Chironomus 23 (Nov 2010) - Insect Division

CHIRONOMUS 

Newsletter on Chironomidae Research 

No. 23 ISSN 0172-1941 November 2010 

CONTENTS 

Editorial: 

Chironomidae and the tree of life 3 

Wolfgang Friedrich Wülker, on the 

occation of his 85th birthday 4 

Professor Iya I. Kiknadze: 80th 

birthday and 55 years in science 

and teaching 10 

Arthur Desmond Harrison 

- an appreciation 14 

Paul Freeman - an appreciation 

of his studies on Chironomidae 17 

Current Research 20 

Hamerlik, L., Christoffersen, K. S. 

& Brodersen, K. P. Short comment 

on chironomid assemblages and 

stratigraphy of high altitude lakes 

from Tibet 20 

Sonoda, K., Malaquias, J. 

& Vettorazzi, C. Chironomidae 

(Insecta, Diptera) from Alto 

Paranapanema Basin, 

Southeastern Brazil 25 

Short Communications 33 

Currrent Bibliography 39 

Sculpture by David Wilson close to the shore of Loch Leven at Kinross. Photo Sarah Stenhouse

CHIRONOMUS Newsletter on Chironomidae Research 

Editors 

Torbjørn EkrEm, Museum of Natural History and Archaeology, Norwegian University of Science and 

Technology, NO-7491 Trondheim, Norway 

Peter H. Langton, 5 Kylebeg Avenue, Mountsandel, Coleraine, Co. Londonderry, Northern Ireland, BT52 

1 JN, Northern Ireland 

Current Bibliography 

Odwin HoffricHtEr, Institut für Biologie I, Albert-Ludwigs-Universität Freiburg, Hauptstrasse 1, D-79104, 

Germany 

The CHIRONOMUS Newsletter on Chironomidae Research is devoted to all aspects of chironomid research 

and aims to be an updated news bulletin for the Chironomidae research community. The newsletter is 

published yearly in October, is open access, and can be downloaded free from this website: http://www. 

ntnu.no/ojs/index.php/chironomus. 

Research articles for the CHIRONOMUS Newsletter are subject to peer-review. The newsletter also contains 

a current bibliography that is maintained by Odwin Hoffrichter, please send complete references of your 

new Chironomidae publications directly to him. 

Contributions to CHIRONOMUS Newsletter on Chironomidae Research should be submitted online through 

the online journal system: http://www.ntnu.no/ojs/index.php/chironomus following the author guidelines. 

Submission deadline for contributions to the newsletter is July 1. 

Would you like to see your picture on the front page? Please send us your favourite midge photograph or 

drawing (Torbjorn.Ekrem@vm.ntnu.no). 

Front page layout: Chironomid in title from photograph by Steve Marshall, Graphic design by Kolbjørn Skarpnes, 

NTNU Information Division. 

Front page photo: Sculpture by David Wilson, commissioned by Scottish Natural Heritage, at Burleigh Sands, Kinross, 

close to the shore of Loch Leven. Photograph by Sarah Stenhouse, SEPA Aberdeen (used with permission and adapted 

from Sue McBean (in Langton & McBean, 2010 Dipterists Digest, in press)). 

2

Editorial 

Chironomidae and the tree of life 

The importance of understanding the genealogical relationships between populations, species and species 

groups can hardly be exaggerated. As reconstructions of evolutionary pathways through time, phylogenetic 

trees provide a framework on which biological hypotheses can be tested; without robust phylogenies we 

will be unable to fully understand the processes behind speciation and biogeographical distributions. It is 

therefore perhaps not surprising that research devoted to systematics and phylogenetics on various branches 

of the tree of life has found increased interest over the last 10-20 years. Easier access to molecular data for 

use in phylogenetic analyses and the development of new analytical tools has promoted what has become a 

huge production of statistically testable hypotheses of evolutionary relationships. 

Diptera is no exception and it was remarkable to see the number of papers dealing with phylogenetic analyses 

at the 7th International Congress of Dipterology in Costa Rica in August this year. In addition to being 

numerous, most of the phylogenetic studies that were presented at this conference used multiple markers, 

often in combination with morphological characters, to test for congruence and investigate the evolution of 

morphological and ecological traits. Datasets comprising more than 2000 characters were not uncommon 

and some even approached 6000! It is obvious in which direction phylogenetic research is going; what was 

regarded as a thorough study with a couple of genes yesterday will be a small pilot study tomorrow. - So 

also for the Chironomidae. 

But, as for other dipteran taxa, morphological characters in phylogenetics are far from dead. As an example, 

Art Borkent at the Diptera Congress presented new evidence from pupal morphology that chironomids 

should be regarded as sister to all the other families in the Culicomorpha. The relationship has been backed 

by Pete Cranston et al.’s molecular phylogeny using molecular characters from four different genes. The 

latter study also has revealed Buchonomyiinae as sister to all other Chironomidae subfamilies, a relationship 

conforming to Murray and Ashe’s earlier observation on the general plesiomorphous morphology of 

adult, pupa and egg of Buchonomyia thienemanni (Murray & Ashe 1981; Ashe & Murray 1983). There are 

many groups with unresolved relationships, however, and there will definitely be many interesting results 

from phylogenetic studies within our family in the years to come. 

I hope that some of these will be presented at the next Chironomidae symposium in Trondheim and that they 

in concert with research results in taxonomy, ecology, genetics, cytology, palaeolimnology and toxicology 

will guarantee for a successful meeting. Please read more about the symposium and how to preregister in 

the Short Communications section in this newsletter. We look forward to see you there! 

Torbjørn Ekrem, Museum of Natural History and Archaeology, Norwegian University of Science and Technology, 

Norway. E-mail: Torbjorn.Ekrem@vm.ntnu.no 

References 

Ashe P., Murray D. A. 1983. Observations on and descriptions of the egg-mass and eggs of Buchonomyia 

thienemanni Fitt. (Diptera: Chironomidae). In: Roback S. S., editors. Proceedings of the 8th International 

Symposium on Chironomidae. Philadelphia: Mem. Am. Entomol. Soc. 34, p 3-13. 

Murray D., Ashe P. 1981. A description of the pupa of Buchonomyia thienemanni Fittkau, with notes on its 

ecology and on the phylogenetic position of the subfamily Buchonomyiinae (Diptera, Chironomidae). 

- Spixiana 4: 55-68. 

3

Wolfgang friedrich Wülker, on the occasion of his 85 th Birthday 

Wolfgang Wülker was born on 25 September 1925 

at Frankfurt, Germany. 

In 1952 he began working as a scientific assistant 

at the Hydrobiological Station Falkau 

(later the Limnological Institute of the University 

of Freiburg, Walter Schlienz-Institut). His initial 

publications were on fish, but papers on chironomids 

began appearing from 1956. He was at Falkau 

for about 10 years and added to the collection of 

the station both specimens from the Black Forest 

area, and specimens collected on his research trips 

to Spain (1954), Fennoscandinavia (1956), Sudan 

(1963), USA and Canada (1964). 

He did his Habilitation in Zoology and Limnology 

on intersexuality in Chironomus and the biology 

of Sergentia at the University of Freiburg, in 

1960. Then in 1962 he joined the University of 

Freiburg. 

After his official retirement Wolfgang continued to 

work at the University, later transferring his laboratory 

to his home. 

Wolfgang was involved in collaborations with colleagues 

in many countries during his long career, 

some of which lasted for many years and produced 

many publications. A photograph of Wolfgang 

with Jim Sublette in Portales, N.M. in 1964 was 

reproduced on page 6 of volume 21 of this newsletter. 

His collaboration with Jim Sublette, Jon 

Martin and later Mac Butler, has produced 16 papers 

from 1967 to the present; that with the Russian 

workers, notably Iya Kiknadze and her group, 

15 since about 1993; seven with the Swiss group 

of Adolf Scholl; four with György Dévai and colleagues 

from Hungary; and numerous others with 

students or colleagues from Germany. Wolfgang’s 

full bibliography of chironomid publications is attached. 

Jon Martin 

Melbourne 

My wife, Mary, and I first met Wolfgang and his 

wife, Dorothea, in Vienna at the 1959 SIL meeting. 

It was a most auspicious meeting for us which 

laid the groundwork for a collaborative 40+ years 

research team of Wuelker, Sublette and Sublette, 

which shortly thereafter was joined by Jon Martin 

who was then doing a postdoctoral in Ottawa 

at Canada Agriculture. The Sublettes were able 

4 

to get grants to visit Wuelker in Freiburg. In their 

brief stay there Jim and Wolfgang held endless discussions 

on midge taxonomy and Mary focused on 

getting a camera lucida drawing of some of the Keyl 

arms. Wolfgang became more entranced with the 

possibility of using polytene chromosomes in Chironomid 

taxonomy and phylogeny and spent some 

time in other labs in Germany becoming acquainted 

with methodology and cytotaxonomy in general. 

