shifts in the biological
communities of Arctic lakes. - Proceedings of the National Academy of Sciences of the United States of
America 102: 4397-4402.
Walker, I.R. and Mathewes, R.W. 1989. Early postglacial chironomid succession in southwestern British
Columbia, Canada, and its paleoenvironmental significance. - Journal of paleolimnology 2: 1-14.
42
Chironomini
Chironomini (Diptera: Chironomidae). Keys to Central European larvae using mainly macroscopic characters.
2nd, revised edition.
By Orendt, C. & Spies, M. (2012).
64 p., more than 450 illustrations.
ISBN 978-3-00-038842-2
The second edition of these richly illustrated keys separates larvae of Central European non-biting midges by subfamilies, and
especially members of the tribe Chironomini by genera or some
smaller taxa. The work is also directed at workers with little previous experience in chironomid larvae. The keys emphasis is on
morphological features that are visible macroscopically without
elaborate preparations.
Chironomus (Meigen). Key to the larvae of importance to biological water analysis in Germany and adjacent areas.
By Orendt, C. & Spies, M. (2012).
Bilingual edition (English/German).
24 p., 47 illustrations.
ISBN 978-3-00-038789-0
This key identifies 11 species and species groups of significance
to water quality evaluation whose ecological requirements are
known sufficiently, at present, and whose larvae are reasonably diagnosable in terms of taxonomy. One of the aims of this work is to
reduce existing uncertainties and misinterpretations, particularly
in biological water analysis.
Chironomidae larvae in brackish waters of Germany and adjacent areas.
By Orendt, C., Dettinger-Klemm, A. & Spies, M. (2012).
Editor: Federal Environment Acency, Berlin.
Editions in English or German.
214 p., 1154 illustrations.
ISSN 2194-7902
The key includes about 90 taxa (genera, species groups, species)
that can be identified in the larval stage and have been recorded
from brackish waters in Germany and adjacent North Sea and
Baltic Sea areas.With its broad geographical scope and practical
approach, this monograph is unique and of equal interest for scientists and technical offices in countries along the North and Baltic
Seas.
Previews, informations and orders: www.hydro-bio.de/chironomidae.htm
43
Keys to
Central European larvae
using mainly
macroscopic characters
Second, revised edition
sponsored by
Deutsche Gesellschaft
fr Limnologie e.V.
Chironomus
Meigen
(Diptera: Chironomidae)
Bestimmungsschlssel zu den
in der biologischen Gewsseranalyse
bedeutenden Larven
Key to the larvae of importance to
biological water analysis
Chironomidae (Diptera)
Claus Orendt
Andreas Dettinger-Klemm
Martin Spies
44
Part 2 of A World Catalogue of Chironomidae (Diptera) is now available and was published on the 1st
October 2012. In Part 2 there are a total of 174 valid genera, 36 subgenera and 2,275 valid species plus 19
subspecies. In addition, there are a further 113 genus-group names and 1,470 species-group names which
are either synonyms, nomina dubia or unavailable. The total number of taxa (genus-group + species-group
names) in Part 2 is 4,087. The full citation is as follows:
Ashe, P. & OConnor, J. P. 2012. A World Catalogue of Chironomidae (Diptera). Part 2. Orthocladiinae.
Irish Biogeographical Society and National Museum of Ireland, Dublin. xvi+968pp.
Due to the large size it is printed in two sections, Section A (i-xvi, 1-468 pp) & Section B (469-968 pp).
The two sections each weigh about 1.5 kg and to save on postage costs each section is posted separately.
The total cost of a copy of Part 2 (including both sections) plus postage (Airmail only) and packing can be
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45
46
Finnish Chironomidae
Lauri Paasivirta
Ruuhikoskenkatu 17, B 5, 24240 Salo, Finland
E-mail: lauri.paasivirta@suomi24.fi
The current list of Chironomidae from Finland comprises 781 species altogether (Table 1). This includes
41 undescribed species, 12 problematic species (cf. or ?), 1 exuvial and 4 larval types. Compared to the
previous version from March 20th 2009, 1015 additions and 23 removals in the provincial records have been
made.
Regional and national check-lists are often based on records from a large number of scientists with variable
knowledge of both species distributions and taxonomy. For the list of species from Finland, over 90 % of
the records are now my own identifications. These are mainly of own sweep-net samples (74%) from the
period 1967 - 2012 and the massive whole-season Malaise-trap sampling by esp. Jukka Salmela during the
last ten years (71% of all Malaise-trap samples). Some additional records come from collections of exuviae
(mainly by Janne Raunio) and larvae (mainly by me). An overview of the sampling efforts divided by bioprovinces (Figure 1) and method is given in table 2.
