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CHIRONOMUS<br />
Newsletter on Chironomidae Research<br />
No. <strong>24</strong> ISSN 0172-1941 (printed) 1891-5426 (online) November 2011<br />
CONTENTS<br />
Editorial:<br />
Science communication 3<br />
18th International Symposium on<br />
Chironomidae, Trondheim Norway<br />
4-6 July 2011 4<br />
Deadheads at high altitude:<br />
Summary from the 10th Workshop<br />
on Subfossil Chironomidae,<br />
Finse, Norway, 2011 8<br />
Current Research 11<br />
Martin, J. <strong>Chironomus</strong> samoensis<br />
is a complex of species 11<br />
da Silva, F. L. Chironomidae types<br />
at the Museum of Comparative<br />
Zoology, Boston 18<br />
Short Communications 23<br />
Pseudodiamesa nivosa or<br />
Pseudodiamesa arctica? 23<br />
Typical types - a swan song? 30<br />
<strong>Chironomus</strong> Newsletter now<br />
compliant with the ICZN 35<br />
Cricotopus brevipalpis mining Potamogeton natans leaves. Photo by Susan Gresens and Torbjørn Ekrem
CHIRONOMUS Newsletter on Chironomidae Research<br />
Editors<br />
Torbjørn EkrEm, Museum of Natural History and Archaeology, Norwegian University of Science and<br />
Technology, NO-7491 Trondheim, Norway<br />
Peter H. Langton, 16, Irish Society Court, Coleraine, Co. Londonderry, Northern Ireland BT52 1GX<br />
The CHIRONOMUS Newsletter on Chironomidae Research is devoted to all aspects of chironomid research<br />
and aims to be an updated news bulletin for the Chironomidae research community. The newsletter is<br />
published yearly in October/November, is open access, and can be downloaded free from this website:<br />
http://www.ntnu.no/ojs/index.php/chironomus.<br />
Research articles for the CHIRONOMUS Newsletter are subject to peer-review.<br />
Contributions to CHIRONOMUS Newsletter on Chironomidae Research should be submitted online through<br />
the online journal system: http://www.ntnu.no/ojs/index.php/chironomus following the author guidelines.<br />
Submission deadline for contributions to the newsletter is July 1.<br />
Would you like to see your picture on the front <strong>page</strong>? Please send us your favourite midge photograph or<br />
drawing (torbjorn.ekrem@ntnu.no).<br />
Printed copies of the CHIRONOMUS Newsletter are available at the following libraries:<br />
Bavarian State Library, Munich, Germany<br />
Biology Library, Univeristy of Lund, Sweden<br />
Natural History Library, Smithsonian Institution, Washington DC, USA<br />
Library and Archives, Natural History Museum, London, UK<br />
Science and Health Library, University of Tromsø, Norway<br />
Science Library, Natural History Museum, University of Oslo, Norway<br />
The Royal Library, Copenhagen, Denmark<br />
<strong>Front</strong> <strong>page</strong> layout: Chironomid in title from photograph by Steve Marshall, Graphic design by Kolbjørn Skarpnes,<br />
NTNU Information <strong>Division</strong>.<br />
<strong>Front</strong> <strong>page</strong> photo: Cricotopus brevipalpis larvae mining Potamogeton natans leaves.<br />
Photo by Susan Gresens and Torbjørn Ekrem.<br />
2
Editorial<br />
Science communication<br />
Communication of research results may take many different forms. Some are modern in expression<br />
and content, while others definitely are more traditional. Traditional ways of communicating<br />
science does not mean that they are unsuitable in a modern research society, however. Printed<br />
articles in journals and newspapers and conferences and symposia are flourishing as never before<br />
and although many of these also have online versions, there is little doubt that many of us prefer<br />
to read text on paper instead of screen - not the least when sitting at the microscope with descriptions<br />
and identification keys. Moreover, certain research results must be either printed on paper<br />
or deposited in at least five publicly accessible libraries to be valid in the scientific community.<br />
This is the case for nomenclatorial changes in zoology as governed by the current International<br />
Code of Zoological Nomenclature (1999). This issue of the CHIRONOMUS Newsletter on Chironomidae<br />
Research will be the first in a number of years that is actually printed in a limited<br />
number of copies and sent to selected public science libraries in Europe and North America (see<br />
previous <strong>page</strong>). We will also send a copy to Zoological Record. This will allow us to publish<br />
descriptions of new species as well as nomenclatorial changes and we hope that you will consider<br />
Current Research in CHIRONOMUS Newsletter as a future venue for publication of your<br />
taxonomic research.<br />
This issue of the newsletter contains multiple additional examples of how science communication<br />
is performed in our community. We have reports from two recent meetings, current research<br />
articles and ongoing taxonomic discussions in our Short Communications section. The Current<br />
Bibliography that has been maintained by Odwin Hoffrichter for so many years, and is a valuable<br />
source of information on chironomid literature, unfortunately is not contained in this issue. We<br />
are currently discussing ways of maintaining this resource for the future, preferably in an online<br />
database, but a final solution is not yet ready.<br />
In the meantime, revival of the tradition of circulating reprints, perhaps by email attachment to<br />
reduce the cost, would ensure that references to papers published, particularly in journals<br />
‘obscure’ outside the country of publication, would not be omitted and citations missed.<br />
We hope that you enjoy this issue of the CHIRONOMUS Newsletter and look forward to see your<br />
contributions for the 2012 issue.<br />
Peter H. Langton 1 & Torbjørn Ekrem 2<br />
1 16, Irish Society Court, Coleraine, Co. Londonderry, Northern Ireland BT52 1GX.<br />
E-mail: PHLangton@kylebegave.fsnet.co.uk<br />
2 Museum of Natural History and Archaeology, Norwegian University of Science and Technology, NO-7491<br />
Trondheim, Norway. E-mail: torbjorn.ekrem@ntnu.no<br />
3
18 th InternatIonal SympoSIum on ChIronomIdae<br />
trondheIm norway 4-6 July 2011<br />
On the 4 th of July 2011 about 80 chironomists from<br />
as far afield as Japan, New Zealand, Argentina and<br />
Canada converged on the Norwegian University of<br />
Science and Technology (NTNU), a short walk uphill<br />
from Trondheim city centre. After registering,<br />
the delegates were warmly welcomed by Kaare<br />
Aagaard, Elisabeth Stur and Torbjørn Ekrem, before<br />
the Symposium was officially opened by the<br />
NTNU Museum Director Prof. Axel Christophersen.<br />
The honorary Thienenamnn lecture was<br />
delivered by Oliver Heiri entitled ‘Traces of past<br />
environments in the chemical composition of chironomid<br />
remains: stable isotopes in chironomid<br />
palaeoecology’. For the oldest of us, what has become<br />
possible in chironomid research – the analysis<br />
of stable isotopes in the fragments of long dead<br />
chironomid larval head capsules in order to reconstruct<br />
past climate changes, nitrogen pollution<br />
and the carbon food sources of lake foodwebs – is<br />
truly mind-boggling. One can only hope that there<br />
will be place still for the amateur chironomist with<br />
minimal equipment and finance. However, this is<br />
what these symposia are all about: widening the<br />
appreciation and knowledge of what can be done<br />
and is being done in the subject to which we are<br />
dedicated. This was an excellent start to a highly<br />
informative and exciting program.<br />
Logo from the symposium. Design: Elin Sandbakk<br />
After a very agreeable lunch in the university canteen<br />
(repeated on the following two days), the general<br />
program took off with two sessions of papers<br />
on palaeolimnology: six papers on what is being<br />
done on little bits of subfossil chironomid larval<br />
head capsules. For those of us who find difficulty<br />
in identifying a complete extant larva the palaeolimnological<br />
identifications appear little more than<br />
4<br />
wishful thinking, but as they so well demonstrated<br />
you do not have to have specific identification to<br />
derive useful information from the exercise. Once<br />
again the use of a carbon isotope to determine the<br />
food source and therefrom the eutrophication states<br />
of a lake, demonstrated just how sensitive the<br />
technique can be: it so happens that during periods<br />
of eutrophication more of the carbon food source<br />
is derived from methanogenic bacteria and this<br />
can be detected in the δ13C values derived from the<br />
subfossil chrironomid larval head capsules.<br />
There followed extreme activity in the lobby area<br />
outside the lecture theatre as posters were rapidly<br />
displayed on the substantial notice boards and the<br />
first period of viewing commenced. There was a<br />
wide variety of presentations including ecology,<br />
biodiversity, systematics traditional (even of some<br />
strange Mesozoic fossils with long biting proboscis)<br />
and more advanced (CAD and CO1 sequencing),<br />
six new species, species reappraised and<br />
species reinstated, and more palaeolimnogy.<br />
That evening an organ recital had been arranged<br />
at the Nidaros Cathedral, which has two organs:<br />
half the program was played on the older Baroque<br />
organ (Bach, including the famous Toccata<br />
and Fugue) and the second half on the main more<br />
recent Steinmeyer organ over the entrance to the<br />
cathedral (Egil Hovland – a composer new to most<br />
of the delegates). The organist was Øyvind Kåre<br />
Pettersen whose virtuosic performance, ending<br />
in a piece of Hovland fireworks, was pure artistic<br />
enjoyment for those of the delegates whose musical<br />
appreciation extends to that of works for large,<br />
loud organs. Rounding the evening off was a wine<br />
and canapés reception in the neighbouring restaurant:<br />
excellent provision for chatting the evening<br />
away with those of like mind.<br />
Tuesday morning was devoted to taxonomy and<br />
systematics. Before the coffee break five papers<br />
based on morphology and after, four based on<br />
molecular techniques and one on morphology<br />
were presented, demonstrating the wide variety of<br />
techniques being used at this time to distinguish<br />
species. The use of the polymerase chain reaction<br />
for separating <strong>Chironomus</strong> species at any stage of<br />
development is a novel addition to the techniques<br />
already in use for the genus.<br />
There followed a period of seventeen presentations<br />
on ecology and biomonitoring that lasted until<br />
lunch of the following day (Wednesday), that
Group photo of symposium participants outside University main building. Photo: Arnstein Staverløkk.<br />
took the delegates across the world from arctic Canada<br />
through Iceland, Europe, India, and Korea to<br />
New Zealand, from alpine pools to lowland lakes<br />
and from freezing temperatures to geothermal waters.<br />
The talks gave a fascinating overview of the<br />
variety of studies that chironomids can stimulate.<br />
These presentations were punctuated by the Tuesday<br />
evening Conference Dinner, held at Ringve<br />
Botanical Garden. The delegates were first shown<br />
round the garden by Vibekke Vange. The garden<br />
includes a large pond, where some delegates took<br />
the opportunity to do some collecting. The meal<br />
was a feast for the eye as well as for the palate, the<br />
evening further lifted by the camaraderie natural in<br />
the chironomist community.<br />
The last session of the oral proceedings began after<br />
lunch on Wednesday on toxicology, cytology and<br />
genetics. The seven presentations covered genetic<br />
control of behaviour, stress response to endocrine-disrupting<br />
compounds, desiccation tolerance<br />
mechanisms, gamma radiation tolerance in <strong>Chironomus</strong><br />
ramosus, DNA, chromosome responses to<br />
toxic contaminants and centromere structure. The<br />
Chironomidae are being comprehensively scrutinized<br />
and exhibiting just how plastic is their general<br />
physiology and cytology.<br />
The final session entitled “Chironomidae Symposium<br />
Forum”, began with in memoriam accounts<br />
5<br />
of chironomists recently deceased: Paul Freeman<br />
and Arthur Harrison by Peter Cranston, and Alevtina<br />
Shilova and Alexander Konstantinov by Andrey<br />
Przhiboro; a minute’s silence was observed<br />
in their memory. The award for student presentation<br />
was awarded to Alyssa Anderson and for the<br />
student poster was presented to Isabelle Proulx.<br />
Two presentations were then given for the venue<br />
of the next symposium: Naime Arslan’s invitation<br />
to Turkey was followed by Jolana Tátosová’s to<br />
the Czech Republic; I could have happily gone to<br />
either location, but the vote went to the Czech Republic.<br />
The Symposium was brought to a very enjoyable<br />
conclusion with a social held in the garden of<br />
Kaare Aagaard’s home. Our grateful thanks go to<br />
Kaare and his family for the warm welcome they<br />
gave and the excellent refreshments.<br />
The day following many of the delegates participated<br />
in the post symposium tour to the UNESCO<br />
cultural heritage site Røros, where we were given<br />
a guided tour of this post-mining community:<br />
exceedingly spartan living seems to have been a<br />
way of life for the miners – the tiny wooden cottages<br />
have left a lasting memory. Thence we travelled<br />
to the Sølendet Nature Reserve, where the<br />
wildflowers punctuated the fen with colour and the<br />
spring stream and fen pools provided the dedicated<br />
collectors an opportunity to get their nets in
action again. Wet but happy we boarded the coach<br />
again to pass over the scenically spectacular<br />
mountains to Tydal Valley and Sylane Mountains,<br />
where we were feasted at a remarkable restaurant<br />
in “the middle of nowhere”. We had a short time<br />
to photograph the wild flowers and do some more<br />
collecting before we were off again, now on our<br />
way back to Trondheim, but there was one more<br />
stop along the River Nea, a river regulated for hydroelectric<br />
power. We were given a guided tour<br />
of the large hydroelectric plant in the bowels of<br />
the mountain, which seems to be under the control<br />
of just two engineers. Culture, nature, spectacular<br />
scenery and pleasant company provided an experience<br />
that participants were very happy not to have<br />
missed.<br />
On behalf of all the delegates I offer our grateful<br />
thanks to Elisabeth Stur, Torbjørn Ekrem, Kaare<br />
Aagaard and their helpers for a most educational,<br />
well organised and memorable symposium and to<br />
the NTNU for providing the venue.<br />
Peter Langton<br />
Londonderry<br />