Wolfgang Wülker. Photo NN 

In 1965 Wolfgang was still working in the field of 

mermithid parasitism in Chironomidae and he was 

funded to come to the U.S. to attempt cross-inoculation 

of mermithid infective larvae into Nearctic 

samples. Wolfgang had worked with Sergentia and 

wanted to locate nearctic populations. He knew 

that John Stahl at the University of Indiana had 

worked on Sergentia in Crooked Lake. The Sublette 

family and Wuelker family met up in Indiana 

and spent a week together at Shafer Lake, where 

Jim’s brother had lakeside acreage and a home. 

The Wuelkers pitched their camping trailer on the 

lawn and Jim and Wolfgang set out in search of 

Sergentia. We located a Crooked Lake on a topographic 

map, near Ft. Wayne. However, sampling 

that lake showed it was a shallow lake with only 

plumosus type larvae. The Wuelkers were anxious

to start a grand tour of the west and bought an old 

VW microbus in Monon from a dealer recommended 

by Jim’s brother. Seven miles from where the 

dealership was, the motor fell out of the old VW. 

Jim’s brother did a bit of arm twisting and they replaced 

the motor gratis. THEN the Wuelkers set of 

on the grand trip. After touring all of the national 

parks in the western United States, the Wuelkers 

ended up in Portales, NM where Jim was teaching. 

The University provided a guest apartment for the 

Wuelkers and they stayed there about 4 months. 

Wolfgang’s infective larvae had moulted once as 

I recall but were still infective. We attempted to 

infect them into a plumosus type larva and a decorus 

type larva and a Glyptotendipes larva. No 

Sergentia were available. Infections occurred at a 

low rate but in all cases the mermithid died before 

moulting again. 

Mary acquired the technique of making polytene 

chromosome squashes and produced several thousand 

over the next several years. Over the years 

Wolfgang returned again to Portales to sort through 

the growing collection of squashes accumulated 

there. We were most fortunate to make the association 

and receive the help from Patrick L. Hudson, 

U.S. Fish and Wildlife Service, who sent innumerable 

mature, fixed larvae with associated adults 

and larval/pupal skins. Much of this material was 

examined by Wolfgang and later by Jon Martin. 

Jon and Wolfgang were responsible for cytology 

and Jim and Mary, the larva, pupa and adults. Jim, 

in ENMU administration, was instrumental in establishing 

an in-house journal, Studies in Natural 

Science (Portales) which served as the publication 

vehicle for several of these joint studies. 

Jim Sublette 

Tucson 

Chironomidae publications of W. Wülker 

1 Wülker, W. 1956. Zur Kenntnis der Gattung 

Psectrocladius Kieff. (Dipt., Chironomidae). 

- Arch. Hydrobiol., Suppl. 24: 1-66 

2 Wülker, W. 1957. Eine spanische Halliella 

(Dipt.Chironomidae). - Arch. Hydrobiol., 

Supp1. 24: 281-296 

3 Wülker, W. 1957. Ober die Chironomiden der 

Parakiefferiella-Gruppe. (Diptera, Tendipedidae, 

Orthocladiinae). - Beitr. Entomol. 7: 411- 

429. 

4 Wülker, W. 1958. Die Bedeutung der Chironomiden 

fiir die limnologisch-tiergeographische 

Charakterisierung des Hochschwarz-waldes. 

5 

- Verh. int. Ver. Limno1. 13: 805-813. 

5 Wülker, W. 1958. Parasitäre Intersexualität bei 

Chironomiden des Sch1uchsees. - Gewässer 

und Abwässer (Krefeld) Jhg. 20: 61-67. 

6 Wülker, W. 1959. Diamesarien-Studien 

(Dipt.,Chironomidae) im Hochschwarzwald. 

- Arch. Hydrobiol., Suppl. 24: 338-360. 

7 Wülker, W. 1959. Drei neue Chironomiden- 

Arten (Dipt.) und ihre Bedeutung fiir das Konvergenzproblem 

der Imagines und Puppen. 

- Arch. Hydrobiol., Supp1. 25: 44-64. 

8 Wülker, W. 1960. Neue Ergebnisse zur parasitären 

Intersexualität der Chironomiden. 

- Naturwiss. 47: 21-22. 

9 Wülker, W. 1960. Studien zur Morphologie, 

Biologie und Verbreitung der Gattung Sergen 

tia Kieff. (Dipt., Chironomidae). - Arch. Hyd 

robiol., Suppl. 25: 307-331. 

10 Wülker, W. 1961. Lebenszyklus und Vertikalverteilung 

der Chironomide (Dipt.) Sergentia 

coracina Zett. im Titisee. - Verh. int. Ver. Lim 

nol. 14: 962-967. 

11 Wülker, W. 1961. Untersuchungen über die 

Intersexualität der Chironomiden (Dipt.) nach 

Paramermis-Infektion. - Arch. Hydrobiol., 

Suppl. 25: 127-181. 

12 Wülker, W. 1961. Parasitäre und nicht parasitäre 

Zwischengeschlechtlichkeit bei Chironomiden 

(Dipt.). - Zool. Anz., Suppl. 25: 132-139. 

13 Wülker, W. 1962. Bedeutung des Wirtes im Leben 

tierischer Endoparasiten. - Naturw. Runds 

chau 15: 469-474. 

14 Wülker, W. 1963. Parasitologische und biochemische 

Verwandtschaft in der Gattung Chi 

ronomus (Dipt.). - Naturwiss. 50: 49-50. 

15 Wülker, W. 1963. Prospects for biological control 

of pest Chironomidae in the Sudan. - Rep. 

World Organization (WHO/EBL) 11: 1-23. 

16 Wülker, W. 1963. Investigations on the chironomid 

fauna of the Nile (Khartoum, Wadi 

Medani, Sennar, Wadi A Halfa). - Ann. Rep. 

Hydrobio1.Res.Unit. Khartoum 9-10: 20-21. 

17 Wülker, W. 1964. Parasite-induced changes of 

external and internal sex characters in insects. 

- Exper. Parasitol. 15: 561-597. 

18 Wülker, W. 1965. Der Mechanismus des Eindringens 

parasitärer Mermithiden (Nematoda) 

in Chironomus-Larven (Dipt., Chironomidae). 

- Z. Parasitenkd. 26: 29-49.

19 Wülker, W., Sublette, J.E., Martin, J. 1967. 

Zur Cytotaxionomie nordamerikanischer Chi 

ronomus-Arten. - Ann. Zool. Fenn. 5: 155- 

158. 

20 Wülker, W., Götz, P. 1968. Die Verwendung 

der Imaginalscheiben zur Bestimmung des Entwicklungszustandes 

von Chironomus-Larven 

(Dipt.). - Z. Morphol. Tiere 62: 363-388. 

21 Wülker, W., Maier, W., Bertau, P. 1969. Untersuchungen 

uber die Hämolymphproteine 

der Chironomiden (Dipt.). - Z. Naturforsch. 

24:110-116. 

22 Wülker, W. 1969. Parasitismus des Nematoden 

Gastromermis rosea in Chironomus anthraci 

nus (Diptera). Unterrichtsfilm C 1024, Institut 

Wiss.Film Göttingen. - Publ. Wiss. Film (Göttingen) 

7: 299-312 1974. 

23 Wülker, W., Winter, G. 1970. Untersuchungen 

über die Ultrastruktur der Gonaden von Chiro 

nomus (Dipt.). 1. Normalentwicklung der Ovarien 

im 4. Larvenstadium. - Z. Zellforsch. 106: 

348-370. 

24 Wülker, W. 1971. Untersuchungen über die 

Ultrastruktur der Gonaden von Chironomus 

(Dipt.). 2. Ovarstruktur nach Schädigung durch 

parasitäre Mermithiden. - Z. Parasitenkd. 36: 

73-93. 

25 Krieger-Wolf, E., Wülker, W. 1971. Chironomiden 

(Diptera) aus der Umgebung von Freiburg 

Bg. (mit besonderer Berücksichtigung der 

Gattung Chironomus). - Beitr. naturk. Forsch. 

SüdwDt1. 30: 133-145. 

26 Wülker, W., Sublette, J.E., Sublette, M.F., Martin, 

J. 1971. A review of the genus Chironomus 

(Chironomidae,Diptera) 1. The staegeri-group. 

- Stud. Nat. Sci. (Portales) 1: 1-89. 

27 Martin, J., Wülker, W. 1971. Inversion polymorphism 

in Chironomus staegeri Lundbeck. 

- Can. J. Gen. Cytol. 13: 306-321. 

28 Wülker, W. 1973. Revision der Gattung Chi 

ronomus Meig. 1II.Europäische Arten des 

thummi-Komplexes. - Arch. Hydrobiol. 72: 

356-374. 

29 Wülker, W., Klötzli, A. 1973. Revision der 

Gattung Chironomus Meig. IV. Arten des lacu- lacunarius 

(commutatus)-Komplexes. - Arch. Hyd 

robio1. 72: 474-489. 

30 Wülker, W., Martin, J. 1974. A review of the 

genus Chironomus (Diptera, Chironomidae). 

VI. Cytology in the Chironomus maturus-complex. 

- Stud. Nat. Sci. (Portales) l(9.:1-17 

6 

31 Martin, J., Wülker, W. 1974. A review of the 

genus Chironomus (Diptera, Chironomidae). 