The material is deposited mainly in my own collection consisting of thousands of slides and alcohol vials.
The hypopygia of the undescribed spp. have been photographied and drawn. DNA barcoding using partial
cytochrome c oxidase subunit 1 (COI) sequences has been done on 226 specimens of 77 species from
southwest Finland, including two types in Procladius choreus-agg. and three types in Chironomus pseudothummi- agg. Papers giving summaries of the local fauna in the main habitat types are now in preparation.
Table 2. Number of species by sampling efforts in bioprovinces.
14 light-trap nights are included in the sweep-net samples. Most
Malaise-traps were operated through the whole season. Only sampling lines/stations are regarded as larval samples.
Bioprov.
Exuval
samples sites
net
traps
traps
Larval
Al (A)
192
22
19
Ab (V)
510
794
78
37
15
310
N (U)
442
134
42
12
14
187
Ka (EK)
378
163
30
40
St (St)
467
149
25
10
114
Ta (EH)
523
365
44
20
52
388
Sa (ES)
414
137
10
160
Kl (LK)
261
39
Oa (EP)
426
302
34
13
18
136
Tb ( PH)
481
241
20
23
139
Sb (PS)
436
189
24
108
Kb (PK)
452
147
33
139
Om (KP)
415
159
16
136
Ok (Kn)
354
33
12
101
Oba (OP)
384
41
32
121
Obb (PeP)
411
33
32
87
Ks (Ks)
459
42
65
319
Lkoc (KiL)
466
186
45
82
Lkor (SoL)
444
52
26
18
78
Le (EnL)
494
137
55
52
Li (InL)
553
471
36
96
Total
781
3836
632
121
193
2802
47
48
X
X
X
Tanypodinae
X
X
Podonominae
95
510
192
Identified by L. Paasivirta ( % )
99
Ab
Al
Biogeographical province
Total number of species
X
X
92
378
Ka
92
442
X
X
X
X
X
X
X
X
X
X
95
523
Ta
98
467
St
X
X
X
X
X
X
X
X
X
X
X
X
X
90
261
Kl
96
414
Sa
X
X
X
X
X
X
X
X
X
X
X
X
X
99
481
Tb
91
426
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
97
452
Kb
97
436
Sb
X
X
X
X
X
X
X
X
E
X
X
X
X
X
X
X
x
X
X
X
X
X
X
99
354
Ok
92
415
Om
X
X
X
X
X
X
E
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
97
411
Obb
X
X
98
384
Oba
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
98
466
Lkoc
X
X
X
X
98
459
Ks
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
96
494
Le
X
X
X
92
444
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
97
553
Li
Table 1. Records of Chironomidae from Finland divided by bioprovinces from south to north (Al - Li). X = ident. by L. Paasivirta, x = ident. by others, M = collection of Museum of Zoology Helsinki
(confirmed by Lauri Paasivirta), E = exuviae collected and identified by J. Raunio (mostly confirmed by Lauri Paasivirta).