Memories of the 18th International Symposium<br />
on Chironomidae<br />
What a great symposium! I would qualify my first<br />
international symposium on Chironomidae as being<br />
refreshing, resourceful and helpful. Being<br />
used to attending much bigger gatherings, I really<br />
appreciated the more one-on-one, friendly and personal<br />
feeling of this symposium. Having the time<br />
to talk to people who work with chironomids and<br />
authors whom I had known only through their<br />
publications was really rewarding. In my doctoral<br />
research, <strong>Chironomus</strong> species identification has<br />
been my biggest challenge, and so I came to this<br />
meeting with the specific objective of getting some<br />
feedback on the work I have been doing. Not only<br />
was I able to get some answers to my questions,<br />
but I also made new and very helpful contacts. I<br />
would really like to thank the organisers for this<br />
wonderful, well-organised and pleasant symposium.<br />
I only have good memories of my experience<br />
in Trondheim!<br />
Isabelle Proulx<br />
INRS Eau Terre Envitonnement<br />
Université du Québec,<br />
Québec (QC), Canada<br />
6<br />
Len Ferrington, Pete Cranston and Anker Halvorsen discussing<br />
midges during a coffee break. Photo: Elisabeth<br />
Stur.<br />
First time impressions of the International<br />
Symposium on Chironomidae<br />
The list of speakers and attendees of the 18th International<br />
Symposium on Chironomidae reads very<br />
much like the bibliography of any rigorous literature<br />
review of the subject. Any misgivings one<br />
might have about attending such a specialized and<br />
esteemed event are quickly forgotten after meeting<br />
the participants. This community of researchers is<br />
very welcoming and it was encouraging to listen<br />
to veteran attendees who told of similar apprehensions<br />
when they arrived for their first symposium.<br />
The expertise each of the speakers was unequivocal<br />
and the breadth of knowledge was clear in the<br />
insightful questions and discussions. However, the<br />
most prominent feature of this group is the passion<br />
that people share for their subject. Not only<br />
do the participants exude zeal for their research,<br />
but many have stories of personal collections, research<br />
performed without funding and for personal<br />
interest, and some of us cannot walk by a pond or<br />
stream without looking for the presence of larvae<br />
or exuviae. The personal investment in this work<br />
seems so great that many of the people in attendance<br />
do not even seem aware of the legacy of<br />
their research. Outside the seminars people stand<br />
around microscopes, trade slides and specimens,<br />
and discuss taxonomic features as if recollecting<br />
the appearance of old friends. Others debate the<br />
best habitats to find certain species and swap stories<br />
of collecting trips. There is no substitution<br />
for being present at this conference. There are no<br />
identification keys precise enough or species list<br />
long enough to replace the value of these shared<br />
discussions. One cannot help but be impressed at<br />
the diversity of session topics and the addition of<br />
the Chironomid Subfossil Workshop held in Finse,<br />
Norway, prior to the conference only serves to in-
crease the number of participants and demonstrate<br />
the versatility of midges in answering ecological,<br />
biogeographical, physiological and genetic questions.<br />
While the monetary costs of attending an<br />
international conference can be high, the value<br />
here is far greater. The hosts of this event deserve<br />
special recognition and gratitude. Elisabeth Stur,<br />
Torbjørn Ekrem and the Museum of Natural History<br />
and Archaeology organised a memorable programme<br />
of talks, outings and social events.<br />
Christopher Luszczek<br />
York University<br />
Toronto, Canada.<br />
The 18th International Symposium on Chironomidae<br />
Participation in the Chironomidae Symposium<br />
in Trondheim was an important step for my PhD<br />
thesis and future work with chironomids. It was<br />
a wonderful opportunity to meet the best experts<br />
from all the fields, in which this amazing animal<br />
group can be used. It was really surprising for me<br />
how friendly and helpful were the people that I<br />
met, especially the “good old” experts. To have<br />
these contacts and to know that there is always<br />
someone you can ask for help, is the most important<br />
thing for a young scientist. Finally, this Symposium<br />
was the easiest way to get to Chironomidae<br />
heaven!<br />
Jarmila Lešková<br />
Commenius University<br />
Bratislava, Slovakia.<br />
7<br />
The 18th International Symposium on Chironomidae<br />
Attending the 18th International Symposium on<br />
Chironomidae in Trondheim, Norway, has given<br />
me the chance to present my own work and listen<br />
to talks on a wide variety of Chironomidae research.<br />
Researchers from all over the world attended<br />
the conference, presenting their work on different<br />
subjects, ranging from taxonomy to ecology. The<br />
conference was very well organized along with social<br />
events, which included an organ recital at the<br />
Nidaros Cathedral, a visit to the botanic gardens<br />
and a day trip to Røros and neighbourhood. I feel<br />
this conference is an important venue for scientists<br />
to meet, present their work, get to know each other<br />
and create collaborations.<br />
Elisabet Ragna Hannesdóttir<br />
University of Iceland<br />
Reykjavik, Iceland.
deadheadS at hIgh altItude: Summary from the 10 th workShop on<br />
SubfoSSIl ChIronomIdae, fInSe, norway, 2011.<br />
The 10th workshop on Subfossil Chironomidae<br />
was held at Bergen Museum/Finse Alpine<br />
Research Center, University of Bergen on June<br />
30. – July 2. 2011, and organised by Dr. Gaute<br />
Velle. There were 23 palaeo-chironomid researchers<br />
present, representing 10 different nationalities<br />
(see group photo). The purpose with these<br />
workshops is to exchange ideas and to have critical<br />
and fruitful discussions about topics ranging<br />
from larval identification to ecological interpretation<br />
– whether people are working with subfossil<br />
or contemporary data. Problems and progresses in<br />
chironomid palaeoecology are discussed among<br />
people with their fingers deep in the mud, and the<br />
informal presentations give a fine overview of current<br />
state-of-the-art research and development.<br />
The pre-workshop day on June 30th Participants at the workshop. Photo: Klaus P. Brodersen.<br />
in Bergen, was<br />
initiated with a welcome by Dr. Kari Hjelle, Head<br />
of Department at Bergen Museum. Dr. Richard<br />
Telford from the Department of Biology gave a<br />
presentation on “Numerical pitfalls in chironomid<br />
palaeoecology”. The talk was an introduction to<br />
the practical on statistics later on during the workshop<br />
(see below). A tour around the laboratory<br />
facilities with demonstration of the museums new<br />
automated identification and enumeration equipment<br />
was lead by Arild Breistøl and Gaute Velle.<br />
8<br />
After lunch, the workshop group went on the scenic<br />
Bergen-Oslo railway line to the field station<br />
at Finse (1222 m a.s.l.). A social ice-breaker hike<br />
to Hardangerjökulen glacier was guided by Gaute<br />
Velle. The hike in the wet and midge rich snow<br />
melt-water landscape, together with discussions<br />
on the glacier advance and retreats, gave a fine inspiration<br />
to the following two days workshop.<br />
The workshop was organised into four sessions<br />
with oral presentations; isotopes, transfer functions,<br />
climate, and environmental change. There<br />
were two practical sessions on taxonomy and statistics,<br />
and one session of general discussion.<br />
A number of ongoing projects (4 presentations)<br />
involve stable isotope analyses of either subfossil<br />
headcapsules, modern headcapsules, living larvae<br />
or food items from different habitats. Many<br />
methods and ideas from macroinvertebrate stable<br />
isotope research over the last decades are now being<br />
evaluated and implemented on subfossil material.<br />
The ongoing work involves optimization of<br />
analytical procedures with very small sample size,<br />
pre-treatment procedures, signal similarities between<br />
headcapsules and larvae, influence of food<br />
sources, in-lake and among lake variability etc.<br />
Combining the ecological information from C-13,
N-15 and O-18 with the information from the longterm<br />
subfossil records may open new windows to<br />
interpretation of catchment-, lake- and ecosystem<br />
development.<br />
The sessions on transfer functions and climate<br />
mainly contributed with both new and well known<br />
thoughts on the strengths and weaknesses of quantitative<br />
chironomid-based temperature models<br />
(e.g., previous workshops, Langdon et al. 2007,<br />
Brodersen et al. 2009, see also Velle et al. 2010).<br />
Interfering and confounding environmental variables<br />
both for the modern transfer functions assemblages<br />
and for the fossil assemblages still requires<br />
serious attention.<br />
The presentations on (human) environmental<br />
change brings the interpretation beyond discussion<br />
of climate and temperature. An interesting attempt<br />
to combine the changes in the chironomid (temperature)<br />
record with numerous findings of pre-historic<br />
human occupation and changing environment in<br />
SW Swedish Lapland, underlines the challenges in<br />
the eventual interpretations. Likewise, a study of a<br />
sediment filled lake basin (a lake that is no longer<br />
there; Danube lowlands, Slovakia) also requires<br />
interpretational aspects that challenge the analytical<br />
skills out of the quantitative modelling. Modern<br />
DNA sequencing methods has also reached<br />
palaeoecology. These methods have recently been<br />
used to confirm taxonomic identifications, such as<br />
a likely placement of the fossil Corynocera oliveri<br />
type into the parthenogenetic Tanytarsus heliomesonyctios<br />
Langton (Stur and Ekrem 2011), parse<br />
out relationships between cryptic species, and harmonize<br />
taxonomic nomenclature. However, the<br />
application of DNA sequencing has more diverse<br />
utilities than taxonomic clarification. Eventually,<br />
the analyses will be able to reconstruct population<br />
differences and link divergence events with<br />
geographic locations and perhaps clear our understanding<br />
of ambiguous indicator species, such as<br />
Corynocera ambigua.<br />
The discussions and challenges mentioned above<br />
perfectly set the scene for the practical session on<br />
statistics, tutored by Dr. Richard Telford. Using the<br />
statistical language R, Telford presented a method<br />
to test the statistical significance of a quantitative<br />
palaeo-environmental reconstruction inferred<br />
from biotic assemblages and transfer functions.<br />
A reconstruction is considered statistically significant<br />
if it explains more of the variance in the<br />
fossil data than most reconstructions derived from<br />
transfer functions trained on random environmental<br />
data (Telford & Birks, 2011). The workshop<br />
participants used the newly developed codes in R<br />
9<br />
(see Telford & Birks, 2011) and tested the significance<br />
of reconstructions resulting from data that<br />
was provided or from their own data (see workshop<br />
photo). Such significance testing fills a major<br />
gap in the range of numeric procedures available<br />
to palaeoecologists and it is recommended that<br />
these tests are used whenever a reconstruction is<br />
published.<br />
Practical on statistics tutored by Richard Telford.<br />
Photo: Klaus P. Brodersen<br />
Workshops are a fine stage for commencing on<br />
position papers. In the discussion session, some<br />
ideas for joint publications were presented and discussed.<br />
Many of us have data-sets that individually<br />
are small, but that together can constitute a wealth<br />
of information. Such data-sets include numerous<br />
data-points at a broad temporal and spatial resolution<br />
that can be used to test hypothesis on topics<br />
such as large-scale environmental influences,<br />
training set diversity, down-core diversity, reconstruction<br />
significance testing, or colonization. The<br />
repeated workshops on subfossil Chironomidae<br />
and the presence of identification guides, such<br />
as Brooks et al. 2007, help ensure that we have a<br />
common platform and that the taxonomy is comparable<br />
among data-sets. We anticipate this and<br />
future workshops will result in shared efforts and<br />
joint papers.<br />
It was generally agreed that having the workshop
at field stations or similar type of residence halls<br />
were a great success, giving rise to a good social,<br />
informal, fruitful and efficient workshop. Then,<br />
there is nowhere to escape from the discussion!<br />
We thank all for active participating and constructive<br />
discussions. The next workshop on subfossil<br />
Chironomidae will be in spring or early summer,<br />
2013, most likely in the UK.<br />
Klaus Peter Brodersen 1 , Ladislav Hamerlik 1 &<br />
Gaute Velle 2<br />
1Freshwater Biological Section, Biological Institute,<br />
University of Copenhagen, Denmark.<br />
E-mail: klaus.brodersen@gmail.com, ladislav.hamerlik@savba.sk.<br />
2Bergen Museum, University of Bergen, Norway.<br />
E-mail: nzlgv@uib.no.<br />
References<br />
Brodersen, K.P., Velle, G., Reuss, N.S. 2009.<br />
Trends and development in chironomid palaeoecology:<br />
Summary from the 9th Workshop on<br />
Subfossil Chironomidae. <strong>Chironomus</strong> 22: 5-7.<br />
10<br />
Brooks, S.J., Langdon, P.G., Heiri, O. 2007. The<br />
identification and use of Palaearctic Chironomidae<br />
larvae in palaeoecology. Quaternary<br />
Research Association Technical Guide 10, 276<br />
pp.<br />
Langdon, P., Holmes, N., Stefánsson, S.M., Hannesdóttir,<br />
E., Axford, Y. 2007. The 8th European<br />
Subfossil Chironomid Workshop, Reykjavik<br />
7–8th May 2007. <strong>Chironomus</strong> 20: 10–13.<br />
Stur, E., Ekrem, T. 2011. Exploring unknown life<br />
stages of Arctic Tanytarsini (Diptera: Chironomidae)<br />
with DNA barcoding. Zootaxa 2743:<br />
27-39.<br />
Telford, R.J., Birks, H.J.B. 2011. A novel method<br />
for assessing the statistical significance of<br />
quantitative reconstructions inferred from biotic<br />
assemblages. Quaternary Science Reviews<br />
30: 1272-1278.<br />
Velle, G., Brodersen, K.P., Birks, H.J.B., Willassen,<br />
E. 2010. Midges as quantitative temperature<br />
indicator species: lessons for palaeoecology.<br />
The Holocene 20: 989-1002.