VIII. Chironomus stigmaterus Say, Cytology. 

- Stud. Nat. Sci. (Portales) 111.: 1-17. 

32 Martin, J., Wülker, W., Sublette, J.E. 1974. Evolutionary 

cytology in the genus Chironomus 

(Diptera, Chironomidae). - Stud. Nat. Sci. 

(Portales) l12.: 1-12. 

33 Wülker, W. 1974. Parasitismus und Symbiose, 

Studieneinheit 19, Funkkolleg Biologie, Systeme 

des Lebendigen, In: Todt, D. (ed.) Begleit 

texte. Beltz-Verlag, Weinheim, Basel. 

34 Wülker, W. 1974. Die groáen Kreisläufe der 

Natur. Studieneinheit 23, Funkkolleg Biologie, 

Systeme des Lebendigen, In: Todt, D. (ed.) Be Be 

gleittexte. Beltz-Verlag Weinheim Basel. 

35 Wülker, W. 1975. Parasite-induced castration 

and intersexuality in insects. In: Reinboth, R. 

(ed.) Intersexuality in the animal kingdom. 

Springer-Verlag Heidelberg. p. 121-134. 

36 Wülker, W. 1976. Influence of mermithids 

(Nematoda) on insect imaginal discs. - Proc. 

1st. Intern Coll. Invertebr. Patho1. Queens 

Univ., Kingston Ontario, p. 255-258. 

37 Wülker, W. 1978. Parasitäre Einflüsse auf undifferenzierte 

Gewebe. - Z. Parasitenkd. 57: 

255-257. 

38 Wülker, W. 1978. Morphology and ultrastructure 

of the ejaculatory duct in normal, parasitized 

and experimentally (UV)-treated Chi 

ronomus. - Acta Univ. Carol. Biol. Praha, 1-2: 

285-290. 

39 Wülker, W., Rössler, R., v. Essen, S. 1979. 

Studies on the development of imaginal discs 

of Chironomus (Dipt.). The female genital disc 

in abdominal segment VIII. - Zool. Jb. Anat. 

101: 136-153. 

40 Wülker, W. 1980. Basic pattern in the chromosome 

evolution of the genus Chironomus 

(Dipt.). - Z. zool. Syst. Evolutionsforsch. 18: 

112-123. 

41 Ryser, H.M., Geiger, H.J., Scholl. A., Wülker, 

W. 1980. Untersuchungen über die Verbreitung 

der Zuckrnückengattung Chironomus in der 

Schweiz, mit besonderer Berücksichtigung von 

drei cytologisch nicht beschriebenen Arten. In: 

Murray, D.A. (ed.) Ecology, Systematics, Cy 

tology and Physiology. Pergamon-Press Oxford 

New York. pp. 17-24. 

42 Schmidt, K., Wülker, W. 1980. Parasitäre Intersexualität 

des Johnstonschen Organes in den

Antennen von Chironomus (Dipt.). - Z. Para 

sitenkd. 64: 1-15. 

43 Wülker, W., Ryser, H.M., Scholl, A. 1981. Revision 

der Gattung Chironomus Meigen (Dipt.). 

V1. C. holomelas Keyl, C. saxatilis n.sp., C. 

melanescens Keyl. - Rev. Suisse Zool. 88: 903- 

924. 

44 Wülker, W., Eisele, H. Rössler, R. 1982. Tar- Tarsal 

sensilla of Chironomus (Diptera: Chironomidae): 

number, parasitogenic changes, 

ultrastructure and function. - 1nt. J. Insect 

Morphol. Embryol. 11(2.: 137-146. 

45 Ryser, H.M., Scholl, A., Wülker, W. 1983. 

Revision der Gattung Chironomus Meigen 

(Diptera) VII: C. muratensis n.sp. und C. nudi 

ventris n.sp., Geschwisterarten aus der plumosus-Gruppe. 

- Rev. Suisse Zool. 90: 299-316. 

46 Wülker, W., Butler, M.G. 1983. Karyosystem- 

Karyosystematics 

and morphology of Northern Chironomus 

(Diptera: Chironomidae): Freshwater species 

with larvae of the salinarius-type. - Ent. scand. 

14: 121-136. 

47 Wülker, W., Ryser, H.M., Scholl. A. 1983. 

Revision der Gattung Chironomus Meigen 

(Diptera), VIII. Arten mit Larven des fluviatilis-Typs 

(obtusidens-Gruppe): C. acutiventris 

n.sp. und C. obtusidens Goetgh. - Rev. Suisse 

Zool. 90: 725-745. 

48 Dévai, G., Wülker, W., Scholl, A. 1983. Revision 

der Gattung Chironomus Meigen (Diptera). 

IX. C. balatonicus sp.n. aus dem Flachsee 

Balaton (Ungarn). - Acta Zool. Acad. Sci. 

Hung. 29: 357-373. 

49 Wülker, W. 1983. Mermithiden (Nematoda) 

- Chironomus spec. (Insecta, Diptera) Kopulation 

und Eiablage der Parasiten, Wirtsfindung 

und Eindringvorgang, parasitäre Intersexualität, 

humorale Einkapselung des Parasiten- In: 

Boeckeler, W., Wülker, W. (eds.) Parasitolo 

gisches Praktikum, Verlag Chemie, Weinheim 

-Deerfield Beach, Florida -Basel. 

50 Wülker, W., Lörincz, G., Dévai, G. 1984. A 

new computerized method for deducing phylogenetic 

trees from chromosome inversion data. 

- Z. zool. Syst. Evolutionsforsch. 22: 86-91. 

51 Wülker, W. 1985. Karyosystematics and morphology 

of two North European species of the 

Chironomus maturus-complex (Diptera: Chironomidae). 

- Entomol. Gener. 10: 125-132. 

52 Wülker, W. 1985. Changes in behaviour, flight 

tone and wing shape in nematode-infested Chi 

7 

ronomus (Insecta, Diptera). - Z. Parasitenkd. 

71: 409-418. 

53 Ryser, H.M., Wülker, W., Scholl, A. 1985. Revision 

der Gattung Chironomus Meigen (Diptera). 

X. Lobochironomus n.subg. (C. montuo 

sus n.sp., C. storai Goetgh., C. mendax Stora). 

- Rev. Suisse Zool. 92: 385-404. 

54 Morisch, U., Wülker, W. 1987. Formation of 

the cerci, abdominal Segment X and postgenital 

plate in the genital imaginal discs of female 

larvae and pupae in Chironomus (Diptera). 

- Ent. scand., Suppl. 29: 91-96. 

55 Kaiser, H., Wülker, W., Skofitsch, G. 1987. 

Limnomermis anthracini n.sp. and L. bathophi 

li n.sp., sympatric species of Mermithidae 

(Nematoda) in different Chironomid (Diptera) 

hosts. - Zoo1 Jb. Syst. 114: 141-156. 

56 Wülker, W. 1987. Der Karyotyp von Chi 

ronomus (Chaetolabis) macani Freeman 1948. 

- Entomol. Gener. 12: 281-286. 

57 Webb, C.J., Martin, J., Wülker, W, 1987. Ultrastructure 

of larval ventromental plates of 

European and North American representatives 

of Chironomus Meigen (subgenus Chaetolabis 

Townes) (Diptera: Chironomidae). - Ent scand. 

18, 205-211. 

58 Wülker, W., Herrmann, I. 1987. Die weiblichen 

Gonodukte in normalen und parasitierten Chi 

ronomus (Dipt.) Imagines. - Spixiana Suppl. 

14: 17-24. 

59 Wülker, W., Sublette, J.E., Morath, E., Martin, 

J. 1989. Chironomus columbiensis n.sp. 

in South America and Chironomus anonymus 

Williston in North America- closely related 

species. - Stud. neotrop. Fauna Environ. 34: 

121-136. 

60 Dévai, G., Miskolczi, M., Wülker, W. 1989. 

Standardization of chromosome arms B, C and 

D in Chironomus (Diptera: Chironomidae). 

- Acta Biol. Debrecina., Suppl. Oeco1. Hungar. 

2: 79-92. 

61 Wülker, W., Dévai, G., Dévai, I. 1989. Computer 

assisted studies of chromosome evolution 

in the genus Chironomus (Dipt.). Comparative 

and integrated analysis of chromosome arms A, 

E and F. - Acta Biol. Debrecina Suppl. Oecol. 

Hungar. 2: 373-387. 

62 Wülker, W., Morath, E. 1989. South American 

Chironomus (Dipt. ) - karyotypes and their relations 

to North America. - Acta Biol. Debre 

cina, Suppl. Oecol. Hungar. 2: 389-397.

63 Wülker, W., Sublette, J.E., Martin, J. 1990. Chi 

ronomus utahensis Malloch and Chironomus 

harpi new species and their karyosystematic 

relationships to other species in the decorusgroup 

of Chironomus. - Spixiana 14: 71-94. 

64 Wülker, W. 1991. Chromosomal and morphological 

differences between Chironomus tenui 

stylus Brundin 1949 and C. longistylus Goetghebuer 

1921 in Fennoscandia and the USA. 