49
P. sp. 3 Kaldoaivi
P. sp. 2 Palsa
P. sp.1 Valassaaret
Al
Biogeographical province
Ab
X
X
Ka
X
X
X
X
X
X
X
X
X
Ta
St
X
X
X
X
X
X
X
X
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
X
X
Sb
X
X
X
X
X
X
X
Ok
X
X
X
X
Om
X
X
E
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
X
X
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Lkoc
X
X
X
X
X
X
Ks
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Le
X
X
X
X
X
X
X
X
X
Lkor
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
Li
50
Al
Biogeographical province
Diamesinae
Telmatogetoninae
Prodiamesinae
Ab
Ka
X
E
Ta
St
X
E
X
X
M
X
Kl
Sa
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
Kb
Sb
X
X
X
X
Ok
Om
X
X
X
X
X
X
X
Obb
X
X
Oba
X
X
X
X
X
X
X
X
X
x
X
X
X
Lkoc
X
X
Ks
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Le
X
X
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
Li
51
Orthocladiinae
Al
Biogeographical province
Ab
Ka
X
X
X
X
Ta
St
X
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
x
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
Ok
X
X
X
Om
X
X
X
X
X
X
X
x
X
X
X
X
X
X
Obb
Oba
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
X
X
Lkoc
Ks
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
X
X
X
X
X
X
X
X
X
X
X
X
Le
Lkor
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
Li
52
B. sp. 5 Pallas
B. sp. 4 Tsarmi
B. sp. 3 Malla
B. sp. 2 Syte
B. sp. 1 Syte
B. sp. 6 Rommas
Al
Biogeographical province
Ab
X
M
X
E
Ka
E,M
X
X
X
X
X
X
Ta
St
Kl
Sa
X
X
X
X
X
X
X
X
X
E
X
X
Tb
Oa
X
X
X
X
X
X
x
X
X
X
Kb
X
X
Sb
X
X
X
X
X
X
Ok
X
X
Om
X
X
X
X
X
X
X
X
X
X
X
X
Obb
X
X
X
X
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Lkoc
X
X
Ks
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
X
X
Le
Lkor
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
Li
53
X
X
C. sp. 2
C. sp. 1 Tvrminne
Al
Biogeographical province
Ab
X
X
X
X
Ka
X
x
X
X
X
X
X
X
X
Ta
X
X
St
X
X
X
X
X
X
X
X
X
X
X
X
X
Kl
Sa
X
X
X
X
X
x
X
X
X
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
X
X
X
X
X
Om
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
X
X
X
X
Oba
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
Lkoc
X
X
X
X
Ks
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
X
X
X
X
X
X
X
X
X
Le
X
X
X
X
Lkor
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
Li
54
Al
X
X
X
x
Biogeographical province
Ab
X
x
Ka
X
x
X
X
X
X
Ta
St
Kl
Sa
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
X
X
X
Om
X
X
X
X
X
X
X
X
x
X
X
X
X
X
E
X
X
X
?E
Obb
Oba
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
X
X
X
Lkoc
Ks
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
X
X
X
X
X
X
x
x
X
X
x
x
X
X
Le
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
Li
55
Al
Biogeographical province
E. sp.pr. gracei
H. sp. 1
H. sp.1 Kolmper
Ab
X
X
Ka
X
X
X
X
Ta
St
Kl
x
X
Sa
X
X
Tb
X
X
Oa
X
X
X
X
X
X
X
X
Kb
X
X
Sb
X
X
X
X
X
X
X
Ok
X
X
Om
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
X
X
X
X
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Lkoc
X
X
Ks
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
Le
X
X
X
X
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Li
56
L. er Saether, 1985
Al
Biogeographical province
Ab
X
X
Ka
X
X
X
X
Ta
St
X
X
X
Kl
Sa
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
X
X
X
Om
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Lkoc
Ks
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
X
X
X
X
X
X
X
X
X
X
X
X
X
Le
Lkor
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
Li
57
Al
Biogeographical province
Ab
Ka
Ta
St
X
M
Kl
Sa
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
Kb
X
X
X
Sb
X
X
X
X
X
E
X
X
X
X
X
Ok
Om
X
X
X
X
X
X
X
X
X
X
E
X
X
X
X
X
Obb
Oba
X
X
X
X
X
X
X
X
X
X
x
X
X
x
X
X
X
X
X
X
X
X
X
Lkoc
Ks
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
X
X
X
X
Le
X
X
Lkor
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
Li
58
Al
Biogeographical province
P. sp. 1
P. sp.1 Luukki
Ab
Ka
X
X
X
X
Ta
St
X
X
X
X
x
X
X
M
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
Om
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
X
X
X
Lkoc
Ks
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
X
X
X
X
X
X
X
Le
X
X
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Li
59
P. sp.1Tervola
Al
Biogeographical province
Ab
Ka
X
X
X
X
X
x
X
X
X
X
Ta
St
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
Kb
X
X
Sb
X
X
X
M
X
X
X
X
X
X
X
X
X
Ok
Om
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
Oba
X
X
X
X
X
X
X
X
X
X
X
M
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Lkoc
Ks
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
X
X
X
Le
X
X
X
X
X
X
X
Lkor
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
Li
60