Current researCh<br />
<strong>Chironomus</strong> samoensis is a complex of species<br />
Jon Martin<br />
Department of genetics, The University of Melbourne, VIC 3010, Australia<br />
E-mail: j.martin@unimelb.edu.au<br />
Abstract<br />
<strong>Chironomus</strong> samoensis, as currently recognised,<br />
is not a single species but a complex of more or<br />
less closely related species. C. samoensis Edwards<br />
1928 is redescribed from additional material, and<br />
considered to occur only in the Pacific region.<br />
Reasons for excluding material from other areas<br />
are given. C. flaviplumus Tokunaga 1940 is confirmed<br />
as the correct name for the Japanese material,<br />
the Indian material described by Chattopadhyay<br />
et al. (1991) is given the new name C. indiaensis,<br />
and new names are required for material from Australia<br />
and additional species from India.<br />
Introduction<br />
<strong>Chironomus</strong> samoensis Edwards 1928 was<br />
originally described from Samoa, American<br />
Samoa (Pago Pago) and Tonga. Since then, the<br />
species has been reported from other parts of<br />
Oceania (Tokunaga 1964, Cranston and Martin<br />
1989), Australia (Martin 2011), Japan (Hashimoto<br />
1977; Sasa and Hasegawa 1983), Taiwan<br />
(Yamamoto 1996), China (Wang 2000) and India<br />
(Chattopadhyay et al. 1991). However, as I will<br />
endeavour to show, most of these reports are<br />
misidentifications, and C. samoensis is probably<br />
limited to Oceania.<br />
I have not had the opportunity to examine the<br />
Edwards types, and his description does not<br />
include many characters important for species<br />
identification and is without illustrations. However,<br />
he did provide sufficient details to exclude many<br />
specimens described as C. samoensis from<br />
belonging to that species. One important feature<br />
is the superior volsella (SV) of the male, which<br />
Edwards compares specifically to the SV figured<br />
for C. imberbis (Kieffer 1917), as well as to other<br />
species such as C. dorsalis, and C. hawaiiensis,<br />
which all have a superior volsella (SV) of the D<br />
type (Strenzke 1959). While Kieffer’s illustration<br />
of the nomen dubium C. imberbis is not all that<br />
clear, all the known <strong>Chironomus</strong> species around<br />
11<br />
the Sydney area, the type location of C. imberbis,<br />
have a SV of the D type. In contrast to the Edwards<br />
(1928) description, some other descriptions of<br />
C. samoensis refer to the male as having a bootor<br />
foot-shaped superior volsella (e.g. Fig. 12a in<br />
Tokunaga 1964 (but see below), Chattopadhyay<br />
et al. 1991), corresponding to Strenzke’s (1959) S<br />
type.<br />
I believe that specimens I collected as larvae<br />
from Mapusaga, Tutuila Island, American Samoa<br />
(-14.29, 170.70), (29 February 1971), as well as<br />
a specimen from Faratogo, Tutuila (coll: N.R.<br />
Spencer, 29 June1964) from the Bishop Museum<br />
collection, are most probably the species described<br />
by Edwards, and they will therefore be more fully<br />
described here. The Bishop Museum specimen is<br />
labelled as C, samoensis, perhaps by Tokunaga,<br />
but this point is not certain. This will include a<br />
description of the immature stages for the first<br />
time. In general, the morphological terminology<br />
follows Sæther (1980), Webb & Scholl (1985)<br />
and Vallenduuk & Moller Pillot (1997). Colour<br />
could not be determined from these slide-mounted<br />
specimens.<br />
Results and Discussion<br />
Male<br />
Head: AR - 2.94 (2.51 - 3.23, 4); frontal tubercles<br />
33 µm (29-38, 4) long and 15µm (14-17, 3) wide;<br />
palpal proportions (micron) - 46 : 46 : 193 : 234 :<br />
354; clypeal setae 17-23.<br />
Thoracic setae: Acrostichal - at least 14 or 15;<br />
dorsolateral - 17-21; prealar - 4-5; scutellar in two<br />
rough rows, ant. 5-12, post. 12-15.<br />
Wing length 2.58 mm (2.40-2.68, 4), width 0.63<br />
mm (0.60-0.66, 4), VR 1.03 (1.02-1.04, 4).<br />
Legs: pale, tarsi slightly darker. Relative length of<br />
leg segments in Table 1.
Table 1. Lengths and ratios of leg segments of <strong>Chironomus</strong> samoensis<br />
Leg Fe Ti Ta1 Ta2 Ta3 Ta4 Ta5 LR F/T BR<br />
PI 1107 1000 1507 810 750 670 330 1.50-1.52 1.08-1.12 1.54-1.75<br />
PII 1170 1040 675 365 <strong>24</strong>5 160 115 0.62-0.67 1.07-1.17<br />
PIII 1290 1<strong>24</strong>5 1185 513 385 233 153 0.78-0.82 1.03-1.05<br />
Abdomen: pale, with darkening as described by<br />
Edward. Hypopygium: (Fig. 1) similar to that of<br />
C. dorsalis, with the SV of the D type, similar to<br />
fig. e of Strenzke (1959). The inferior volsella<br />
(IV) has mainly simple, curved setae, but a small<br />
number appear to have a small simple fork near the<br />
tip. About 4-6 setae on the 9th tergite near the base<br />
of the anal point.<br />
Female:<br />
No adult females are available amongst the present<br />
material, but one pupa with a pharate female<br />
was present. An important character is the relative<br />
proportions of the fore leg, particularly the tarsi,<br />
as Tokunaga (1964) notes that the Ta4 of specimens<br />
he assigned to C. samoensis was unusually<br />
long. The approximate lengths of these segments<br />
were measured (in micron) as: Fe 900 ; Ti 750 ;<br />
Ta1 1020 ; Ta2 620 : Ta3 470 : Ta4 610 : Ta5 340;<br />
Ta4 about same length as Ta2, and about one third<br />
longer than Ta3.<br />
Pupa: (Fig. 2)<br />
Exuviae length 6.8 (6.5-7.0, 3) mm (male), inner<br />
margin of wing case about 1.34 (1.27-1.42, 3) mm<br />
(male). Pale, with darkened caudolateral spurs.<br />
Cephalic tubules 87 (76-115, 3) µm long and 66<br />
Figure 1. Male hypopygium of <strong>Chironomus</strong> samoensis (left), superior volsella (right). Note the partly beaked apex of<br />
SV in the lower figure.<br />
12<br />
(56-80, 3) µm across the base, subterminal bristle<br />
about 68-80 µm in length. Basal ring about 151<br />
(129-164, 3) long and 70 (54-85, 3) µm. About<br />
67-77 hooks in row on segment II. Slight development<br />
of Pedes spurii B on segment II, progressive<br />
development of Pedes spurii A from segments IV<br />
to VI. Caudolateral spur of segment VIII about<br />
180 (155-200, 3) µm long, with 1 to 3 spines. 78-<br />
88 taeniae on each side of the anal lobe of male.
Figure 2. Pupal exuviae of <strong>Chironomus</strong> samoensis. Cephalic<br />
tubercles (above) and variations of spines on caudolateral<br />
spurs of segment VIII (below).<br />
Larva: (Fig. 3)<br />
A medium sized plumosus-type; length about<br />
12.5-12.7 mm (female) and 10.8-11.8 mm (male);<br />
lateral tubules about 280-360 µm; ventral tubules<br />
relatively long (anterior 1.76-2.16 mm; posterior<br />
1.80-2.68 mm), posterior pair longer and coiled;<br />
anal tubules moderately long (about 1.6-2.6 times<br />
longer than wide), dorsal pair (<strong>24</strong>0-410 µm) slightly<br />
longer than ventral pair (215-370 µm). Head<br />
capsule pale with darkening of the posterior half of<br />
the gula, frontal apotome sometimes pale but mostly<br />
with slight darkening, ventral head length 261-<br />
318 µm. Distance between antennal bases greater<br />
than the distance between the S4 setae. Mentum<br />
wider than usual, about 0.6 of ventral head length;<br />
centre trifid teeth with c2 teeth well developed (essentially<br />
type IV); 4th laterals reduced to about the<br />
level of 5th laterals (type II), 6th lateral variable,<br />
sometimes arising at same level as other laterals<br />
but generally appearing to be at a slightly lower<br />
level, apparently due to breakage resulting from<br />
wear. Ventromental plates separated by about 35-<br />
41% of the width of the mentum; each with about<br />
32-35 striae. Pecten epipharyngis with about 13<br />
(10-16, 8) sharp pointed teeth. Premandible with<br />
sharp teeth, outer tooth shorter than inner tooth,<br />
which is about twice as wide as the outer tooth.<br />
Mandible about 208-228 mm long, with 3rd inner<br />
tooth relatively pale and only partly separated<br />
(type II), about 13 (12-14, 8) striae on inner margin<br />
13<br />
Figure 3. Features of the larval head capsule of <strong>Chironomus</strong><br />
samoensis. a. Labrum and pecten epipharyngis,<br />
b. Premandible, c. Antenna, d. Mentum, e. Ventromentum,<br />
f. Mandible.<br />
at base, pecten mandibularis sparse, with about 8<br />
(7-10, 5) setae. Antenna five segmented, with A1<br />
almost 4 times longer than wide, RO between 0.4<br />
and 0.5 up from the base of the segment; relative<br />
length of antennal segments (micron) 110 : <strong>24</strong> : 6 :<br />
11: 7 ; AR 2.03-2.30.<br />
Cytology<br />
The polytene chromosomes (Fig. 4) available<br />
from six salivary gland squashes, prepared by the<br />
technique of Martin et al. (2006), are not of high<br />
quality, but are sufficiently good to show significant<br />
differences to those of other species that have<br />
been called C. samoensis. The arm combination<br />
is similar, being pseudothummi-cytocomplex (AE,<br />
BF, CD, G). There are at least two nucleoli, one<br />
central in arm G, and one about region 20 of arm<br />
F, with a large puff that may be a nucleolus near<br />
the middle of arm C. There are two Balbiani rings<br />
near one end of arm G. The only banding pattern<br />
that could be completely identified was that of<br />
arm A, which is the basic pattern of C. holomelas<br />
(Wuelker 1980).<br />
Based on these descriptions, diagnostic features of<br />
the species are: <strong>Front</strong>al tubercles relatively long;<br />
LR about 1.50 -1.52, fore Ta5 about one third of<br />
the length of the fore tibia, SV of the D-type, or<br />
“beaked”; in female fore Ta4 longer than Ta3 and<br />
about the same length as Ta2. In larva, antennal<br />
segment 3 relatively short, usually shorter than A5.<br />
In the polytene chromosomes, the nucleolus in arm<br />
G is median, and there is a further nucleolus about<br />
region 20 of arm F and usually a large puff in arm<br />
C.
Figure 4. Salivary gland chromosome complement of <strong>Chironomus</strong> samoensis. A-G - chromosome arm identifications,<br />
N - Nucleolus, BR - Balbiani ring.<br />
The first re-description of C. samoensis was by<br />
Tokunaga (1964). While the male colouration,<br />
AR and LR are quite similar to those described by<br />
Edwards, and Tokunaga describes the SV as similar<br />
to C. dorsalis, his figure has a triangular apex<br />
which can be misinterpreted as an S-type. However,<br />
Tokunaga’s illustration of the SV is presumably<br />
intended to depict the somewhat “beaked” SV seen<br />
in some specimens (for which there is no equivalent<br />
in Strenzke’s SV types). Tokunaga makes<br />
particular comment on the relative length of the<br />
tarsal segments of the fore leg of the female - “in<br />
female the fore tarsus with segment 4 far longer<br />
than 3 and slightly longer than 2”. He then gives<br />
the relative lengths of the segments as Fe 110 ; Ti<br />
86.5 Ta1 163.5, Ta2 84.5 ; Ta3 81.5 ; Ta4 88.5 ;<br />
Ta5 38.5. The relative lengths are about the same<br />
as those found for the Samoan specimen. The<br />
specimens described by Tokunaga from Micronesia<br />
are probably C. samoensis, although their LR<br />
is higher (1.75-1.84) and Tokunaga mentions the<br />
female abdomen as having faint oval spots on the<br />
terga, while Edwards states only that it is without<br />
distinct markings. In the event that re-examination<br />
of Tokunaga’s specimens (the specific location of<br />
which is not given) indicates that this is a different<br />
species, the name <strong>Chironomus</strong> eximius Johannson<br />
(1946) might be an available name. Otherwise,<br />
the Tokunaga description provides details of adult<br />
morphology not included in the present re-description.<br />
14<br />
On the other hand it can be shown that the species<br />
identified as C. samoensis from other geographic<br />
areas do not fit these descriptions of the species.<br />
Material from Japan has been described both as C.<br />
samoensis (Hashimoto 1977) and as C. flaviplumus<br />
Tokunaga 1940, and, while there is agreement that<br />
there is only one species, there is uncertainty as<br />
to which name should be used. Sasa (1978) used<br />
the name C. flaviplumus on the basis that the AR<br />
of Japanese specimens was higher (about 4.0) than<br />
that of C. samoensis, and the fore Ta5 was longer<br />
compared to the fore Ti (about 0.42). Although he<br />
gave the lengths of the leg segments of the female,<br />
he did not note that those of the fore tarsi did not<br />
agree with those of C. samoensis as specified by<br />
Tokunaga (1964). Ta4 is only the same length as<br />
Ta3 and shorter than Ta2. Despite this, Sasa and<br />
Hasegawa (1983) accepted the synonymy of the<br />
two species and it has been used in this way by<br />
many authors (Sasa and Kawai 1987; Elbetieha<br />
and Kalthoff 1988; Kuhn et al. 1987; Wuelker et<br />
al. 1989). Wuelker et al. recognized the synonymy<br />
as doubtful, but incorrectly stated the probable<br />
correct name was C. fulvipilus. These authors also<br />
gave the banding sequences of chromosome arms<br />
A, E. and F., and photographs of the other arms<br />
were kindly made available to the author. These<br />
show that at least arms A and G differ from those<br />
of C. samoensis. Arm A does not have the basic<br />
sequence of C. holomelas, but differs by complex<br />
inversions: flaA1, 1a-i, 2k-d, 9 - 4, 13 - 14, 3h-i, 12<br />
- 10, 2c - 1k, 3a-g, 15 – 19 (Wuelker et al. (1989).