Ent. scand. 22: 231-24. 

65 Wülker, W. 1991. Chironomus fraternus n.sp. 

and C. beljaninae n.sp., sympatric sister species 

in Fennoscandian reservoirs. - Ent. Fenn. 

2: 97-109. 

66 Wülker, W., Weiser, J. 1991. Helmichia glan 

dulicola n. sp. (Microspora, Thelohanidae): 

Morphology, development and influence on 

salivary glands of Chironomus anthracinus 

(Diptera, Chironomidae). - Parasitol. Res. 

77:335-340. 

67 Wülker, W. 1992. Problems of karyosystematics 

in the Genus Chironomus Meigen (In Russian). 

In: Biologie der Siissgewässer. Inform. 

Bull. Russ. Acad.Sci. St. Petersburg 93:57-62. 

68 Kiknadze, I.I., Siirin, M.I., Wülker, W. 1993. 

Siberian species of the riihimakiensis-group in 

the genus Chironomus (Diptera, Chironomidae) 

1. Karyotypes and morpho1ogy. - Neth 

er1. J. Aquat. Ecol. 26: 163-171. 

69 Kiknadze, I.I., Siirin, M.I., Wülker, W. 1994. 

Siberian species of the riihimakiensis-group of 

the genus Chironomus (Diptera, Chironomidae) 

2. Inversion polymorphism and cytophy- 

1ogeny. - Spixiana Suppl. 20:115-125. 

70 Wülker, W. 1996. Chironomus pilicornis Fabricius 

1787 and C. heteropilicornis n.sp (Diptera, 

Chironomidae) in Fennoscandian reservoirs: 

karyosystematic and morphological resu1ts. 

- Aquat. Insects 18:209-221. 

71 Wülker, W. 1996. Zoogeographische Verbreitung 

sibirischer Chironomus-Arten (In Russian). 

In: Shobanov, N.A., Zinchenko, T.D. 

(eds.) Ecology, evolution and systematics of 

chironomids. Togliatti and Borok Russ. Acad. 

Sci., pp. 24-27. 

72 Wülker, W. 1997. Chironomus esai n.sp (Diptera:Chironomidae) 

in lakes and reservoirs of 

central Fennoscandia. - Ent. Fenn. 8: 171-176. 

73 Wülker, W., Kerkis, I.E., Kiknadze, I.I., Nevers, 

P. 1998. Chromosomes, morphology, ecology 

and distribution of Sergentia baueri n.sp., 

8 

S. prima Proviz and Proviz 1996 and S. corac 

ina Zett. 1824. - Spixiana 22: 69-81. 

74 Wülker, W., 1998. A Lobochironomus-species 

with 3 chromosomes (2n=6. - the true 

Chironomus (Lobochironomus) mendax Stora 

(Diptera, Chironomidae). - J. Kansas Entomol. 

Soc. 71: 304-314. 

75 Wülker, W. 1999. Fennoscandian Chironomus 

species (Dipt., Chironomidae) - identified by 

karyotypes and compared with the Russian and 

Central European fauna. - Stud. dipt. 6: 425- 

436. 

76 Butler, M.G., Kiknadze, I.I., Golygina, V.V., 

Martin, J., Istomina, A.G., Wülker, W., Sublette, 

J.E, Sublette, M.F. 1999. Cytogenetic 

differentiation between Palearctic and Nearctic 

populations of Chironomus plumosus L. (Diptera, 

Chironomidae). - Genome 42: 797-815. 

77 Wülker, W., Martin, J. 2000. Northernmost 

Chironomus karyotypes (Insecta, Diptera, Chironomidae). 

In: Baehr, M., Spies, M. (eds). 

Contributions to chironomid research in mem 

ory of Dr. Friedrich Reiss (Insecta, Diptera). 

- Spixiana 23: 151-156. 

78 Butler, M.G., Kiknadze, I.I., Golygina, V.V., Istomina, 

A.G., Wülker, W., Martin, J., Sublette, 

J.E, Sublette, M.F. 2000. Macrogeographic patterns 

of banding sequences in Holarctic Chi 

ronomus plumosus L. In: Hoffrichter, O. (ed.) 

Late 20th Century Research on Chironomidae: 

An Anthology from the 13th International Sym 

posium on Chironomidae, Freiburg, 5-9 September, 

1997, Shaker Verlag GmbH, Aachen: 

51-57. 

79 Kiknadze, I.I., Butler, M.G., Golygina, V.V., 

Martin, J., Wülker, W., Sublette J.E., Sublette, 

M.F. 2000. Intercontinental karyotypic differentiation 

of Chironomus entis Shobanov, a 

Holarctic member of the C. plumosus - group 

(Diptera, Chironomidae). - Genome 43: 857- 

873. 

80 Kiknadze, I.I., Butler, M.G., Golygina, V.V., 

Wülker, W., Martin, J., Sublette J.E., Sublette, 

M.F. 2000. Macrogeographic patterns of banding 

sequences in Holarctic Chironomus entis 

Shobanov. In: Hoffrichter, O. (ed.) Late 20th 

Century Research on Chironomidae: An An 

thology from the 13th International Symposi 

um on Chironomidae, Freiburg, 5-9 September, 

1997, Shaker Verlag GmbH, Aachen: 135-141. 

81 Istomina, A.G., Siirin, M.T., Makarevich, 

I.F., Scherbik, S.V., Wülker, W., Polukonova,

N.V. 2000. The fixation of polytene chromosome 

species-specific band sequences is the 

main differentiating factor in the obtusidens 

group of the genus Chironomus (Diptera, Chironomidae). 

In: Biodiversity and dynamics of 

ecosystems in North Eurasia. V. 1. Part 3. Mo 

leculargenetic bases of biodiversity (animals 

and plants), Inst. Tsitol. Genet., Sib. Otd. Russ. 

Akad. Nauk, Novosibirsk: 57-59. 

82 Spies, M., Sublette, J.E., Sublette, M.F., Wülker, 

W., Martin. J., Hille, A., Mitchell, M. A., 

Witt, K. 2002. Pan-American Chironomus 

calligraphus Goeldi, 1905 (Diptera, Chironomidae): 

species or complex? Evidence from 

external morphology, karyology and DNA sequencing. 

- Aquat. Insects 24: 91-113. 

83 Shobvanov, N., Wülker, W., Kiknadze, I.I. 

2002. Chironomus albimaculatus sp.n. and C. 

trabicola sp.n. (Diptera, Chironomidae) from 

polar Russia. - Aquat. Insects 24: 169-188. 

84 Kiknadze, I.I., Wülker, W., Istomina, A.G., Andreeva, 

E.N. 2005. Banding sequences pool of 

Chironomus anthracinus Zett. (Diptera, Chironomidae) 

in Palearctic and Nearctic. - Eu 

roasian Ent. J. 4: 13-27. 

85 Martin, J., Andreeva, E.N., Kiknadze, I.I., 

Wülker, W. 2006. Polytene chromosomes and 

phylogenetic relationships of Chironomus 

atrella (Diptera: Chironomidae) in North 

America. - Genome 49: 1384-1392. 

86 Wülker, W., 2007. Two new Chironomus species 

with fluviatilis-type larvae from the near- 

9 

shore sandy sediments of Lake Michigan (Diptera: 

Chironomidae). In: Andersen, T. (ed.) 

Contributions to the Systematics and Ecology 

of Aquatic Diptera A Tribute to Ole A. Sæther, 

The Caddis Press, Columbus, Ohio: 321-333. 

87 Kiknadze, I.I., Gunderina, L., Butler, M.G., 

Wülker, W., Martin, J. 2008. Chromosomes 

and continents. In: Dobretsov, N., Kolchanov, 

N., Rozanov, A., Zavarzin, G. (eds.) Biosphere 

origins and evolution, Springer, New York, 

N.Y.: 349-369. 

88 Wülker, W., Martin, J., Kiknadze, I.I., Sublette, 

J.E., Michiels, S. 2009. Chironomus blaylocki 

n.sp. and C. bifurcatus n.sp., North American 

species near the base of the decorus-group. 

- Zootaxa 2023: 28-46. 

89 Wülker, W. 2010. The role of chromosomes in 

chironomid systematics, ecology and phylogeny. 

In: Ferrington, L.C.,Jr. (ed.) Proceedings 

of the XV International Symposium on Chi 

ronomidae. Chironomidae Research Group, 

University of Minnesota, Saint Paul, Minnesota: 

1-13. 

90 Kiknadze, I.I., Istomina, A.G., Wülker, W., 

Vallenduuk, H.J. 2010. The karyotype of Chi 

ronomus uliginosus Keyl (Diptera, Chironomidae). 

- VOGiS Herald 14: 22-30 

Jon Martin 

Melbourne 

Odwin Hoffrichter 

Freiburg

Professor iya i. kiknadze: 80 th Birthday and 55 years 

Professor Iya Ivanovna Kiknadze is a leading expert 

in cell biology, cytogenetics, karyosystematics, 

and comparative and evolutionary genomics of 

Chironomidae. 