S. sp.1 s. Tuiskunen
S. sp. 2 Jehkats
Ab
Al
Biogeographical province
Ka
X
M
X
X
Ta
St
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
Ok
X
X
Om
X
X
X
X
X
X
X
X
X
X
Obb
X
X
X
X
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Lkoc
X
X
X
X
X
Ks
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Le
X
X
X
X
Lkor
X
X
X
X
M
X
X
X
X
X
X
x
X
X
X
X
Li
61
Chironominae, Chironomini
Al
Biogeographical province
Ab
X
X
Ka
Ta
St
Kl
Sa
Tb
Oa
X
X
Kb
X
X
Sb
X
X
X
X
Ok
Om
X
X
X
X
X
X
Obb
Oba
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
Lkoc
Ks
X
X
X
X
X
X
x
X
X
X
X
X
X
X
X
X
X
X
X
X
Le
X
X
X
X
X
X
X
X
X
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Li
62
X
X
C. saxatilis agg.
Al
Biogeographical province
Ab
X
X
Ka
X
X
X
X
Ta
St
x
X
Kl
Sa
X
X
X
X
x
X
X
x
x
X
x
x
Tb
Oa
X
X
X
X
x
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
Ok
X
X
X
x
Om
X
X
X
X
X
X
X
X
X
X
X
X
Obb
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
X
X
X
X
Lkoc
X
X
x
X
Ks
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Le
Lkor
X
X
X
X
X
X
X
X
x
X
X
X
X
x
X
Li
63
X
Al
Biogeographical province
Ab
X
X
Ka
X
X
X
X
X
X
E
X
X
Ta
St
X
X
X
X
X
X
X
E
X
X
x
X
x
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
E
E
X
X
X
X
X
X
x
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
E
X
X
X
X
Ok
X
X
Om
X
X
X
X
X
X
X
X
X
X
E
X
X
X
X
X
X
X
X
X
X
X
Obb
X
X
Oba
X
X
X
X
X
X
X
X
x
X
X
X
X
X
Lkoc
X
X
x
Ks
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Le
x
X
X
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
Li
64
Al
Biogeographical province
Ab
X
X
X
X
Ka
X
X
X
X
X
x
X
X
Ta
X
X
St
X
X
X
X
X
X
X
X
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
Tb
X
X
X
X
Oa
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
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
E
X
X
X
X
X
X
X
X
X
X
Ok
Om
X
X
X
X
X
X
X
X
X
X
E
X
X
Obb
Oba
X
X
X
X
M
X
X
X
X
X
X
X
X
X
X
X
Lkoc
Ks
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
X
X
X
Le
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Li
65
X
X
X
X
P. sp. 1 Tornionjoki
P. sp. 1 Iijoki
Al
Biogeographical province
Ab
Ka
X
X
X
X
X
X
X
X
X
X
X
Ta
St
X
X
X
X
X
X
X
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
X
X
X
x
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
X
x
Om
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
Oba
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
Lkoc
Ks
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
X
X
Le
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Li
66
Al
x
X
X
X
X
X
X
Biogeographical province
Chironominae, Pseudochironomi
Chironominae, Tanytarsini
Ab
X
M
X
X
Ka
X
X
X
X
X
X
X
X
x
X
Ta
St
X
M
X
X
X
X
X
X
X
E
X
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
x
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
X
X
x
Om
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Obb
Oba
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Lkoc
x
X
X
X
Ks
X
X
X
X
X
X
X
X
X
Le
X
X
X
X
X
X
X
X
X
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
x
X
X
x
X
X
Li
67
Al
Biogeographical province
Ab
X
X
Ka
X
X
Ta
X
X
St
X
X
X
X
X
Kl
Sa
X
E
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
X
X
X
Kb
Sb
X
X
X
X
x
X
X
X
X
X
X
X
X
X
X
X
Ok
Om
X
X
X
X
X
X
X
X
X
X
X
x
X
X
X
X
X
x
X
Obb
Oba
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
X
X
X
X
X
Lkoc
X
X
X
X
X
X
Ks
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
X
X
x
x
X
X
Le
X
X
X
X
Lkor
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
Li
68
Al
Biogeographical province
Ab
X
X
Ka
X
X
X
X
X
X
X
X
X
Ta
St
X
X
X
X
X
X
X
X
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
Om
X
X
X
X
X
X
X
X
X
x
X
X
X
X
x
X
X
Obb
Oba
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
Lkoc
Ks
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
Le
Lkor
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
Li
69
X
X
X
Ab
Al
Biogeographical province
Ka
X
X
X
X
X
X
X
X
X
X
Ta
St
X
X
X
X
X
X
X
X
X
X
X
X
Kl
Sa
X
X
X
X
X
X
X
X
X
X
X
X
X
Tb
X
X
Oa
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Kb
X
X
Sb
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ok
X
X
Om
x
X
X
X
X
X
Obb
X
X
Oba
X
X
X
X
X
X
x
X
X
X
X
X
X
X
X
X
Lkoc
Ks
X
x
X
X
X
X
X
X
X
X
x
X
X
X
X
X
X
X
Le
X
X
X
X
Lkor
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Li
70
X
X
X
Ka
E
X
X
X
Ta
St
E
X
Kl
Sa
X
X
X
X
X
X
X
Tb
Oa
X
X
X
X
X
X
X
X
Kb
Sb
Ok
X
X
X
Om
X
X
X
E
X
X
X
X
Oba
X
X
X
X
X
Obb
X
X
X
X
Ks
X
X
X
X
Lkoc
X
X
X
Lkor
x
X
X
X
Le
X
X
X
X
X
X
Li
Gika, W. 2011. Six unusual Cladotanytarsus Kieffer: towards a systematics of the genus and resurrection of Lenziella Kieffer (Diptera: Chironomidae: Tanytarsini ). - Zootaxa 3100:
1-34.