The nucleolus in arm G is virtually terminal, not<br />
central.<br />
Given the differences noted above, it is clear that<br />
Sasa’s 1978 assessment was correct and the Japanese<br />
material should be called C. flaviplumus.<br />
<strong>Chironomus</strong> samoensis has also been reported<br />
from Taiwan (Yamamoto 1996) and China (Wang<br />
2000), but no details of the specimens were given.<br />
It is therefore unclear whether they are C. flaviplumus<br />
or another species.<br />
A species occurring in northern Australia was initially<br />
identified as C. samoensis (Martin 2011), but<br />
the morphology and cytology now indicate that<br />
this is incorrect. The Australian species is a close<br />
relative of C. flaviplumus, but is not identical. The<br />
AR of the adult males is lower (2.4-2.9), the LR<br />
is higher (1.82-1.96 cf. 1.63) and the fore Ta5 is<br />
slightly shorter (about 0.37 of Ti in males, 0.34 in<br />
females). The tarsal proportions of the female are<br />
similar to those of C. flaviplumus. Cytologically,<br />
the nucleolus in arm G is virtually terminal, but<br />
arm A carries the basal sequence as in C. samoensis,<br />
and arm F differs from that of C. flaviplumus<br />
by a simple inversion In14d-9. A new name will<br />
be required for this species.<br />
Finally, C. samoensis has also been reported from<br />
India, with a detailed description of West Bengal<br />
specimens by Chattopadhyay et al. (1991). I have<br />
been sent material identified as C. samoensis from<br />
Jammu and Kashmir, but these specimens do not<br />
agree with the description of Chattopadhyay et al.,<br />
or that of C. flaviplumus. The Jammu and Kashmir<br />
material may be comprised of two different species,<br />
although one may be C. incertipenis Chaudhuri<br />
and Das, 1996. The essential difference from<br />
C. samoensis evident in the description of Chattopadhyay<br />
et al. (1991) is that the superior volsella is<br />
a definite boot-shape (S-type), rather than similar<br />
to that of C. dorsalis, and so differs from any of the<br />
species discussed above. The fore Ta5 is slightly<br />
shorter, at 0.28 of Ti, but the description makes<br />
no mention an unusually long Ta4 in the female.<br />
While colour can be variable, it may be noted that<br />
the Indian specimens are described as generally<br />
brown rather than the green or yellowish colour<br />
of C. samoensis or C. flaviplumus. There are also<br />
differences in the larva. The larval head capsule<br />
is described as pale, the premandible has the outer<br />
tooth longer, the AR is lower (only 1.86) and the<br />
A3 segment is relatively longer (longer than A4,<br />
not shorter). There is no cytological data definitely<br />
associated with this species. This species therefore<br />
requires a new name, and is renamed C. indiaensis.<br />
15<br />
The material from Jammu and Kashmir requires<br />
further study as it comprises mostly larvae, with<br />
only a few adults. The situation is compounded<br />
by the fact that there are a number of described<br />
Indian species which are close cytologically, and<br />
have adults similar to those of the C. samoensis<br />
group. These include C. incertipenis Chaudhuri<br />
and Das 1996, which differs mainly in the dark,<br />
pointed anal point of the adult male and the shorter<br />
blunt inner tooth of the larval premandible, and<br />
C. ramosus Chaudhuri, Das and Sublette 1992,<br />
where the most obvious differences are the higher<br />
number of teeth in the larval pecten epipharyngis<br />
and the essentially equal teeth of the premandible.<br />
DNA sequence of the mitochondrial COI gene is<br />
available for a number of the Jammu and Kashmir<br />
specimens, including three adult males, and these<br />
indicate relationship to C. flaviplumus, with only<br />
5-7% base differences. The adult males (Fig. 5),<br />
while close to the C. samoensis group, are not C.<br />
samoensis and probably not C. flaviplumus. These<br />
adults are missing many leg segments or the antennae,<br />
which makes comparison difficult. They all<br />
have a “beaked” superior volsella. The AR is lower<br />
(about 3); LR is about 1.6 on the only specimen<br />
with the fore tarsi, and fore Ta5 is about 0.4 of Ti.<br />
Figure 5. Male hypopygium of a <strong>Chironomus</strong> species<br />
from Farooq Nagar, Jamu and Kashmir, India (Coll: P.<br />
Khanna) related to C. samoensis. Inset: “beaked” superior<br />
volsella.<br />
The larvae from Jammu and Kashmir do not seem<br />
to belong to either C. samoensis or C. flaviplumus,<br />
or to C. indiaensis. The head capsules generally<br />
have a darkened gula and the FA is very dark,<br />
sometimes mainly at posterior. The antennae
seem variable, with only some having a very short<br />
segment A3, others being longer than A5; AR<br />
about 1.9-2.0; the premandible teeth are nearer<br />
to equal length, and the outer tooth is about three<br />
quarters as wide as the inner tooth.<br />
Cytologically, most of these specimens have the<br />
arm F sequence flaF1 of Wuelker et al. (1989), but<br />
some specimens have the basal arm A sequence<br />
homA1, as in C. samoensis, while others a sequence<br />
that differs from flaA1 by a simple inversion. The<br />
nucleolus in arm G is generally subterminal, but<br />
the presence of a nucleolus cannot be definitely<br />
established in some specimens. While related<br />
to C. incertipenis and C. ramosus, they do not<br />
appear to be either of those species. C. ramosus<br />
characteristically has a nucleolus in arm B, and<br />
not in arm G (Nath and Godbole 1997), while C.<br />
incertipenis is present in Jammu and Kashmir, but<br />
differs in the mtCOI sequence (unpublished data).<br />
The presence of C. flaviplumus in India cannot<br />
be definitely ruled out, but it is more likely that<br />
at least two new names will be required for the<br />
material from Jammu and Kashmir.<br />
Summary<br />
These observations indicate that C. samoensis<br />
has not been found in any area other than the on<br />
Islands of the Pacific Ocean. C. flaviplumus could<br />
be more broadly distributed than its current range<br />
in Japan and Korea, the species described as C.<br />
samoensis by Chattopadhyay et al. (1992) can be<br />
renamed as C. indiaensis, while two or three new<br />
names will be required for other specimens from<br />
India, and Australian material.<br />
Acknowledgements<br />
I am grateful to many people for assistance<br />
in obtaining the materials used in this study:<br />
including P.M. Beales at Mapusaga who reared<br />
some of my larval sample to adults, M. Yamamoto<br />
for specimens of C. flaviplumus, J.P. Gupta for<br />
Varanasi samples, P. Khanna for Jammu and<br />
Kashmir samples, B.B. Nath for chromosome<br />
photographs of C. ramosus. Financial support for<br />
travel to Tutuila was provided by the University of<br />
Melbourne.<br />
References<br />
Chattopadhyay, S. Mazumdar, A., and Chaudhuri,<br />
P.K. 1991. Life stages and biology of<br />
<strong>Chironomus</strong> samoensis Edwards (Diptera:<br />
Chironomidae). - Proceedings of the National<br />
Academy of Sciences of India 61: 291-301.<br />
16<br />
Chaudhuri, P.K., Das, S.K., and Sublette, J.E. 1992.<br />
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Chironomidae). - Development 101: 591-603.<br />
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edu.au/Martin/AustChironfile/AustChiron.htm<br />
(accessed <strong>24</strong> June 2011).<br />
Martin, J., Andreeva, E.N., Kiknadze, I.I., and<br />
Wuelker, W. F. 2006. Polytene chromosomes<br />
and phylogenetic relationships of <strong>Chironomus</strong><br />
atrella (Diptera: Chironomidae) in North<br />
America. - Genome 49: 1384-1392.<br />
Nath, B.B. and Godbole, N.N. 1997. Chromosomal<br />
characterization of a tropical midge. - Cytobios<br />
91: 25-31.<br />
Sæther, O.A. 1980. Glosasary of chironomid<br />
morphology terminology (Diptera:<br />
Chironomidae). - Entomologica scandinavica<br />
Supplement 14: 1-51.<br />
Sasa, M. 1978. A comparative study of adults and<br />
immature stages of nine Japanese species of the<br />
genus <strong>Chironomus</strong> (Diptera, Chironomidae).<br />
- Research Report from The National Institute<br />
for Environmental Studies 3: 1-63.
Sasa, M. and Hasegawa, H. 1983. Chironomid<br />
midges of the tribe Chironomini collected from<br />
sewage ditches, eutrophicated ponds, and some<br />
clean streams in the Ryuku Islands, southern<br />
Japan. - Japanese Journal of Sanitary Zoology<br />
34: 305-341.<br />
Sasa, M. and Kawai, K. 1987. Studies on the<br />
chironomid midges of Lake Buwa (Diptera:<br />
Chironomidae). - Lake Buwa Studies<br />
Monographs 3: 1-120.<br />
Strenzke, K. 1959. Revision der Gattung<br />
<strong>Chironomus</strong> MEIG. I. Die Imagines von 15<br />
norddeutschen Arten und Unterarten. - Archiv<br />
für Hydrobiologie 56: 1-42.<br />
Tokunaga, M. 1940. Chironomidae from Japan<br />
XII. New or little-known Ceratopogonidae<br />
and Chironomidae. - Philippines Journal of<br />
Science 72: 255-311.<br />
Tokunaga, M. 1964. Chironomidae of Micronesia.<br />
- <strong>Insect</strong>s of Micronesia 12: 485-628.<br />
Wang, X. 2000. A revised checklist of chironomids<br />
from China (Diptera). In Hoffrichter, O. (Ed.)<br />
Late 20th Century Research on Chironomidae:<br />
an Anthology from the 13th International<br />
17<br />
Symposium on Chironomidae, Shaker Verlag,<br />
Aachen, pp. 629-652.<br />
Vallenduuk, H.J. and Moller Pillot, H.K.M. (1997)<br />
Key to the larvae of <strong>Chironomus</strong> in Western<br />
Europe. - RIZA Rapport 97.053: 1-13 + appendices.<br />
Webb, C.J. and Scholl, A. 1985. Identification<br />
of larvae of European species of <strong>Chironomus</strong><br />
Meigen (Diptera: Chironomidae) by morphological<br />
characters. - Systematic Entomology<br />
10: 353-372.<br />
Wuelker, W., Devai, Gy. and Devai, I. 1989.<br />
Computer assisted studies of chromosome<br />
evolution in the genus <strong>Chironomus</strong> (Dipt.)<br />
Comparative and integrated analysis of<br />
chromosome arms A, E, and F. - Acta Biologica<br />
Debrecina Supplementum Oecologica.<br />
Hungarica 2: 373-387.<br />
Yamamoto, M. 1996. New record of four<br />
<strong>Chironomus</strong> (Diptera, Chironomidae) from<br />
Taiwan. - Japanese Journal of Entomology<br />
64: 618.
Chironomidae Types aT The museum of ComparaTive Zoology, BosTon<br />
Fabio Laurindo da Silva<br />
Laboratory of Aquatic Entomology, Department of Hydrobiology, Federal University of São Carlos, P.O.<br />
Box 676, 13565-905, São Carlos, SP, Brazil.<br />
Section of Natural History, Museum of Natural History and Archaeology, Norwegian University of Science<br />
and Technology, NO-7491, Trondheim, Norway.<br />
Ernst Mayr Fellow, Museum of Comparative Zoology, Harvard University, Massachusetts, USA.<br />
E-mail: fabelha@hotmail.com<br />
Abstract<br />
The Chironomidae (<strong>Insect</strong>a: Diptera) type collection<br />
at the Museum of Comparative Zoology<br />
(MCZ) is reviewed. It comprises 23 primary types,<br />
as well as paratypes and paralectotypes for an additional<br />
29 species, mostly resulting from research<br />
by H. Loew and H. K. Townes, respectively. Notes<br />
updating the taxonomic status are provided for<br />
several species.<br />
Introduction<br />
The chironomid collection of the Museum of<br />
Comparative Zoology (MCZ) at Harvard University<br />
in Boston, Massachusetts (USA), includes<br />
about 1,200 specimens mostly stored dry on pins,<br />
with a small number mounted on slides. It is historically<br />
relevant but not well known. The main<br />
parts of this material have resulted from respective<br />
research endeavors by H. Loew (1807-1879) and<br />
H. K. Townes (1913-1990). Its geographic emphasis<br />
lies in the United States and Central America.<br />
The annotated checklist presented below (Table 1)<br />
was generated during a research visit to the Museum<br />
of Comparative Zoology in February 2011<br />
that was partly financed by an Ernst Mayr Travel<br />
Grant in Animal Systematics from MCZ/Harvard<br />
University.<br />
Results<br />
The collection includes 52 type specimens now<br />
classified as 10 holotypes, 5 lectotypes and 8 syntypes,<br />
as well as paratypes and paralectotypes of<br />
an additional 29 Chironomidae species (Table 1).<br />
The majority of the types belong to the subfamilies<br />
Chironominae and Tanypodinae.<br />
All names of species and references were checked<br />
with the corresponding literature. The notes on the<br />
species’ taxonomic status are based on Systema<br />
Dipterorum (Thompson & Pape 2010), as well as<br />
on published papers, catalogs (Oliver et al. 1990,<br />
Spies & Reiss 1996, Ashe & O’Connor 2009) and/<br />
or information on the collection labels. Some of<br />
the primary types were found labeled merely with<br />
“type”. The present respective interpretations of<br />
such specimens as holotypes, syntypes, etc., are<br />
based on all available data, e.g. those in the respective<br />
corresponding publications.<br />
The information presented here will be included<br />
in the database of primary types of Chironomidae<br />
already accessible online as part of the MCZ Entomology<br />
database (http://insects.oeb.harvard.edu/<br />
mcz).<br />
Table 1. Chironomidae species with types deposited in the collection of the Museum of Comparative Zoology<br />
at Harvard University, Boston. Abbreviations: H = holotype, L= lectotype, P = paratype, PL= paralectotype,<br />
S = syntype, M = male, F = female, AU = Australian, NE = Nearctic, NT = Neotropical<br />
Type<br />
number<br />
MCZT<br />
07433<br />
MCZT<br />
19425<br />
MCZT<br />
15655<br />
Original genus<br />
Original<br />
species<br />
<strong>Chironomus</strong> brachialis<br />
<strong>Chironomus</strong> bulbosa<br />
<strong>Chironomus</strong> imperator<br />
CHIRONOMINAE<br />
Author/<br />
Reference<br />
Coquillett,<br />
1901: 607<br />
Gerry, 1933:<br />
97<br />
Walley, 1926:<br />
64<br />
18<br />
Type Sex Preparation Distribution Note<br />
S M, F Pinned NE 1<br />
S M, F Pinned NT 2<br />
P M, F Pinned NE 3
MCZT<br />
194<strong>24</strong><br />
MCZT<br />
25385<br />
MCZT<br />
07434<br />