On February 9, 2010, this honoured worker of science 

of the Russian Federation celebrated her 80th anniversary. She is among the oldest researchers at 

the Institute of Cytology and Genetics (ICG), Novosibirsk, 

and Professors of the Novosibirsk State 

University (NSU). In addition, in 2010 it will be 

55 years since the beginning of her research work, 

of them, three years at the Laboratory of Cytology, 

Zoological Institute, Leningrad, later transformed 

into the Institute of Cytology. She has been working 

at ICG, Novosibirsk, for 52 years, since January, 

1958. For 32 years, from1962 till 1994, she 

headed a laboratory at the institute and from 1963 

to 1986, at NSU. 

Iya Kiknadze is the founder in Russia of the new 

research field: functional organization of chromosomes 

and differential gene activity in Diptera. 

She developed the notion of puffing as the basis 

for differential gene transcription and described 

the pattern and dynamics of puffs at major stages 

of chironomid development. With her supervision, 

the role of tissue-specific puffs was analyzed and 

an insight into the genetic control of tissue-specific 

secretory proteins was obtained. She pioneered in 

microdissection of the disk containing genes of 

the tissue-specific Balbiani ring, investigation of 

the molecular study of these genes, description of 

transposable elements in Chironomidae, and study 

of the cytogenetic control of the fine structure of 

chironomid salivary glands in the course of induced 

gene repression and expression. This study 

provided grounds for the original hypothesis of 

periodic genome reprogramming. Construction of 

high-resolution cytophotographic maps of chironomid 

polytene chromosomes allowed these species 

to be enrolled on the list of models for studies of 

evolutionary transformations in genomes and genetic 

impact of industrial factors on organisms. 

In particular, these results were extensively used 

in the assessment of radioactive pollution in the 

integrated projects: Remote Consequences of the 

Radioactive Impact of Nuclear Tests at the Semi 

palatinsk Test Site on the Population of the Altai 

Territory and Study of the Genetic Consequences 

of Nuclear Tests at the Semipalatinsk Test Site on 

Plant and Animal Populations. Iya Kiknadze supervised 

the development of inversion genomics of 

chironomids, based on global analysis of the disk 

in science and teaching 

10 

(gene) sequence polymorphism on various continents: 

Eurasia, North and South America, Africa, 

and Australia. Putative ancestral disk sequences 

were recognized in each of the chromosome arms 

of the genome (primitive karyotype). Phylogenetic 

trees were constructed for the first time for the 

genus Chironomus on the basis of inversion polymorphism 

in cooperation with researchers of the 

Institute of Mathematics, Novosibirsk, and the cytogenetic 

history of the genus was reconstructed. 

Iya I. Kiknadze at the Symposium of Cytogenetics of 

Invertebrates, August 2010. Photo W. Wülker. 

Iya Kiknadze was the first to obtain experimental 

evidence against the involvement of endomitosis 

in somatic polyploidization. It allowed revision 

of the endomitosis concept. Functional organization 

of chromosomes and differential gene activity 

are also among the subjects of Kiknadze’s seminal 

studies. Her monograph “Functional organization 

of polytene chromosomes” summarizes studies 

in this field. Iya Kiknadze applied the polytene 

chromosome model to the development of the 

essentials of the functional organization of interphase 

chromosomes and introduced the notion of 

chromomeres as functional units of these chromosomes. 

When working at ICG, Iya Kiknadze commenced 

studies of interphase chromosomes, chromomeres, 

and nucleoli. In collaboration with Dr. E. S. Belyaeva, 

she proved that the nucleolus was a transcriptionally 

active region of an interphase chromosome.

One more field of research conducted under 

Kiknadze’s supervision since 1980s is the molecular 

and cytological organization of specific regions 

in eukaryotic chromosomes, including the organization 

of multigenic loci and their transformation in 

the course of evolution. 

Long-term monitoring of chromosome pools of 

natural chironomid populations is conducted in 

permafrost regions of Yakutia in cooperation with 

the Institute of Ecology, Yakutian Academy of Sciences. 

Iya Ivanovna Kiknadze was born in the old merchant 

city of Tyumen on February 9, 1930. Her 

mother Antonina Reshetnikova and father Ivan 

Balakin were clerks. 

Iya spent all her childhood and school years in Tyumen, 

never going further than ten miles from it. She 

entered first School No. 1 and graduated from girl’s 

school No. 25. In wartime, Tyumen gave home not 

to only evacuated industrial enterprises but also 

many higher educational institutions from Moscow 

and other Russian cities. They included the 

staff of the Moscow Medical Institute. As it later 

turned out, the embalmed body of Vladimir Lenin 

was kept there during the Second World War. The 

cultural standard of the provincial town was notably 

improved by performances of companies of the 

Moscow Academic Art Theatre and other theatres. 

The metropolitan culture influenced the artistic 

taste of Tyumen inhabitants, in particular, teenagers. 

In 1947, Iya graduated school with a School 

Gold Medal and went to the “northern capital” to 

enter the Leningrad State University. 

In 1952, Iya Kiknadze graduated from the Faculty 

of Biology and Soil Science cum laude and became 

specialist in a field rare at that time: Darwinist–Geneticist. 

She started her research activity when 

learning at the University. From 1952 to 1955, she 

took a postgraduate course at the Leningrad State 

University and defended a candidate’s dissertation 

entitled Dynamics of DNA and RNA in oogenesis 

and early cleavage in invertebrates. Then she obtained 

her first position of Junior Researcher at the 

Laboratory of Cytology, Zoological Institute (In 

1956, the laboratory was transformed into the Institute 

of Cytology). Her research advisor at the University 

and Zoological Institute was Ivan I. Sokolov, 

a prominent cytologist. For years, he remained her 

tutor and standard of dedication and ethics. In 1957 

Iya was advised by A. A. Prokof’eva–Bel’govskaya 

to move to the just established Akademgorodok 

in Novosibirsk and obtain a job at ICG. She was 

promised she would have an apartment and interesting 

work. Iya was a daughter of Siberia, and she 

11 

did not resist it. In 1957 Iya Kiknadze, her husband 

the botanist Georgii Sergeevich Kiknadze, and their 

small daughter Irene left for Novosibirsk. 

At ICG, Iya Kiknadze became Junior Researcher 

at the Department of Physical, Chemical, and Cytological 

Basics of Heredity. It was headed by Prof. 

Ivan Dmitrievich Romanov till 1961. It was a remarkable 

time with a remarkable scientist. Even 

now, his portrait is on the wall in Iya Kiknadze’s 

study. During her first decade in Novosibirsk, 

Kiknadze made friends with older colleagues, Vera 

V. Khvostova and Raisa P. Martynova, and colleagues 

of her age, Ninel B. Khristolyubova and 

Klavdia K. Sidorova. 

In 1961–1962, Kiknadze worked as a Senior Researcher. 

Since October, 1962, she has headed the 

Laboratory of General Cytology. She defended 

her doctoral dissertation Functional Organization 

of Chromosomes in 1967, at a session of the Joint 

Dissertation Council in Biological Sciences, Siberian 

Branch of the USSR Academy of Sciences. 

She was awarded Professor’s rank at the Chair of 

Cytology and Genetics on December 10, 1970. 

On January 21, 1988, the Department of Cell Biology 

was founded on the base of the Laboratory 

of General Cytology. It included the Laboratory 

of Evolutionary Cytogenetics and several sectors: 

Genetics of Tissue-Specific Traits, Molecular Neurogenetics, 

and Genomics. 

Since 1994, Prof. Kiknadze has held the position 

of Chief Researcher at the Laboratory of Evolutionary 

Biology, ICG. 

Professor Iya Kiknadze convened several All- 

Union and international conferences: the 2nd All- 

Union Symposium Chromosome Structure and 

Function (Novosibirsk, 1970) and the All-Union 

Symposium Chironomidae Evolution, Speciation, 

and Systematics (Novosibirsk, 1985). In 1982, 

the international symposium Organization and 

Expression of Tissue-Specific Genes was held in 

Akademgorodok in 1982. It marked the beginning 

of regular workshops on chironomid Balbiani 

rings. At present, I. Kiknadze is participating 

in the preparation of the conference Invertebrate 

Karyosystematics (Novosibirsk) as Chairlady of 

the Organizing Committee. 

Professor Iya Kiknadze is the founder of the 

Novosibirsk school of dipterologists, experts in 

Chironomidae and Drosophilidae. Twenty-nine 

candidate’s dissertations were defended with her 

scientific supervision. She generously presents her 

ideas to her students. She has dozens of followers 

and disciples who have chosen their own research

ways in many countries. Many of Kiknadze’s students 

have raised their own students. 

Professor Iya Kiknadze applied much effort to 

teaching of cytology at the Chair of Cytology and 

Genetics, Biological Department, Faculty of Sciences, 

NSU. From the foundation of the Chair of 

Cytology and Genetics in 1962 to 1986, she performed 

the offices of Vice-Chairholder. She was 

the first not only at NSU but in the USSR to develop 

and deliver the course Cytology/Cell Biol 

ogy after decades of the stranglehold of Michurin’s 

biology. She also held and supervised seminars 

and laboratory courses on branches of cytology 

and genetics. 