Stur, E. and Spies, M. 2011. Description of Chaetocladius longivirgatus sp. n., with a review of C. suecicus (Kieffer) (Diptera: Chironomidae). - Zootaxa 2762: 37-48.
Spies, M. and Sther, O.A. 2004. Notes and recommendations on taxonomy and nomenclature of Chironomidae ( Diptera ). - Zootaxa 752. 1-90.
Makarchenko, E.A. 1994. Chironomids of the Subfamily Diamesinae ( Diptera, Chironomidae ) from Japan. II. Sympotthastia Pagast, 1947. - Bulletin of the National Science Museum,
Tokyo, Serie A, 20 (1): 51-58.
Ekrem, T. 2006. A redescription of Neozavrelia cuneipennis (Edwards) comb. nov., with a checklist of Neozavrelia species of the world (Diptera : Chironomidae). - Zootaxa 1153: 1-16.
Ekrem, T. 2004. Immature stages of European Tanytarsus species I. The eminulus-, gregarius-, mendax- and lugens species groups (Diptera: Chironomidae). - Deutsche entomologische
Zeitschrift 51: 109-158.
Cranston, P.S., Oliver, D.R. and Sther, O.A. 1989. The adult males of Orthocladiinae (Diptera: Chironomidae) of the Holarctic region - Keys and diagnoses. In Wiederholm, T. (Ed.)
Chironomidae of the Holarctic region - Keys and diagnoses. Part 3. Adult males. - Entomologica scandinavica Supplement 34: 165-352.
References
Notes: 1: ?syn. with takatensis (Tokunaga, 1936) (Makarchenko 1994); 1a: mostly hardly separable from Pseudokiefferiella; 2: ?syn. with submontanus; 2a: ?syn. with flavifrons (Johannsen, 1905). The differences in the colour and AR of the species (Sublette 1967) may be due to the water temperature (different generations); 3: might be earlier confused with subvernalis; 4: obviously belongs to Zalutschia; 4a: ?syn. with dissipates; 4b: includes just splitted longivirgatus Stur & Spies, 2011 (Stur & Spies 2011); 5a: syn. claripes Hirvenoja, 1973;
5b: syn. polychaetus Hirvenoja, 1989 (Spies & Sther 2004); 6: ?syn. brumalis; 6b: partly masked by minimus; 7: repl. bicolor (Zetterstedt, 1838) (Spies & Sther 2004), includes
colour form anderseni Sther, 1977; 7a: see Cranston, Oliver & Saether (1989): Fig. 9.58 sp. D and F; 8: ?syn. with platypus; 8a: in the provinces Lkoc-Li hypopygium resembles
fennicus; 8b: syn. Chironomus (Loboch.) dissidens (Walker, 1856); 9: syn. longipes Staeger, 1839; 10: syn. lateralis (Goetghebuer, 1934); 11: ?syn. nigronitens (Edwards, 1929); 12:
syn. gripekoveni (Kieffer, 1913); 13: syn. mancunianus (Edwards, 1929); 14: hypopygium separable from imbecilis s. Langton & Pinder 2007; 14a: syn. arcuatus (Goetghebuer, 1919);
15: syn. nudisquama (Edwards, 1929) and transcaucasicus Chernovski, 1949; 16: syn. uncinatum (Goetghebuer, 1921) (Dettinger-Klemm 2002); 16a: syn. wexionensis Brundin, 1947
(Gilka 2011); 17: syn. longappendiculata Albu, 1980 (Ekrem, 2006); 18: syn. Stempellinella distinctissimus Brundin, 1947; 19: replaces minor (Edwards, 1929) (Spies & Sther 2004);
20: syn. decipiens Lindeberg, 1967 and palmeni Lindeberg, 1967 (Ekrem 2004); 21: syn. simulans Lindeberg, 1967 and socialis Lindeberg, 1967 (Ekrem 2004).
Ab
Al
Biogeographical province