MCZT<br />
25404<br />
MCZT<br />
25393<br />
MCZT<br />
25381<br />
MCZT<br />
25389<br />
MCZT<br />
25398<br />
MCZT<br />
25401<br />
MCZT<br />
25387<br />
MCZT<br />
25394<br />
MCZT<br />
25399<br />
MCZT<br />
25386<br />
<strong>Chironomus</strong> jamaicensis<br />
<strong>Chironomus</strong> nigricans<br />
<strong>Chironomus</strong> taeniapennis<br />
Glyptotendipes testaceus<br />
Harnischia amachaerus<br />
Harnischia argentea<br />
Harnischia carinata<br />
Harnischia cuneata<br />
Harnischia potamogeti<br />
Omisus pica<br />
Polypedilum simulans<br />
Pseudochironomus banksi<br />
Pseudochironomus crassus<br />
MCZT Stenochironomus albipalpus<br />
MCZT Stenochironomus fuscipatellus<br />
MCZT Stenochironomus woodi<br />
MCZT<br />
27258<br />
MCZT<br />
25391<br />
MCZT<br />
25403<br />
MCZT<br />
25384<br />
MCZT<br />
25395<br />
MCZT<br />
25397<br />
MCZT<br />
25382<br />
Type<br />
number<br />
MCZT<br />
01260<br />
MCZT<br />
10388<br />
Tanytarsus subtendens<br />
Tribelos protextus<br />
Tendipes atroviridis<br />
Tendipes biseta<br />
Tendipes carus<br />
Tendipes ochreatus<br />
Tendipes tuberculatus<br />
Original genus<br />
Original<br />
species<br />
Chasmatonotus bimaculatus<br />
Chasmatonotus unimaculatus<br />
Gerry, 1933:<br />
96<br />
Johannsen,<br />
1905: 219<br />
Coquillett,<br />
1901: 607<br />
Townes, 1945:<br />
140<br />
Townes, 1945:<br />
168<br />
Townes, 1945:<br />
164<br />
Townes, 1945:<br />
158<br />
Townes, 1945:<br />
163<br />
Townes, 1945:<br />
159<br />
Townes, 1945:<br />
27<br />
Townes, 1945:<br />
43<br />
Townes, 1945:<br />
17<br />
Townes, 1945:<br />
15<br />
Borkent,<br />
1984: 66<br />
Borkent,<br />
1984: 64<br />
Borkent,<br />
1984: 91<br />
Townes, 1945:<br />
65<br />
Townes, 1945:<br />
69<br />
Townes, 1945:<br />
114<br />
Townes, 1945:<br />
127<br />
Townes, 1945:<br />
118<br />
Townes, 1945:<br />
115<br />
Townes, 1945:<br />
128<br />
ORTHOCLADIINAE<br />
Author/<br />
Reference<br />
Osten-Sacken,<br />
1877: 191<br />
Loew, 1864:<br />
50<br />
19<br />
S M, F Pinned NT 4<br />
P M, F Pinned NE 5<br />
S F Pinned NE 6<br />
P M, F Pinned NE<br />
P M Pinned NE 7<br />
P M Pinned NE 8<br />
P M Pinned NE 9<br />
P M Pinned NE 10<br />
P M Pinned NE 11<br />
P M Pinned NE<br />
P M Pinned NE<br />
H, P M Pinned NE<br />
H M Pinned NE<br />
P M, F Slide NE<br />
P M Slide NE<br />
P M, F Slide NE<br />
P M, F Pinned NE 12<br />
P M Pinned NE 13<br />
P M. F Pinned NE 14<br />
P M, F Pinned NE 15<br />
P M Pinned NE/NT 16<br />
P M Pinned NE 17<br />
H, P M, F Pinned NE 18<br />
Type Sex Preparation Distribution Note<br />
S M Pinned NE<br />
S M, F Pinned NE
Type<br />
number<br />
Original genus<br />
Original<br />
species<br />
MCZT Ablabesmyia parajanta<br />
MCZT Ablabesmyia tarella<br />
MCZT<br />
31778<br />
MCZT<br />
10367<br />
MCZT<br />
19423<br />
MCZT<br />
07431<br />
MCZT<br />
15660<br />
MCZT<br />
15657<br />
MCZT<br />
10368<br />
MCZT<br />
15658<br />
MCZT<br />
10369<br />
MCZT<br />
10376<br />
MCZT<br />
15661<br />
MCZT<br />
15662<br />
MCZT<br />
15656<br />
MCZT<br />
10370<br />
MCZT<br />
10371<br />
MCZT<br />
15659<br />
MCZT<br />
10372<br />
MCZT<br />
10373<br />
MCZT<br />
31777<br />
MCZT<br />
10374<br />
MCZT<br />
10375<br />
Type<br />
number<br />
MCZT<br />
27639<br />
Coelotanypus cletis<br />
TANYPODINAE<br />
Author/<br />
Reference<br />
Roback, 1971:<br />
373<br />
Roback, 1971:<br />
368<br />
Roback, 1963:<br />
174<br />
Type Sex Preparation Distribution Note<br />
P M Slide NE<br />
P M Pinned/Slide NE 19<br />
H M Slide NT<br />
Tanypus bellus Loew, 1866: 4 L, PL M, F Pinned/Slide NE 20<br />
Tanypus brooksi<br />
Tanypus concinnus<br />
Tanypus cornuticaudatus<br />
Tanypus currani<br />
Tanypus flavicinctus<br />
Tanypus garretti<br />
Gerry, 1933:<br />
95<br />
Coquillett,<br />
1895: 308<br />
Walley, 1925:<br />
277<br />
Walley, 1925:<br />
276<br />
Loew, 1861:<br />
309<br />
Walley, 1925:<br />
275<br />
S M Pinned NT 21<br />
S M, F Pinned NE/NT 22<br />
P M, F Pinned/Slide NE 23<br />
P F Pinned NE <strong>24</strong><br />
H M Slide NE 25<br />
P F Pinned/Slide NE 26<br />
Tanypus hirtipennis Loew, 1866: 5 H M Slide NE 27<br />
Tanypus humeralis Loew, 1866: 3 L, PL M, F Pinned/Slide NE/NT 28<br />
Tanypus mallochi<br />
Tanypus peleensis<br />
Tanypus pilicaudatus<br />
Walley, 1925:<br />
273<br />
Walley, 1926:<br />
64<br />
Walley, 1925:<br />
277<br />
P M Pinned NE 29<br />
P M, F Pinned/Slide NE/NT 30<br />
P M Pinned/Slide NE 31<br />
Tanypus pilosellus Loew, 1866: 5 L, PL F Pinned/Slide NE/NT 32<br />
Tanypus pinguis<br />
Tanypus prudens<br />
Loew, 1861:<br />
308<br />
Walley, 1925:<br />
275<br />
H F Pinned NE 33<br />
P M Slide NE 34<br />
Tanypus pusillus Loew, 1866: 5 H M, F Pinned NE 35<br />
Tanypus scapularis Loew, 1866: 2 L, PL M, F Pinned/Slide NE/NT 36<br />
Tanypus telus<br />
Roback, 1971:<br />
61<br />
H, P M. F Pinned/Slide NE<br />
Tanypus thoracicus Loew, 1866: 4 L, PL M, F Pinned/Slide NE 37<br />
Tanypus tricolor<br />
Original genus<br />
Original<br />
species<br />
Thalassomyia setosipennis<br />
Loew, 1861:<br />
309<br />
TELMATOGETONINAE<br />
Author/<br />
Reference<br />
Wirth, 1947:<br />
121<br />
20<br />
H F Pinned NE/NT 38<br />
Type Sex Preparation Distribution Note<br />
P M, F Pinned AU 39
Notes on Table 1<br />
<strong>Chironomus</strong> brachialis Coquillett, 1901 is now Demeijerea brachialis (Coquillett, 1901).<br />
<strong>Chironomus</strong> bulbosa Gerry, 1933, the spelling of which has been corrected to C. bulbosus, is a nomen dubium in Chironominae.<br />
<strong>Chironomus</strong> imperator Walley, 1926 is a junior synonym of <strong>Chironomus</strong> plumosus (Linnaeus, 1758).<br />
<strong>Chironomus</strong> jamaicensis Gerry, 1933 is a nomen dubium in Chironominae.<br />
<strong>Chironomus</strong> nigricans Johannsen, 1905 is now Endochironomus nigricans (Johannsen, 1905).<br />
<strong>Chironomus</strong> taeniapennis Coquillett, 1901 is a junior synonym of Stenochironomus hilaris (Walker, 1848).<br />
Harnischia amachaerus Townes, 1945 is now Cladopelma amachaerus (Townes, 1945).<br />
Harnischia argentea Townes, 1945 is now Cyphomella argentea (Townes, 1945).<br />
Harnischia carinata Townes, 1945 is now Parachironomus carinatus (Townes, 1945).<br />
Harnischia cuneata Townes, 1945 is now Demicryptochironomus cuneatus (Townes, 1945).<br />
Harnischia potamogeti Townes, 1945 is now Parachironomus potamogeti (Townes, 1945).<br />
Tanytarsus subtendens Townes, 1945 is now Endochironomus subtendens (Townes, 1945).<br />
Tanytarsus protextus Townes, 1945 is now a junior synonym of Tribelos jucundum (Walker, 1848).<br />
Tendipes atroviridis Townes, 1945 is now <strong>Chironomus</strong> atroviridis (Townes, 1945).<br />
Tendipes biseta Townes, 1945 is now <strong>Chironomus</strong> biseta (Townes, 1945).<br />
Tendipes carus Townes, 1945 is now Goeldichironomus carus (Townes, 1945).<br />
Tendipes ochreatus Townes, 1945 is now <strong>Chironomus</strong> ochreatus (Townes, 1945).<br />
Tendipes tuberculatus Townes, 1945 is now <strong>Chironomus</strong> tuberculatus (Townes, 1945).<br />
Ablabesmyia tarella Roback, 1971 is a junior synonym of Ablabesmyia mallochi (Walley, 1925).<br />
Tanypus bellus Loew, 1866 is now Procladius bellus (Loew, 1866).<br />
Tanypus brooksi Gerry, 1933 is nomen dubium in Pentaneurini.<br />
Tanypus concinnus Coquillett, 1895 is now Coelotanypus concinnus (Coquillett, 1895).<br />
Tanypus cornuticaudatus Walley, 1925 is now Helopelopia cornuticaudata (Walley, 1925).<br />
Tanypus currani Walley, 1925 is now Conchapelopia currani (Walley, 1925).<br />
Tanypus flavicinctus Loew, 1861 is a junior synonym of Clinotanypus pinguis (Loew, 1861).<br />
Tanypus garretti Walley, 1925 is a junior synonym of Psectrotanypus dyari (Coquillett, 1902).<br />
Tanypus hirtipennis Loew, 1866 is a junior synonym of Macropelopia decedens (Walker, 1848).<br />
Tanypus humeralis Loew, 1866 is now Coelotanypus humeralis (Loew, 1866).<br />
Tanypus mallochi Walley, 1925 is now Ablabesmyia mallochi (Walley, 1925).<br />
Tanypus peleensis Walley, 1926 is now Ablabesmyia peleensis (Walley, 1926).<br />
Tanypus pilicaudatus Walley, 1925 is now Helopelopia pilicaudata (Walley, 1925).<br />
Tanypus pilosellus Loew, 1866 is now Labrundinia pilosella (Loew, 1866).<br />
Tanypus pinguis Loew, 1861 is now Clinotanypus pinguis (Loew, 1861).<br />
Tanypus prudens Walley, 1925 is a junior synonym of Ablabesmyia pulchripennis (Lundbeck, 1898).<br />
Tanypus pusillus Loew, 1866 is a junior synonym of Procladius bellus (Loew, 1866).<br />
Tanypus scapularis Loew, 1866 is now Coelotanypus scapularis (Loew, 1866).<br />
Tanypus thoracicus Loew, 1866 is a junior synonym of Clinotanypus pinguis (Loew, 1861).<br />
Tanypus tricolor Loew, 1861 is now Coelotanypus tricolor (Loew, 1861).<br />
Thalassomyia setosipennis Wirth, 1947 is valid with the spelling Thalassomya setosipennis.<br />
21
Acknowledgements<br />
I am very grateful to Dr. Philip Perkins, Museum<br />
of Comparative Zoology, for his hospitality during<br />
my stay, and to Dr. James Hanken and Catherine<br />
Weisel for helping me organize my visit.<br />
Thanks also to Martin Spies, Dr. Marion Kotrba,<br />
Dr. Susana Trivinho-Strixino and Dr. Dalva Matos<br />
for supporting my application for an Ernst Mayr<br />
Travel Grant, and to Dr. Alaide Fonseca-Gessner<br />
for valuable suggestions on the manuscript. Finally,<br />
I would like to thank Zaki Habib-Gómez for<br />
checking the English text. The author has received<br />
financial support from the Ernst Mayr Fund (MCZ<br />
- Harvard University) and the National Council for<br />
Scientific and Technological Development (CNPq<br />
- Brazil).<br />
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22<br />
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- Am. Midl. Nat. 34: 1-206.<br />
Walley, G. S. 1925. New Canadian Chironomidae<br />
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Wirth, W. W. 1947. Notes on the genus Thalassomyia<br />
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Short CommuniCationS<br />
Pseudodiamesa nivosa or arctica? A confounded story of a midge moustache and an<br />
attempt at some taxonomic orthodontics<br />
Endre Willassen<br />
University Museum of Bergen, Postboks 7800, NO5020 Bergen, Norway.<br />
E-mail: endre.willassen@zmb.uib.no<br />
Midges of Pseudodiamesa are fascinating examples of insect adaptations to cold environments. Adults may<br />
be observed when they have emerged shortly after spring ice-thaw in mountain lakes and brooks. In extreme<br />
cases this may be as late as mid October in western Norwegian mountains when winter precipitation<br />
with snow has been particularly high. Adults may be swarming when temperature and wind allow them.<br />
Otherwise they will sit in the snow or some other substrate waiting for better times. The immature stages<br />
can also stand up against some rough treatment from the environment. I have seen larvae, deflated and looking<br />
like miniature sausage peels, trapped and completely surrounded by ice, when they recover from the<br />
melting ice and regain their body shape and vitality within minutes after the ice has thawed around them.<br />
Oliver (1959) advanced the idea that the North American Pseudodiamesa arctica is so morphologically<br />
distinctive by comparison with the European P. nivosa that it qualified for a subgenus of its own which he<br />
called Pachydiamesa. In the larvae, the alleged exclusive possession of a pair of labral lamellae (Oliver<br />
1959, 1983) has since figured in some papers as the key character to identify P. arctica from other Pseudodiamesa<br />
larvae.<br />
It was particularly the observation of such labral lamellae (see Figure 1C) that led Schnell and Willassen<br />
(1991) in a technical report to literally follow Oliver (1959, 1983) with some hesitation and to identify<br />
Pseudodiamesa arctica in Norwegian mountains. Moreover, since we were unable to observe clear cut D.<br />
nivosa characteristics as laid out by Oliver (1959) in the details of wing morphology, leg ratio and male genitalia<br />
we even suggested that the two taxa might possibly represent the same species. These were arguments<br />
to back up our identification but they were never intended as a formal nomenclatorial act of synonymisation.<br />
We did not have N. American material available for comparison and we also pointed to the taxonomic<br />
challenge that the candidate senior synonym, P. arctica, was originally described from a female. Basically<br />
we simply applied the authoritative taxonomic identification literature of that time to conclude that Pseudodiamesa<br />
arctica in the sense of Oliver was also observable in Norway. The only problem was that we could<br />
not see the difference from P. nivosa. When I brought specially fixed larvae to Novosibirsk for karyotype<br />
identification, Dr. Irina Kerkis identified the species to P. nivosa from chromosome characteristics. It is still<br />
not clear to me whether there was also a karyotype characterisation of something called P. arctica in place<br />
to compare with, but the cytotaxonomic interpretation of P. nivosa (at least at the time) corresponded to a<br />
species that actually has the “moustache” we would call labral lamellae (Figure 1C), although according to<br />
Oliver (1959, 1983) nivosa was not supposed to have one.<br />
Curiously, in another paper by Oliver (1976:1054) also including a record of P. arctica from Peary Land<br />
Greenland, there is no mention of labral lamella as a distinguishing character, - simply a statement about<br />
nivosa and arctica being very similar and that: “Based on a single larva of nivosa available for study, it may<br />
be possible to separate the larvae of the two species by the shape of the procercus.”.<br />
In the description of P. nivosa larvae from the Alps, Schmid’s drawings (1993:49, fig.21D) clearly show the<br />
labral lamellae that P. nivosa is not supposed to have. Although Schmid refers to Oliver (1983) in his list<br />
of taxonomic literature he states, contrary to Oliver without further comment, that the labral lamellae are<br />
the character that separates P. nivosa from P. branickii. So how can we actually identify arctica and nivosa<br />