Iya Kiknadze introduced her experience acquired 

from the Leningrad biological research school 

to ICG and NSU. She started with the system of 

biologist tutorship. Major and minor laboratory 

courses on cytology and genetics and summer 

practicals in the field were conducted in the image 

and likeness of corresponding activities at the 

Chair of Biology of the Leningrad University. This 

was how the famous Leningrad school of cytology 

and genetics, created by the famous scientists 

Yu. A. Filipchenko, M. E. Lobashov, D. N. Nasonov, 

and M. S. Navashin, sprouted in Siberia. 

Since 1965, Iya Kiknadze has been a member of 

the Joint Academic Board in Biology, Siberian 

Branch of the USSR Academy of Sciences. She 

was a permanent member of the Academic Board 

of ICG since its establishment, a member of the 

Academic Board of the Faculty of Sciences, NSU, 

and a member of the Dissertation Council at the 

Novosibirsk State Medical University. Now Prof. 

Kiknadze is also a member of the Dissertation 

Council at the Institute of Systematics and Ecology 

of Animals. 

For 28 years, from 1974 to 2001, Iya Kiknadze 

was a member of the Editorial Board of the journal 

Tsitologiya (Cytology), and for 5 years, from 

1984 to 1989, of the Editorial Board of Ontogenez 

(Development). Since 2003, she has been a member 

of the Editorial Board of Evroaziatskii Ento 

mologicheskii Zhurnal (Eurasian Entomological 

Journal), and since 2008, of the Editorial Board of 

Comparative Cytogenetics. 

The Honoured Worker of Science of the Russian 

Federation (since 1998), Professor Kiknadze was 

awarded the Order of the Badge of Honour in 1967 

for participation in the development of the Novosibirsk 

Research Centre and scientific contribution; 

in 1970, the Medal for Valorous Labour. She has the 

titles of Honoured Veteran of the Siberian Branch 

of the USSR Academy of Sciences and Honoured 

12 

Veteran of Labour. Certificates of Merit: from SB 

RAS on the occasion of the 275 th Anniversary of 

the Academy of Sciences in 1999, from the Ministry 

of Education and Science on the occasion of 

the 50 th Anniversary of the Siberian Branch of the 

RAS in 2007, and from the Novosibirsk Governor 

V. A. Tolokonsky in 2010. 

Iya Kiknadze is an earnest and fruitful worker. It is 

apparent from her numerous publications in the recent 

decade and enormous work on reviewing and 

editing of research papers. Her desk is always covered 

by fans of photographs of her favourite polytene 

chromosomes. Their banding patterns serve as 

barcodes. They allow the features and evolutional 

history of each species to be understood. With the 

help of her students, A. G. Istomina, L. I. Gunderina, 

V. V. Golygina, and A. D. Broshkov, as well 

as with numerous Russian and foreign colleagues, 

Iya Kiknadze works in the enormous field of chromosome 

pools, chromosome polymorphism, and 

speciation in Holarctic Chironomidae species. 

These studies contribute to cytogenetics of natural 

chironomid populations in the context of the 

role of chromosome rearrangements in population 

adaptation and chromosome set divergence during 

speciation. They have revealed profound cytogenetic 

divergence between Palearctic and Nearctic 

populations of Holarctic species. 

Iya Kiknadze is among authors of over 350 publications, 

including 10 monographs. 

During all scientific activity Prof. I. Kiknadze had 

fruitful contacts with many specialists from different 

countries. She took part in many international 

symposia and other scientific meetings. While 

studying the structure and function of chironomid 

polytene chromosomes she had associated 

with Prof. B. Daneholt, Prof. E.R. Schmidt, Prof. 

M. Lezzi, Prof. J.E. Edström, Prof. U. Grossbach, 

Prof. R. Panitz, Prof. S. Case, H. Bäumlein, U. 

Wobus. Joint work with Prof. W. Wülker, Prof. J. 

Martin, Prof. M. Butler, Prof. X. Wang, Prof. R. 

Contreras-Lichtenberg, H. Vallenduuk, H. Moller- 

Pilot result in the description of new species, their 

karyotypes and chromosomal polymorphism. Iya 

Kiknadze collected chironomid larvae with M. 

Butler in North America. The fruitful interactions 

with Prof. W. Wülker and Prof. J. Martin resulted 

in revision and making cytomaps of polytene 

chromosomes of many Chironomus species more 

precise. In joint work with Prof. W. Wülker, Prof. 

J. Martin, Prof M. Butler and Dr. L. Gunderina it 

has been shown that the banding sequence pools of 

populations of the same species on different continents 

differed much in the sets and frequencies of

gene inversion orders. Banding sequence pools of 

populations on each continent were found to contain 

continent-specific banding sequences in addition 

to sequences occurring on several continents. 

We wish Iya Kiknadze many more interesting 

years full with new ideas and successes. 

I.K. Zakharov, A.G. Istomina, W. Wülker 

Iya Kiknadze’s selected publications 

Kiknadze I.I. 1972. The functional organization of 

the chromosomes. - L. Nauka, 212p. (In Russian). 

Kiknadze I.I., Shilova A.I., Kerkis I.E. et al. 1991. 

Karyotypes and larval morphology in tribe 

Chironomini. An Atlas. Novosibirsk: Nauka. 

113 p. (In Russian with English Summary). 

Kiknadze, I.I., Istomina, A.G., Gunderina, L.I., 

Salova, T.A., Aimanova, K.G. & Savvinov 

D.D. 1996. The banding sequences pool of 

chironomids from permafrost zone of Yakutia: 

tribe Chironomini. Nauka Publishing, Novosibirsk, 

166 pp. (In Russian). 

Kiknadze I.I., Kokoza V.A., Kolesnikiv N.N., Karakin 

E.I., Scherbakov D.Yu., Aimanova K.G., 

Agapova O.A., Zainiev G.A., Kopantsev E.P., 

Sebeleva T.E. 1985. Organization and expres 

sion of tissue specific genes in Diptera. Novosibirsk: 

Nauka Publ. 237 p. (In Russian). 

Kiknadze I.I., Blinov A.G., Kolesnikov N.N. 1989. 

The molecular-cytological organization of chironomid 

genome; (pp. 4-59) in Shumny V.K. 

(Ed.) Structural and functional organization of 

genome. Novosibirsk, 189 p. (In Russian with 

English summary). 

Kiknadze I.I. 1967. The chromosomes of Diptera. 

The evolutionary and practical significance of 

karyotype study. - Genetika 3: 145-165. (In 

Russian with English Summary). 

Kiknadze I.I. 1970. The transcriptional activity of 

chromosomes during differentiation. - Soviet 

Journal of Developmental Biology, 1: 7-27. (In 


Kiknadze .I.I. 1971. Polytene chromosomes as a 

model of interphase chromosome. - Tsitologia 

13: 716-732. (In Russian with English Summary). 

Kiknadze I.I., Perov N.A., Chentsov Yu.S. 1976. 

Ultrastructural organization of salivary gland 

polytene chromosomes of Chironomus thum 

mi. - Chromosoma 55: 91-102. 

13 

Kiknadze I.I. 1976. Comparative characteristics of 

puffing pattern in salivary gland Chironomus 

during larval development and metamorphosis. 

I. Puffing pattern in chromosome IV. - Tsito 

logia 18: 1322-1329. (In Russian with English 

Summary). 

Kiknadze I.I. 1978. Comparative characteristics of 

puffing pattern in salivary gland Chironomus 

during larval development and metamorphosis. 

II. Puffing pattern in chromosomes I, II, III. - 

Tsitologia 20: 514-521. (In Russian with English 

Summary). 

Kiknadze I.I., Valeeva F.S., Vlasova I.E., Panova 

T.M., Sebeleva T.E., Kolesnikov N.N. 1979. 

Puffing and specific function of salivary gland 

in Chironomus thummi. I. Quantitative changes 

of proteins and glycoproteins in the salivary 

gland at different larval stages. - Soviet Jour 

nal of Developmental Biology 10: 161-172. (In 


Kiknadze I.I. Chromosomal polymorphism in natural 

populations of the plumosus species-group 

of West Siberia (Diptera: Chironomidae). - En 

tomol. Scand. 1987, Suppl. 29: 113-121. 

Kiknadze I.I., Kolesnikov N.N., Panova T.M., Gydamakova 

E.K., Blinov A.G., Filippova M.A. 

1987. Transposable elements in Chironomidae 

I. Location of pCth C1.2HR sequence in the 

polytene chromosomes of closely related subspecies 

Chironomus thummi thummi Kieffer, 

C. th. piger Strenzke and their hybrids. - Gene 

tika 23: 1366-1376. (In Russian with English 

Summary). 

Kiknadze I.I., Butler M.G., Aimanova K.G. et al. 

1998. Divergent cytogenetic evolution in Nearctic 

and Palearctic populations of sibling species 

in subgenus Camptochironomus Kieffer. 

- Can. J. Zool. 76; 361–376. 

Butler M.G., Kiknadze I.I., Golygina V.V. et al. 

1999. Cytogenetic differentiation between 

Palearctic and Nearctic populations of Chi 

ronomus plumosus L. (Diptera, Chironomidae) 

- Genome 42: 797–815. 

Kiknadze I.I., Butler M.G., Golygina V.V. et al. 

2000. Inter-continental cytogenetic differentiation 

in Chironomus entis Shobanov, a Holarctic 

species in the plumosus-group (Diptera, Chironomidae) 

- Genome 43: 857–873. 

Kiknadze I.I., Gunderina L.I., Istomina A.G. et al. 

2003. Similarity analysis of inversion banding 

sequences of Chironomus species (breakpoint 

phylogeny) In: Kolchanov N., Hofestaedt R..

(eds) Bioinformatics of genome regulation and 

structure Boston/Dordrecht/London: Kluwer 

Acad. Publ.. pp. 245–253. 

Kiknadze I.I. 2008. The role of chromosome polymorphism 

in divergence of populations and 

species of the genus Chironomus. - Entomol. 

Rev. 88: 509–524. 

Kiknadze I.I., Golygina V.V., Broshkov F.D. et al. 

2008. Mystery of Chironomus dorsalis karyo- 

arthur desmond harrison (24.12.1921 - 30.12.2007) – an aPPreciation 

Arthur D. Harrison, the doyen of African limnology 

and studies of the Afrotropical Chironomidae, 

died in Canada over 3 years ago with little or no 

posthumous scientific recognition. This piece is an 

attempt to rectify the situation, and to recognise 

the significance of Arthur as a clear-thinking and 

often pioneering biogeographer, entomologist, educator 

and above all as a limnologist in the broadest 

sense. His life and career spanned Africa and 

Canada, although his influence remains very much 

associated with Africa, from Ethiopia to the Cape. 

Arthur D. Harrison. Photo Helen James 

Arthur was born in the Western Cape of South Africa, 

at Kalk Bay, on 24 th December 1921, and he 

lived for a period at Fish Hoek (where he returned 

in retirement – see photograph). He attended school 

in Rondebosch and then studied at the University 

of Cape Town where he gained B.Sc and M.Sc. He 

then obtained teaching qualifications (B.Ed) and 

took up a teaching appointment before deciding 

that research was to be his career. He gained a po- 

14 

type (Diptera, Chironomidae) - Comp. Cy 

togenet. 2: 21–35. 

Kiknadze I.I., Gunderina L.I., Butler M.G. et al. 

2008. Chromosomes and continents. In: Dobretsov 

N., Rozanov A., Kolchanov N., Zavarzin 

G., (eds.) Biosphere Origin and Evolution. 

Springer-Verlag pp. 349-369. 

sition at CSIR (South Africa’s Council for Scientific 

and Industrial Research) undertaking limnological 

surveys in rivers and estuaries. This led to 

the award of a Ph.D. for his pioneering studies of 

the Great Berg River (of the Western Cape), published 

in two parts in the Transactions of the Royal 

Society of South Africa (Harrison 1958; Harrison 

& Elsworth 1958) and revisited subsequently (e.g. 

Harrison 1964). These papers (‘beacons’ in the 

literature according Allanson 2003) remain wellcited 

and show Arthur’s early recognition of the 

downstream effects (zonation) in river structure 

and function from headwaters to estuaries. From 

the outset Arthur balanced his limnological studies 

with the applied – one of his earliest papers concerned 

the effects of acidic mine pollution on the 

streams of the Transvaal. Throughout his career, 

by himself or others with his assistance, revealed a 

range of human impacts on aquatic ecosystems. 

Fish Hoek. Photo P. Cranston 

After relocations with CSIR (Witwatersrand University, 

then Pretoria) Arthur took up a Rockefeller-funded 

position studying Bilharzia at the 

University of Salisbury, in what was then (early- 

1960s) Rhodesia. Arthur clearly had some time on 

his hands aside from studying the effects of molluscicides. 

He observed the recovery of a Rhodesian 

stream, post-drought, including documenting

the chironomids, and finding the adults of the first 

known podonomine midge from Africa, described 

as Afrochlus harrisoni by Freeman (1964). Following 

the Unilateral Declaration of Independence 

(from the UK) in November 1965, Rockefeller 

funding was withdrawn under sanctions, 

and Arthur returned to South Africa to become 

Professor of Zoology at the University of Natal, in 

Pietermaritzburg. 

This was a short-lived appointment as Arthur accepted 

an invitation to join the faculty of the University 

of Waterloo in Ontario, Canada, where 

under Noel Hynes he joined a dynamic group of 

tropical limnologists. At this time Arthur’s deepening 

personal interest in the chironomids became 

more evident. Although he had collected the 

midges throughout his career, including finding in 

1954 Harrisonina petricola Freeman in an ephemeral 

stream in Oliphants Valley and recognising it 

as curious, he passed many of his specimens onto 

others, first to Marjorie Scott, and then to Paul 

Freeman at the Natural History Museum, London 

(BMNH). However his own first publications on 

the group started to appear in the early 1970s, with 

his interest in the Tanypodinae evident when he 

took Sepp Fittkau’s (1962) revision of the Tanypodinae 

and placed the somewhat neglected African 

species into the modern generic concepts (Harrison 

1971). In the course of this study, he described 

his first midge genus (Lepidopelopia) for the ‘one 

that didn’t fit’ the Fittkau scheme (Harrison 1970). 

It was at this time that I first met Arthur as he came 

through London to view the types of the African 

Tanypodinae held in the BMNH – he asked me to 

make microscope slide preparations in advance of 

his visit so that he could see features such as the 

spurs on the adult legs that characterised Fittkau’s 

new taxa. Arthur was a stickler for correct preparations, 

and it gave me some pleasure to ‘pass the 

test’. He was very alert to the ‘modern’ means of 

doing chironomid taxonomy, and made slides of 

most of his specimens as he collected them. Further, 

he reared much and tried to incorporate the 

immature stages. 

For many subsequent years our paths did not cross, 

as Arthur spent his time either in Waterloo, or seconded 

to the University of Addis Ababa in Ethiopia. 

From 1981 to 1989 he was a major contributor 

to the Canadian International Development Agency 

(CIDA)-funded Institutional Enhancement 

project, spending 4 years living there. Another of 

the Waterloo faculty engaged in this project, Herbert 

Fernando, recalled ‘At the best of times Ethiopia 

is not an easy country to work in. But these 

were not good times. We needed permission from 

15 

the highest government authorities to leave Addis 

Ababa even for field work.’ Despite this situation, 

experienced Africa-hand Arthur succeeded in doing 

much publishable research, and he produced 

a series of papers on the Chironomidae of Ethiopian 

lakes, extending distributions of known taxa, 

and describing new species and new life histories. 

He managed to get himself to the high elevation 

streams as well as the Rift Valley lakes, and published 

general invertebrate reviews with Noel 

Hynes and some particular chironomid papers. 

Arthur published this work especially in Archiv für 

Hydrobiology and Spixiana, and always he tried to 

get his research out to the appropriate audience. 

When Aquatic Insects started, he was a contributor 

and supporter from the outset (e.g. Harrison 2000). 

Further, he seemed never to decline an invitation 

to summarise his compendious and very broad 

understanding of the ecology and distribution of 

African invertebrates (e.g. Harrison 1978, 1995). 

He wrote fluently and with a highly readable style 

– and was a frequent correspondent and an early 

adopter of e-mail. 

On his retirement from Waterloo, he lived on Vancouver 

Island for some years, but he felt the health 

of his wife, Jessie, might benefit from returning to 

South Africa where more help was available. Although 

she died in 1994, Arthur remained in Fish 

Hoek, making periodic visits to his family in Canada 

and to Perth, Australia. It was in the Western 

Cape, post-apartheid, that I resumed ‘in person’ 

acquaintance with Arthur as I started to visit the 

‘rainbow nation’ and adjacent countries. Arthur 

guided me by hand-drawn maps and photographs 

and detailed verbal instructions to localities including 

for Afrochlus in Zimbabwe (see photograph 

of Ngoma Kurira) and Aphrotenia in the Western 

Cape. He had immense knowledge (and profound 

memory) of aquatic locations and their inhabitants 

throughout southern Africa. In the 1990s and 

early years of this century, this knowledge continued 

to be extended as he identified chironomids, 

seemingly for all aquatic research groups in South 

Africa. Fortunately he continued to publish from 

his immense collections and those provided by 

his collaborators, although his field work was curtailed. 

When Don Edward (University of Western 

Australia) and I visited the Cape in 1998, Arthur 

accompanied us to the upper Eerste River in Jonkershoek. 

We collaborated subsequently, including 

over an orthoclad, Elpiscladius, a member of the 

Brillia group for which Arthur had a pharate male 

(Harrison & Cranston 2007). Little did we know 

but the then-unknown larva was in the Eerste– mining 

in immersed wood as its phylogeny predicted 

(Cranston 2008).

Ngome Kurira. Photo: P. Cranston. 