from branickii larvae?<br />
In a more recent taxonomic review of Pseudodiamesa Ilyashuk et al. (2010) refer to Makarchenko’s (1985<br />
23
C<br />
D<br />
A<br />
Figure 1. A) Epipharynx of Pseudodiamesa nivosa larva as pictured by Ilyashuk et al. (2010: fig. 1C). The present<br />
author has added arrows to indicate the two pairs of chaetulae basales flanking seven scalelike teeth (also marked by<br />
the present author). B) Detail of Pseudodiamesa nivosa larva according to Ilyashuk et al .’s (2010) fig. 1D. The present<br />
author has pointed an arrow to the labral lamella which according to Ilyashuk et al. is absent in P. nivosa. Details of<br />
Pseudodiamesa larva from Norway that would key out to P. arctica from Oliver (1983) and to P. nivosa from Schmid<br />
(1993) Epipharynx has seven scale like teeth (1-7) and a pair of comb shaped labral lamellae (LL). Notice also that<br />
the right sensilla S1 is apically bifid while the left is simple. D) Photostack of the ventral part of epipharynx showing<br />
chaetulae laterales, the two pairs of chaetulae basales (arrows, also see Figures 1A, D, E) and teeth number 1 and 7. E)<br />
The same as D but with focus constrained more dorsally showing odd number symmetry of (totally 7) teeth next to tooth<br />
number 4 (Photos: C-E by E. Willassen)<br />
<strong>24</strong><br />
B<br />
E
et seq.) studies of the pecten epipharyngis stating that: “The pecten epipharyngis of the<br />
P. nivosa-group larvae consists of an even number, namely 3–4 pairs, of broad, elongate, apically blunt<br />
scales (fig. 1c, d), but that of the P. branickii-group larvae is characterized by an odd number, namely seven<br />
scales (fig. 1e, f)”. However, a closer look at their fig. 1c (see Figure 1A) shows that there are indeed seven<br />
epipharynx teeth (my numbering) in the specimen they identified as P. nivosa. There is no doubt that the<br />
counting of these structures can sometimes be a challenge, particularly in somewhat squashed microslide<br />
preparations. But in this case it seems pretty obvious to me. If we use the pairs of distinctive chaetulae basales<br />
as landmarks to define the outer margins of the pecten (Figures 1A, D), we see that there is symmetry<br />
around only one median tooth, the one which is numbered 4 in Figures 1 C-D. In other words, the assertion<br />
by Ilyashuk et al (2010) stating that all previous authors (including Oliver, Schmid, Janecek) were wrong<br />
when counting seven epipharynx scales in species other than those of the branickii – group does not seem<br />
justified.<br />
Ilyashuk et al. (2010) are also maintaining the notion that the labral lamellae are a unique feature of P. artica<br />
larvae. If so, we should conclude that Schmid also has documented presence of P. arctica in the Alps, but<br />
then what should we do with P. nivosa? I find it intriguing that while Ilyashuk et al. (2010) repeatedly state<br />
that P. arctica is the only species with labral lamellae, they also show a photo (Ilyashuk et al. 2010:fig.1D)<br />
of what they call P. nivosa in which the specimen clearly seems to possess a labral lamella (Figure 1B).<br />
Was this taxonomically important detail (as this paper testifies) overlooked by the authors or was it perhaps<br />
being interpreted as another type of structure? It is tempting to suspect that when Oliver (1983) described<br />
some Pseudodiamesa larvae (other than arctica) as having lamelliform SI setae, he may perhaps actually<br />
have referred to labral lamellae. Could this be one of the loose ends of the confounding problems with the<br />
taxonomy of Pseudodiamesa? With admittedly limited experience with Pseudodiamesa larvae from other<br />
parts of the world I have never come across what I would describe as lamelliform SI setae in Pseudodiamesa.<br />
What I have seen, however, is that the apically bifid SI claimed to be unique (Ilyashuk et al. 2010) to the<br />
American endemic P. pertinax may also be observed in Norwegian larvae with labral lamellae (Figure 1C).<br />
The specimen pictured here actually has a split on the right SI only. It is a sort asymmetry that is not uncommon<br />
in setae of immature Diamesinae so I would be sceptical to use it a litmus test of species identity.<br />
The observation that Ilyashuk et al. (2010) may have got the numbers of epipharynx teeth wrong in what<br />
they call nivosa unfortunately also leaves their claim that arctica has eight epipharynx teeth somewhat<br />
suspect. So are arctica and nivosa really separable as larvae? In his key to subgenera Oliver (1983) also<br />
included eight anal setae on the procercus as a distinctive feature for arctica. Figure 2A shows a specimen<br />
from Norway that would key out to Pachydiamesa arctica because it has 8 anal setae and labral lamellae.<br />
So why is it a problem with this record of P. arctica from Europe? Obviously because the differences between<br />
the larvae of arctica and nivosa described by Oliver (1983) and recently reiterated by Ilyashuk et al.<br />
(2010) don’t appear to hold up and the relationship between the two nominal taxa is still in need of critical<br />
review.<br />
But Pseudodiamesa arctica and P. nivosa could of course be inseparable in the larval stage and still be different<br />
species if we could observe distinctive and consistent differences in the adults. Ilyashuk et al. (2010)<br />
refer to a poster presentation by Makarchenko (2009) in which the statement is that P. arctica differs from P.<br />
nivosa by a more finger like distal part of the gonostylus and a posterolateral projection of the ninth tergite.<br />
However, the latter feature (Figure 2C) is also observable in specimens from Norway (Figure 2B) and does<br />
not seem to be a consistent difference although a few idiosyncratic drawings in the taxonomic literature<br />
could perhaps leave that impression. Moreover, when comparing Oliver’s (1959) drawing of P. arctica<br />
(Figure 2C) with specimens from Norway (Figures 2B) I am unable to see that P. arctica in the sense of<br />
Oliver (1959) is strikingly different and more finger like in the shape of the gonostylus. Is P. arctica sensu<br />
Oliver (1983) actually the same as P. arctica sensu Makarchenko (2009) and how do they relate to P.arctica<br />
sensu Malloch and to P. nivosa sensu authors?<br />
I am probably contributing my share to what seems to be a present state of chaos in Pseudodiamesa taxonomy<br />
by maintaining that I still cannot see the difference between the species that European workers have<br />
called P. nivosa and P. arctica in the sense of Oliver (1959). In anticipation of a more substantial documentation<br />
than a conference abstract I would also feel uncomfortable by having to choose between arctica<br />
and nivosa based on a critial value of 0.56 in leg ratio (Makarchenko 2009). It seems to me that molecular<br />
data could help in getting us out of some of the troubled waters that Pseudodiamesa taxonomy appears to<br />
be in these days. But even more so important is it that we try to resolve the taxonomic problems that may<br />
25
A<br />
B C<br />
Figure 2. A) Procerci of Pseudodiamesa larva from Norway that would key out to P. arctica from Oliver (1983). Arrows<br />
point to eight anal setae. B) Male hypopygium of Pseudodiamesa from Norway with arrow pointing to caudolateral<br />
projection of tergite which according to Makarchenko (2009) is a key character to separate P. arctica from P. nivosa. C)<br />
Drawing of P. arctica hypopygium from Oliver (1959). The present author disputes that the gonostyli are conspicuosly<br />
more fingerlike than in the species that has been known as P. nivosa in Europe. (Photos: A,B by E. Willassen)<br />
be rooted in ill-defined characters, misconceptions based on mismatching terminologies and particularly<br />
from blind acceptance of old taxonomic decisions that did not seem to stand the test of additional material<br />
and new observations. We have never been better equipped technologically and electronic pictures are a<br />
blessing.<br />
References<br />
Ilyashuk, B.P., Ilyashuk, E.A., Makarchenko, E.A., Heiri, O. 2010. Midges of the genus Pseudodiamesa<br />
Goetghebuer (Diptera, Chironomidae): current knowledge and palaeoecological perspective. - Journal<br />
of Paleolimnology 44: 667-676. DOI: 10.1007/s10933-010-9446-0. (For higher resolution pictures see:<br />
http://www.springerimages.com/img/Images/Springer/JOU=10933/VOL=2010.44/ISU=2/ART=9446/<br />
MediaObjects/LARGE_10933_2010_9446_Fig1_HTML.jpg)<br />
Makarchenko, E.A. 1985. Chironomids of the Far East of USSR. The subfamilies Podonominae, Diames-<br />
26
inae, and Prodiamesinae (Diptera, Chironomidae). DVNC AN SSSR, Vladivostok.<br />
Makarchenko, E.A. 2009. Pseudodiamesa nivosa (Goetghebuer, 1928) is not synonym of Pseudodiamesa<br />
arctica (Malloch, 1919) (Diptera: Chironomidae). XVII International Symposium on Chironomidae,<br />
July 5–10, 2009, Nankai University, Tianjin, China. Program and Abstracts, p 71. http://entomology.<br />
nankai.edu.cn/17chiro/program_abstr_Ver.6.30.pdf<br />
Oliver, D.R. 1959. Some Diamesini (Chironomidae) from the Nearctic and Palearctic. - Entomologisk Tidskrift<br />
80:48–64.<br />
Oliver, D.R. 1976. Chironomidae (Diptera) of Char Lake, Cornwallis Island, N.W.T., with descriptions of<br />
two new species. - Canadian Entomologist 108:1053–1064.<br />
Schmid, P.E. 1993. A key to the larval Chironomidae and their instars from Austrian Danube Region<br />
streams and rivers. Part I, Diamesinae, Prodiamesinae and Orthocladiinae. Federal Institute for Water<br />
Quality of the Ministry of Agriculture and Forestry, Wien, Austria.<br />
Schnell, Ø.A., Willassen, E. 1991. Fjærmyggarten Pseudodiamesa arctica (Malloch) i to høyfjellsreservoarer<br />
(The chironomid species Pseudodiamesa arctica (Malloch) in two high mountain reservoirs).<br />
Report No 76, Freshwater Ecology and Inland Fisheries Laboratory, Department of Zoology, University<br />
of Bergen, Bergen, Norway.<br />
Moustached Pseudodiamesa is still in waiting for a modern cytogenetic approach<br />
and a taxonomic revision: a reply to Willassen<br />
Boris P. Ilyashuk1 1, 2<br />
& Elena A. Ilyashuk<br />
1 Institute of North Industrial Ecology Problems, Kola Science Centre, Russian Academy of Sciences, 14<br />
Fersman St., Apatity, Murmansk reg. 184209, Russia. E-mail: ilboris@yandex.com<br />
2 Institute of Ecology, University of Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria.<br />
E-mail: elena.ilyashuk@uibk.ac.at<br />
Beginning with the study of chironomid subfossils in Schwarzsee ob Sölden, the Austrian Alps (Ilyashuk<br />
et al. 2011), we have tried to compile all available information about aquatic invertebrates in this high-alpine<br />
lake. To our surprise, we found rather contradictory data on the Pseudodiamesa species. Janecek (1998)<br />
reported that P. nivosa inhabits the lake whereas Raddum et al. (2004) have found that the Pseudodiamesa<br />
subfossils from a short core taken in the lake are represented by P. branickii. To all appearances, difficulties<br />
in the identification of Pseudodiamesa subfossils resulted in the mentioned taxonomic inconsistency. The<br />
following study of contemporary Pseudodiamesa larvae and their subfossils showed that the genus is represented<br />
by P. nivosa in this lake (Ilyashuk et al. 2011). However, we encountered a number of contradictions<br />
in the available literature for identification of contemporary Pseudodiamesa larvae. Some of them are noted<br />
in Willassen (2011). In Ilyashuk et al. (2010), we summarise current experience in splitting the genus into<br />
two intra-genus morphotypes within the subfossil material from different Arctic and Alpine regions. Later,<br />
working with subfossils from another high-alpine lake, we got chironomid remains of much better preservation<br />
and found that the P. nivosa subfossils have the labral lamellae and the pecten epipharyngis consisting<br />
of seven scales (Fig. 1), as is described in Schmid (1993). The observations by Willassen (2011) based on<br />
contemporary material confirm it as well. Nevertheless, we would like to emphasize that the mentum is a<br />
well-preserved structure in subfossil specimens and can be one of the best morphological characters for differentiating<br />
Pseudodiamesa species-group morphotypes in subfossil material (Ilyashuk et al. 2010).<br />
The current state of knowledge on the genus Pseudodiamesa does not allow us to confirm or disprove<br />
hypotheses concerning synonymisation of some species. Schnell and Willassen (1991) suppose that P.<br />
nivosa is synonymous with P. arctica and Makarchenko (1998) assumes that P. nepalensis is a synonym of<br />
P. nivosa. Both hypotheses are realistic and testable, taking into account modern cytogenetic approaches<br />
and techniques. Unfortunately, the discussion in Willassen (2011) does not add new data to check the hypothesis<br />
about synonymisation of P. nivosa and P. arctica. However, there are the first successful steps in<br />
resolving some Pseudodiamesa taxonomic problems. A recent comparative study of karyotypes of some<br />
27
Figure 1. Pseudodiamesa nivosa (Goetghebuer) subfossils from a high-mountain lake in the Italian Alps: A) pecten<br />
epipharyngis (PE); B) pecten epipharyngis (PE) and labral lamella (LL); C) labral lamella.<br />
Pseudodiamesa species, conducted by Ermolaeva (2005) and supervisored by Dr. E. Makarchenko and Dr.<br />
I. Kiknadze, provides evidence that P. nivosa, P. stackelbergi, and P. latistyla are valid species. Moreover,<br />
comparative karyological analysis of P. branickii, collected from different regions in Eurasia (Germany,<br />
Bulgaria, Kyrgyzstan, Russian Far East, and Japan) revealed that there are at least four distinct chromosomal<br />
races, which may represent sibling species corresponding to the P. branickii morphotype (Ermolaeva<br />
2005). Hopefully, other species of this genus will be included in subsequent comparative karyological<br />
analyses.<br />
References<br />
Ermolaeva, O.V. 2005. Citotaksonomicheskiy analiz vidov podsemeystv Diamesinae i Prodiamesinae:<br />
(Diptera, Chironomidae) [Cytotaxonomic analysis of species of the subfamilies Diamesinae and Prodiamesinae<br />
(Diptera, Chironomidae)]. Ph.D. Thesis, Novosibirsk State University, Novosibirsk, Russia.<br />
239 p.<br />
Ilyashuk, B.P., Ilyashuk, E.A., Makarchenko, E.A. and Heiri, O. 2010. Midges of the genus Pseudodiamesa<br />
Goetghebuer (Diptera, Chironomidae): current knowledge and palaeoecological perspective. - Journal<br />
of Paleolimnology 44: 667–676.<br />
Ilyashuk, E.A., Koinig, K.A., Heiri, O., Ilyashuk B.P. and Psenner, R. 2011. Holocene temperature variations<br />
at a high-altitude site in the Eastern Alps: a chironomid record from Schwarzsee ob Sölden, Austria. -<br />
Quaternary Science Reviews 30: 176–191.<br />
Janecek, B. 1998. Fauna Aquatica Austriaca, Taxonomie und Ökologie aquatischer wirbelloser Organismen,<br />
Teil V. Diptera: Chironomidae (Zuckmücken), Larvenstadien mitteleuropäischer Gattungen und<br />
österreichischer Arten, Bestimmung von 4. Universität für Bodenkultur, Abteilung Hydrobiology, Wien,<br />
Austria. 117 p.<br />
Makarchenko, E.A. 1998. Chironomidy podsemeystva Diamesinae (Diptera, Chironomidae) severnogo polushariya:<br />
sistematika, biologiya, biogeografiya [Chironomids of the subfamily Diamesinae (Diptera,<br />
Chironomidae) of the Northern Hemisphere: systematics, biology, and biogeography]. Dr.Sci.Thesis,<br />
FEB of Russian Academy of Sciences, Vladivostok, Russia. 518 p.<br />
Raddum, G., Erikson, L., Fott, J., Halvorsen, G.A., Heegaard, E., Kohout, L., Kifinger, B., Schaumberg, J.,<br />
Maetze, A. and Zahn, H. 2004. Recovery from acidification of invertebrate fauna at ICP Water sites in<br />
Europe and North America. ICP Waters report 75/2004, Norwegian Institute for Water Research, Oslo,<br />
Norway.<br />
Schmid, P.E. 1993. A key to the larval Chironomidae and their instars from Austrian Danube Region<br />
streams and rivers. Part I, Diamesinae, Prodiamesinae and Orthocladiinae. Federal Institute for Water<br />
Quality of the Ministry of Agriculture and Forestry, Wien.<br />
Schnell, Ø.A. and Willassen, E. 1991. Fjærmyggarten Pseudodiamesa arctica (Malloch) i to høyfjellsreservoarer<br />
(The chironomid species Pseudodiamesa arctica (Malloch) in two high mountain reservoirs).<br />
Report No 76, Freshwater Ecology and Inland Fisheries Laboratory, Department of Zoology, University<br />
of Bergen, Bergen, Norway.<br />
Willassen, E. 2011. Pseudodiamesa nivosa or arctica? A confounded story of a midge moustache and an<br />
attempt at some taxonomic orthodontics. - <strong>Chironomus</strong> Newsletter on Chironomidae Research <strong>24</strong>: XX.<br />
28
Pseudodiamesa nivosa (Goetghebuer, 1928) is not a synonym of Pseudodiamesa<br />
arctica (Malloch, 1919), but what about the separation of immature stages of these<br />
species?<br />
Eugenyi A. Makarchenko<br />
Institute of Biology and Soil Science of Russian Academy of Sciences, Vladivostok, Russia.<br />
E-mail: makarchenko@biosoil.ru<br />
As I showed in my poster “Pseudodiamesa nivosa (Goetghebuer, 1928) is not synonym of Pseudodiamesa<br />
arctica (Malloch, 1919)” at the XVIIth Chironomid symposium in Nankai University (Makarchenko,<br />
2009), both are a good and valid species and can be separated by some features of male imagines adduced<br />
below.<br />
1. LR 0.52–0.54. Basal lobe of gonocoxite with narrow and triangular or roundish triangular apex. Tergite<br />
1<br />
IX with projection of posterior-lateral angle. Gonostylus in apical 1/3 finger-shaped, with roundish apex ...<br />
P. arctica (Mall.) (Fig. 3).<br />
– LR 1 0.65–0.69. Basal lobe of gonocoxite with wide and roundish apex. Tergite IX without projection<br />
of posterior-lateral angle. Apical 1/3 of gonostylus of different shape, with beak-shaped apex ... P. nivosa<br />
(Goetgh.) (Figs 1–2).<br />
Figures 1–3. Male imagines of Pseudodiamesa nivosa (Goethebuer) from Norway (1–2) and Pseudodiamesa arctica<br />
(Malloch) from Devon Island of Canada (3). 1, 3 – total view of hypopygium, from above; 2 – gonocoxite and gonostylus.<br />
Scale bars are 100 µm.<br />
29
The hardest problem is to separate larvae of P. nivosa and P. arctica. Nobody has studied numerous specimens<br />
of larvae and has not checked variation of some larval features, namely shape, presence or absence of<br />
labral lamellae. When I studied larvae of the nivosa group, sometimes I checked labral lamellae, sometimes<br />
not, but in the same population of the same river. Yes, for deciding taxonomic problems of these and other<br />
species of Pseudodiamesa we need to revise the species by larvae, pupae and imago from various places of<br />
the Holarctic region using traditional morphological methods and karyology and DNA analysis. Also we<br />
need to study biology and ecology of these species because it can help us in deciding taxonomic problems.<br />
At first we need to get fresh material of P. nivosa and P. arctica from type localities, which for the first<br />
is the French Alps and for the second is Arctic Canada. I think it is a good joint work for an international<br />
project and a team of chironomidologists, karyologists, molecular biologists and ecologists. I think we must<br />
and can decide this interesting problem but only all together.<br />
Let us co-operate!<br />
References<br />
Makarchenko, E.A. 2009. Pseudodiamesa nivosa (Goetghebuer, 1928) is not synonym of Pseudodiamesa<br />
arctica (Malloch, 1919) (Diptera: Chironomidae). Program and abstracts XVII International<br />
Symposium on Chironomidae. July 5–10, 2009. Nankai University, Tianjin, China: 71.<br />
Typical types – a swan song? Observations on chironomids in the Linnean collections,<br />
and corresponding general considerations<br />
Martin Spies<br />
Zoologische Staatssammlung München, Münchhausenstr. 21, 81<strong>24</strong>7 München, Germany.<br />
E-mail: spies@zi.biologie.uni-muenchen.de<br />
Introduction<br />
At the 18th International Symposium on Chironomidae in Trondheim, Norway, this past July I gave a<br />
short presentation to draw attention to an online resource that has become available recently. On web<strong>page</strong>s<br />
provided by the Linnean Society of London, under http://www.linnean-online.org/view/insects/tipula.html,<br />
a list of electronic links can be found which use scientific species names made available by Linnaeus in<br />
original combination with the genus name Tipula Linnaeus. Each link leads to a series of digital images of<br />
pinned adult specimens still preserved in the Linnean collections under the corresponding species name.<br />
On a visit to London in August I then had the opportunity – thanks to the Society’s honorary curator of<br />
insects and two curators of Diptera at The Natural History Museum – to examine all Linnean specimens<br />
known to be extant and to represent species of Chironomidae. Some of the results as presented below are<br />
more or less preliminary, as so far the animals could be viewed merely at relatively low magnification, in<br />
the dry-pinned and shriveled condition in which they have been preserved all along. State-of-the-science<br />
microscopic (and possibly genetic) analysis could be performed only if permission for ‘destructive sampling’<br />
is applied for, and granted by, the Linnean Society.<br />
Chironomids in the Linnean collections<br />
The chironomid material comprises five specimens under three species names: 1 male each under Tipula<br />
littoralis L., 1758 and T. monilis L., 1758, and 1 male plus 2 females under T. plumosa L., 1758. Only one<br />
specimen per species name is accompanied by a hand-written name label (see the online images referred to<br />
above; the scale rulers shown are graded in mm) but, unless there is evidence to the contrary in a particular<br />
case, all specimens are equally eligible for consideration as original type material (Day & Fitton 1978; the<br />
second author now is the responsible curator).<br />
The male under T. littoralis is well-preserved in general, but the posterior end of the abdomen is missing<br />
and probably lost. The wing crossvein RM is darkened; the venation pattern and tibial armament are as in<br />
30
Chironomini; the fore tarsi are distinctly bearded. The taxonomically relevant evidence is consistent with<br />
what Linnaeus treated as the one of his two variant forms of T. littoralis that had been collected in forests<br />
(“habitat in nemoribus”), e.g. under the variant designators “#1136” in Linnaeus (1746) and “β” in Linnaeus<br />
(1760). An alternative possibility of connection to the name Tipula arundineti L., 1760 – raised by<br />
a second name label apparently added to the pin relatively recently – can be ruled out, as several observable<br />
morphological features differ significantly from Linnaeus’ diagnosis for T. arundineti. In any case,<br />
without the specimen’s hypopygium, current knowledge of external morphology does not allow positive<br />
identification. In summary, T. littoralis should remain a nomen dubium in <strong>Chironomus</strong> Meigen or, more<br />
conservatively, in Chironomini.<br />
The male under T. monilis L., judging from what can be evaluated in the pinned condition, looks consistent<br />
with the current and long-standing interpretation of the described species as a member of Ablabesmyia<br />
Johannsen. However, there is reason to suspect that it might not represent the taxonomic species for which<br />
the name A. monilis has been used. For instance, compared to the figures of tibial spurs in Fittkau (1962:<br />
422), especially the midleg spur configuration of the Linnean specimen is like the one ascribed to A. phatta<br />
(Egger), not like that labeled “Ablabesmyia monilis” by Fittkau. A slide-mount of the hypopygium would<br />
be necessary and likely sufficient to clarify whether this Linnean specimen belongs to any currently recognized<br />
species of Ablabesmyia and, if so, to which one.<br />
The third Linnean lot, under T. plumosa L., may raise the most critical questions. As I had suspected from<br />
the online images and mentioned at the Trondheim symposium, the three corresponding specimens appear<br />
to belong to a species in the currently recognized subgenus <strong>Chironomus</strong> (Camptochironomus). If so,<br />
they would be incompatible with the long-standing application of the name C. plumosus (L.) to a species<br />
complex or species in <strong>Chironomus</strong> (<strong>Chironomus</strong>). From inspection of the pinned material the Camptochironomus<br />
identification is certain so far only for the male in the series, but the two females have not shown<br />
any contradicting evidence. Further support comes from the information on the species’ larva and sampling<br />
sites given by Linnaeus (e.g. 1746: 333 under “#1135”, referred to by Linnaeus 1758: 587 under “plumosa”;<br />
see also the next-following reference there). The adult and larval specimens of Tipula plumosa personally<br />
examined by Linnaeus (1758) had come from two sites in the Baltic Sea along the coasts of southern Sweden<br />
(see also Hirvenoja 2006: 374, and 376 left column). Linnaeus (1746) described the larvae as showing<br />
ventral tubules of the posterior abdomen longer than the posterior parapods. This fits the ‘plumosus type’<br />
known, e.g., from larvae currently placed in Camptochironomus, but not the ‘semireductus type’ shown by<br />
brackish-water specimens subsumed in recent taxonomic concepts to which the name C. plumosus (L.) has<br />
been applied.<br />
Figure 1. <strong>Chironomus</strong> tentans Fabricius; paralectotype (ZMUC, Copenhagen), lateral view. The lectotype, also an<br />
adult female, has been slide-mounted (Hirvenoja 2006). Photo by M. Kotrba & M. Spies.<br />
31
Figure 2. <strong>Chironomus</strong> tentans Fabricius; paralectotype,<br />
dorsal view. Photo by M. Kotrba & M. Spies.<br />
As in the case of Tipula monilis, positive species<br />
identification of the Linnean specimens of T. plumosa<br />
would require microscopic evaluation of at least<br />
partial slide mounts. However, the special permission<br />
that would have to be obtained for such analysis<br />
is not the only obstacle here, as shall be discussed in<br />
the following text sections.<br />
Types to guide usage, or vice versa?<br />
The International Code of Zoological Nomenclature<br />
certainly is not the best guideline one could think<br />
of, but we cannot do without such a framework, and<br />
the Code is the only one around that has long been<br />
applied and can be applied reasonably. In its currently<br />
effective edition (ICZN 1999) the Code has<br />
attempted to serve the interests of scientific communication<br />
by allowing the application of taxon<br />
names to be influenced more by their recent usage<br />
rather than exclusively by criteria tied to the respective<br />
original publication. While this more flexible<br />
approach can be beneficial in some cases (for one<br />
example, see Spies & Sæther 2004: 27, second paragraph<br />
from top), it seems questionable whether the<br />
apparent conflict between the Linnean specimens of<br />
Tipula plumosa and the usage of <strong>Chironomus</strong> plumosus<br />
constitutes such a case.<br />
After my presentation in Trondheim I heard the ex-<br />
pected argument that any change to the application of the name <strong>Chironomus</strong> plumosus to make the latter<br />
reflect the taxonomic identity of the Linnean specimens would cause significant difficulties with accessing<br />
corresponding data published earlier not only in taxonomy, but also in ecology, faunistics, etc. While this<br />
argument, of course, has its merit in principle, it seems to miss the essential mark here. With the possible<br />
exception of identifications based on karyological/cytological evidence (i.e. on the larval giant chromosome<br />
banding patterns), ‘recent usage’ (whichever way one might define this term) of the name C. plumosus<br />
can hardly be seen as anywhere near unanimous and stable. Instead, wherever identification as C. plumosus<br />
was not based on karyological determinations, and especially wherever the methods or references used to<br />
arrive at such identification were not made sufficiently clear, one cannot determine reliably to which of the<br />
species in the <strong>Chironomus</strong> plumosus sibling species aggregate (e.g., Hirvenoja 2006) the corresponding<br />
data actually apply. Incidentally, for the same reason we used to be unable to address the issues – much<br />
more interesting to the biologist than matters of nomenclature – which species Linnaeus had encountered,<br />
whether it still lives where he found it, why not if it does not, and so on.<br />
Consequently, it is quite doubtful that there has been any “prevailing usage” (ICZN 1999) of the name C.<br />
plumosus, and thus any corresponding body of reliable biological data, that is definable and significant<br />
enough so that it could or should be protected in place of original facts such as the taxonomic identity of<br />
the type material. On the contrary, it seems that recognizing the Linnean specimens as syntypes would not<br />
decrease but even increase stability of nomenclature and quality of the corresponding dataset, by finally<br />
providing a basis for reproducible species identifications in all life stages, where previously only larvae<br />
could be assigned to a ‘cytospecies’, the name for which has not been tied to any reproducible voucher.<br />
It can be debated whether the current guidelines governing typification – e.g. the selection, designation,<br />
safe storage and consultation of physical vouchers such as type specimens – have been sufficient or require<br />
adaptation to progress in taxonomic methods. Conceivably, the assignment of type status could be made<br />
more flexible, instead of forcing taxonomists to work with or around specimens that are original types but<br />
no longer sufficiently informative. Such modifications could allow subsequent assignment of voucher status<br />
to carriers of information not included among the original type material, e.g. to specimens of other life<br />
stages reliably associated later, or to samples on the molecular level. However, this topic would be one for<br />
a separate discussion (see below) beyond the scope of the present paper.<br />
32
Evidence versus hearsay<br />
Have you ever played or watched the game variously called ‘(broken) telephone’, ‘grapevine’, ‘whisper<br />
down the lane’, etc.? In this pastime enjoyed in numerous variations the world over, information is transmitted<br />
along a chain of people, with each individual transfer involving two persons only, such that other<br />
players up or down the line cannot perceive or control the content of any transfer in which they are not<br />
directly involved. The attraction of this setup to an audience watching this game, e.g. on television, is as<br />
follows. “Errors typically accumulate in the retellings, so the statement announced by the last player differs<br />
significantly, and often amusingly, from the one uttered by the first. Some players also deliberately alter<br />
what is being said in order to guarantee a changed message by the end of it.” (http://en.wikipedia.org/wiki/<br />
Chinese_whispers).<br />
Does this, by ‘chance’, remind you of any tradition that has been going on in chironomid research in general,<br />
and especially in the large body of literature using names in <strong>Chironomus</strong>? If it does not, then here is<br />
an example which also illustrates why identification of the ‘Camptochironomus’ species represented by the<br />
Linnean specimens under Tipula plumosa is not as easy as one should expect in a taxon, in which no more<br />
than two valid species names are currently recognized in Europe (Spies & Sæther 2011).<br />
Basically, all taxonomic distinctions of Camptochironomus species to this day derive from the very brief<br />
characterizations proposed by Edwards (1929) for two species concepts under the names C. tentans Fabricius,<br />
1805 and C. pallidivittatus Malloch, 1915. Edwards (1929: 382) stated: “I find there is a small but<br />
constant difference in the hypopygium, and therefore treat the two as distinct. They superficially resemble<br />
C. plumosus.” [The latter fact, by the way, could help explain why workers after Linnaeus have misapplied<br />
Tipula plumosa for species in <strong>Chironomus</strong> s. str.] Edwards’ diagnoses hardly presented any character state<br />
distributed discretely between the two supposed species; instead, one was said to be “rather smaller and<br />
lighter”, with an “anal point rather differently shaped”, and so on. The illustrations provided (op.cit.: 381,<br />
fig. 12a, b) do not show any full hypopygium as given for many other species in that work, but limit themselves<br />
to details of the anal point and flanking parts of the anal tergite. As could have become obvious a long<br />
time ago, the appearance of such parts in a microscope preparation can vary significantly with orientation of<br />
the specimen in the mount, degree of maceration, pressure when applying the cover slip, etc. No number of<br />
specimens analysed was given by Edwards to support his claim of a difference “constant” across variation<br />
to be expected among preparations studied, individuals in a sample, or populations of one or more species.<br />
Apparently, Edwards (1929) had not seen any type of C. tentans Fabricius or C. pallidivittatus Malloch, but<br />
merely applied those names from the literature to his British material. In spite of this and the quite obviously<br />
less than water-tight identifications (see, e.g., Hein & Schmulbach 1971: 458), everyone in the subsequent<br />
long chain of authors on these and other Camptochironomus species (Townes 1945, Beermann 1955,<br />
Hein & Schmulbach 1971, Sæther 1977, Shobanov et al. 1999, Langton & Pinder 2007 – to name just a few<br />
cornerstone examples) have merely continued the game started (in this case) by Edwards, working mostly<br />
from more or less uncritically accepted information handed down along that line of letters on patient paper.<br />
To my knowledge I am now the first ever to gather all relevant voucher material (in this case by Fabricius,<br />
Malloch, and Edwards) for direct comparison. [Incidentally, I also seem to be the first chironomid worker<br />
since Linnaeus to have looked at his specimens of non-biting midges.]<br />
As shown repeatedly (e.g. in Spies & Sæther 2004) there is one case like this after another that would have<br />
to be looked at seriously – which would be necessary if, in contrast to audiences watching ‘whisper’ games,<br />
our purpose in chironomid research runs a little deeper than trivial amusement. Taking such a hard look,<br />
then, one cannot help but find that anything we present without truly reproducible evidence is tantamount<br />
to hearsay or speculation, i.e. essentially unreliable, and does not qualify to be labeled and employed as science.<br />
If significant portions of the data in any system such as chironomid research are non-reproducible, e.g.<br />
because they are not based on evidence that remains accessible and observable, then the system is degraded<br />
from one built on, and aiming for, evidence-based knowledge to one running on mere beliefs. If members of<br />
a research community like ours are inclined or forced to blindly follow what they have seen in publications<br />
only, rather than being willing and able to critically test what they examine directly, then entropy inadvertently<br />
growing underneath a seemingly orderly surface is bound to cause the corresponding data and system<br />
to deteriorate rather than increase in practical value.<br />
33
Towards a discussion<br />
Against the above background, please consider the following questions.<br />
— Has there not been long-standing, significant imprecision and inconsistency in the application of the<br />
name <strong>Chironomus</strong> plumosus?<br />
— Are situations like this not exactly what typification has been invented for, and with good reason?<br />
— Does reinterpretation of the name Tipula plumosa L. in accordance with original type specimens not<br />
constitute a good chance to increase or, rather, finally establish nomenclatural stability in this case, in which<br />
we have had only superficial semblance of stability, at best?<br />
— If using and quoting taxonomic names with credit to the authors who have made them available is supposed<br />
to honor these people, would Linnaeus consider himself honored if we knowingly apply the name<br />
<strong>Chironomus</strong> plumosus to a species different from the one he coined it for?<br />
— How can museums and other institutions – to which many of us would like to be able to look for employment<br />
– justify to funding agencies, etc., the efforts necessary to build, maintain or even improve the<br />
collections we would need and like to have at our disposal, if we do not actively and scientifically support<br />
them in this? Which parts of such collections could be more important to use and support than the holdings<br />
of type material?<br />
— Are we allowing too much of chironomid research to run like a game of ‘whispering down the lane’? Or<br />
are we doing enough to make and keep every important bit we publish scientifically reproducible, so that<br />
we and others depending on input from us may work with a well-founded and well-growing body of reliable<br />
and useful information that deserves to be called knowledge?<br />
It would be most welcome if answers to these questions, arguments supporting or countering any of those<br />
presented here, or any other constructive contributions, could be exchanged in a wide-open discussion. One<br />
appropriate forum for this could be the Chironomidae mailing-list (to sign up see https://lists.ansatt.ntnu.no/<br />
vm.ntnu.no/info/chironomidae), even though so far the list has seen as poor participation as several other<br />
potentially highly useful services offered to the chironomid ‘community’.<br />
In closing, I would like to declare that responses, as well as silence, on the issues raised in this contribution<br />
will play no small part in determining whether and how I will proceed concerning the Linnean chironomids<br />
and similar matters. Consequently, if you have an opinion and would like to see it count, then by all means<br />
do let it be known.<br />
References<br />
Beermann, W. 1955. Cytologische Analyse eines Camptochironomus-Artbastards. I. Kreuzungsergebnisse<br />
und die Evolution des Karyotypus. – Chromosoma 7: 198-259.<br />
Day, M.C. and Fitton, M.G. 1978. Re-curation of the Linnaean Hymenoptera (<strong>Insect</strong>a), with a reassessment<br />
of the taxonomic importance of the collection. – Biological Journal of the Linnean Society, 10: 181-198.<br />
Edwards, F.W. 1929. British non-biting midges (Diptera, Chironomidae). – Transactions of the Royal Entomological<br />
Society of London 77: 279-430.<br />
Fittkau, E.J. 1962. Die Tanypodinae (Diptera: Chironomidae). (Die Tribus Anatopyniini, Macropelopiini<br />
und Pentaneurini). – Abhandlungen zur Larvalsystematik der Insekten 6: 1-453.<br />
Hein, J. and Schmulbach, J.C. 1971. Intraspecific and interspecific breeding behavior of <strong>Chironomus</strong> pallidivittatus<br />
(Dipt., Chironomidae). – Canadian Entomologist 103: 458-464.<br />
Hirvenoja, M. 2006. Contributions to the taxonomy and ecology of the <strong>Chironomus</strong> plumosus sibling species<br />
aggregate (Diptera: Chironomidae): the brackish water populations in the Tvärminne area, Finland.<br />
– Entomologica Fennica 17: 373-380.<br />
ICZN = International Commission on Zoological Nomenclature. 1999. International Code of Zoological<br />
Nomenclature. Fourth Edition. – International Trust for Zoological Nomenclature, London. xxix<br />
+ 306 p.<br />
34
Langton, P.H. and Pinder, L.C.V. 2007. Keys to the adult male Chironomidae of Britain and Ireland. Volumes<br />
1 and 2. - Freshwater Biological Association, Scientific Publication 64: 239 + 168 p.<br />
Linnaeus, C. 1746. Fauna svecica sistens animalia sveciae regni: ... – L. Salvius, Stockholm. [xxviii] +<br />
411 p., 2 pls.<br />
Linnaeus, C. 1758. Systema naturae per regna tria naturae, ... Tomus I. Editio decima, reformata. – L.<br />
Salvius, Stockholm. [iv] + 8<strong>24</strong> p.<br />
Linnaeus, C. 1760 (“1761”). Fauna svecica sistens animalia sveciae regni: ... Editio altera, auctior – L.<br />
Salvius, Stockholm. [48] + 578 p.<br />
Sæther, O.A. 1977. Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies,<br />
keys. – Bulletin of the Fisheries Research Board of Canada 197: 1-209.<br />
Sæther, O.A. and Spies M. 2011. Fauna Europaea: Chironomidae – In: Beuk, P. & T. Pape (eds.) Fauna<br />
Europaea: Diptera Nematocera. Fauna Europaea version 2.4. Available at: http://www.faunaeur.org/<br />
Shobanov, N.A., Kiknadze, I.I. and Butler, M.G. 1999. Palearctic and Nearctic <strong>Chironomus</strong> (Camptochironomus)<br />
tentans (Fabricius) are different species (Diptera: Chironomidae). – Entomologica scandinavica<br />
30: 311-322.<br />
Spies, M. and Sæther, O.A. 2004. Notes and recommendations on taxonomy and nomenclature of Chironomidae<br />
(Diptera). – Zootaxa 752: 1-90.<br />
Townes, H.K., Jr. 1945. The Nearctic species of Tendipedini (Diptera; Tendipedidae (= Chironomidae)). –<br />
American Midland Naturalist 34: 1-206.<br />
CHIRONOMUS Newsletter now compliant with the International Code of<br />
Zoological Nomenclature<br />
Torbjørn Ekrem<br />
Co-editor <strong>Chironomus</strong> Newsletter on Chironomidae Reseach<br />
Museum of Natural History and Archaeology, Norwegian University of Science and Technology, Trondheim,<br />
Norway. E-mail: torbjorn.ekrem@ntnu.no<br />
It is a pleasure to inform you that as of this issue, the newsletter complies with article 8.6 of the International<br />
Code of Zoological Nomenclature. This means that texts published in the newsletter are to be regarded as<br />
a published work also for the purposes of zoological nomenclature, and that we now welcome papers that<br />
involve nomenclatorial acts. In order to comply with article 8.6, copies of the newsletter must be deposited<br />
in at least five major publicly accessible libraries. This and future issues of the CHIRONOMUS Newsletter<br />
will therefore be printed in a limited number of paper copies and distributed to the libraries listed on <strong>page</strong><br />
two of this issue. We are very pleased that all libraries we contacted were willing to receive and store copies<br />
of the CHIRONOMUS Newsletter and look forward to receive more taxonomy manuscripts for our Current<br />
Research section.<br />
35