My last meeting with Arthur in person was in Fish 

Hoek in early 2004 when he announced that he 

was to return to be nearer to the trusted medical 

facilities of Vancouver. Arthur continued to write 

wonderful e-mails, full of biogeographic and taxonomic 

insights from the vast range of organisms 

with which he was familiar. If a communication 

silence went on too long he would write simply 

to enquire ‘where in the world was I?’. The last 

silence though was on his part: the precursor to his 

death at the end of 2007, although sadly this news 

was slow to spread. 

Arthur Harrison was an immensely knowledgeable, 

insightful and very productive scientist to the end. 

Justifiably, his major works continue to be well 

cited – he was a limnological pioneer in Africa at 

the time the field was in its infancy. His breadth of 

knowledge of invertebrates and their distributions 

was unrivalled, and his biogeographic insights 

(e.g. Harrison 1978) have stood the test of time. 

Further, although he had a tremendous empathy 

with local peoples, he showed that quality biological 

research can be produced under most arduous 

circumstances. We will not see his like again. 

I am grateful to Ferdy de Moor for provoking this 

article – I hope that my heartfelt appreciation is 

better late than not at all. 

Peter S. Cranston 

University of California, Davis 

References 

Allanson, B.R. 2003. Beacons in the limnological 

landscape. - African Journal of Aquatic Sci 

ence 28: 89-101 

Cranston, P.S. 2008. Phylogeny predicts larval biology 

in a wood-mining chironomid (Diptera: 

Chironomidae). - African Entomology 16: 1-6. 

Fittkau, E.J. 1962 Die Tanypodinae (Diptera: Chironomidae). 

(Die Tribus Anatopyniini, Macropelopiini 

und Pentaneurini). - Abhandlungen 

zur Larvalsystematik der Insekten 6: 1–453. 

16 

Freeman, P. 1964. Notes on Chironomidae (Dip(Diptera, Chironomidae). - Proceedings of the Roy 

al Entomological Society of London (B) 33: 

147-150. 

Harrison, A.D. 1958. Hydrobiological studies on 

the Great Berg River, Western Cape Province. 

Part 2. Quantitative studies on sandy bottoms, 

notes on the tributaries and further information 

on the fauna, arranged systematically. - Trans 

actions of the Royal Society of South Africa 35: 

227-276. 

Harrison, A.D. 1964. An ecological study of the 

Berg River, pp.143-158, in Ecological Stud 

ies in Southern Africa, ed. D.H.S. Davis, The 

Hague: W. Junk, 415pp. (Monographiae Biologicae 

14). 

Harrison, A.D. 1970. Lepidopelopia, a new chironomid 

genus with scale-like setae on the 

legs (Dipt., Chironomidae). - Journal of the 

Entomological Society of South Africa 33: 295- 

301. 

Harrison, A.D. 1971. A conspectus of the Macropelopiini 

and Pentaneurini (Tanypodinae: 

Chironomidae) of Africa south of the Sahara. 

- Canadian Entomologist 103: 386-390. 

Harrison, A.D. 1978. Freshwater invertebrates (except 

Molluscs), pp. 1139-1152, in Biogeogra 

phy and Ecology of Southern Africa, ed. M.J.A. 

Werger, with the assistance of A. C. Van Bruggen. 

2 parts. The Hague: Dr W. Junk. 1-660 

p. (pt. 1); 661-1439 p. (pt. 2). (Monographiae 

Biologicae, 31). 

Harrison, A.D. 1995. Northeast Africa Rivers and 

Streams, pp. 507-517, in Ecosystems of the 

world: 22. River and stream ecosystems, eds. 

C. E. Cushing, K. W. Cummins, and G. W. 

Minshall, Amsterdam, The Netherlands, Elsevier, 

817 pp. 

Harrison, A.D. 2000. Four new genera and species 

of Chironomidae (Diptera) from southern Africa. 

- Aquatic Insects 22: 219-236. 

Harrison, A.D. & Cranston, P.S. 2007. Elpiscladius 

Harrison and Cranston, a new orthoclad (Diptera: 

Chironomidae) in the Brillia group from 

South Africa. - Annals of the Eastern Cape Mu 

seum 6: 1-11. 

Harrison, A.D. & Elsworth, J.F. 1958. Hydrobiological 

studies on the Great Berg River, Western 

Cape Province. Part I General Description, 

Chemical Studies and Main Features of the 

Flora and Fauna. - Transactions of the Royal 

Society of South Africa 35: 125-226.

Paul freeman – an aPPreciation of his studies on chironomidae 

Paul Freeman, August 1960. Photo G.W. Byers 

Dr. Paul Freeman died at the age of 94 at the end of 

July this year. To many contemporary chironomidologists 

he will be known principally for his contribution 

to untangling the taxonomy of the sub- 

Saharan African Chironomidae. The results were 

published in 4 parts in the Bulletin of the British 

Museum (Natural History), Entomology series, between 

1955 and 1958 (Freeman 1955, 1956, 1957, 

1958), followed by the Chironomidae of New Zealand 

(Freeman 1959) and of Australia (Freeman, 

1961). One third of Freeman’s scientific publications 

(of 86 in total) concerned the Chironomidae: 

the others ranged across several other families of 

nematocerous Diptera. After his first publication 

on chironomid midges (adding two new species to 

the United Kingdom list in 1948), the remainder 

concerned non-European species, especially, but 

not exclusively, from Africa, Australia and New 

Zealand. Freeman apparently never visited these 

countries but relied on an extensive network of colleagues 

who sent him adult midges. Amongst these 

scientists based in Africa were Arthur Harrison 

(whose commemoration can be found elsewhere 

in this issue) and Margaret (K.M.F.) Scott from 

the University of Cape Town, who are expressly 

thanked at the outset of the sub-Saharan studies for 

their big collection of adult midges ‘in excellent 

condition’. From the Sudan, David (D.J.) Lewis 

is acknowledged for much material including the 

asthma-inducing Nile midge that Freeman named 

for its collector (Cladotanytarsus lewisi (Freeman, 

1950)). From throughout colonial Africa people 

sent material to London, providing the impetus 

for a series of short papers on particular National 

Parks, especially those in central Africa. This material 

led Freeman to understand the problematic 

influence of the Abbe J.J. Kieffer – whose work 

17 

he described as ‘very erratic’, noting the ‘very uncertain’ 

concepts of genera, paucity of illustrations 

and redescription of the same species ‘over and 

over again not only in different papers but even in 

the same one’ (Freeman 1955: pp. 5–6). Although 

Freeman examined as many of Kieffer’s types as 

could be found (many are lost amongst the 300 

Kieffer described from the region), and he disentangled 

the taxonomic confusion as best he could, 

he concluded that more collecting was needed in 

type localities, picking out Kribi (tropical Cameroon) 

as especially important. Sadly this situation 

remains essentially as true today as it was during 

the 1950s. 

Freeman’s African studies were important in placing 

the midges of a large continent into more modern 

generic (including subgeneric) concepts, yet he 

published at a time of turn-over in our ideas and 

methods. Freeman used pinned adults, but prepared 

hypopygial mounts and drew quite detailed 

and accurate figures of these structures. The days of 

routine slide mounting of the complete adult were 

yet to come, but the warning that Freeman gave 

of the tendency of coverslips to ‘distort’ genitalia 

remains as pertinent today as then. The post-WWII 

years was the time when European entomologists 

were building on an increased understanding of the 

significance of the immature stages in classification 

– led by what has been termed the ‘Thienemann 

school’ of ecologists-turned-taxonomists. These 

disciples often reared their larvae to adulthood, 

retained the immature stages (the larvae and the 

quite critical pupal exuviae) and had sent the adults 

to Kieffer for description. The outcome became a 

more synthetic (and coherent) generic concept, often 

parallel to that derived from adults alone, but 

generally narrower. What is more, the Hennigian 

revolution was starting ‘on the continent’ with 

early adherents amongst some of the chironomid 

workers. One could cite Strenzke’s (1960) explicit 

application to the chironomids (for Clunio and 

relatives), and phylogenetic thinking was evident 

earlier amongst the Thienemann group. Actually 

such thinking pre-dated Willi Hennig, as F.W. Edwards, 

one of Freeman’s predecessors in studying 

Nematocera in the British Museum (Natural History), 

was remarkably prescient about these issues 

(Edwards, 1926). However Freeman’s African 

studies were at the cusp of this transformation, and 

his higher level taxonomic work remained closer 

to the traditional adult-based scheme of Goetghebuer. 

This is not to criticise these studies for not 

being ‘ahead of their time’ from an ecological or 

phylogenetic perspective – history certainly has

een kind to Freeman’s species concepts: his keys 

work, there is little or no synonymy, and the ‘tidying-up’ 

of so much of Kieffer’s African concepts 

was of immense value to later taxonomists. For 

a more modern allocation of the taxa to genera, 

Freeman (with some help from an acolyte) did this 

in the Catalogue of the Diptera of the Afrotropical 

Region (Freeman & Cranston 1980) – his last publication 

on the family. 

After the major African publication, Freeman continued 

to receive additional chironomids of interest, 

notably an Afromontane Diamesa from Mount 

Kenya, the southernmost representative of the genus 

(Freeman 1964a) and the first African species 

belonging to the ‘cool stenothermic’ subfamily 

Podonominae (Freeman 1964b). Paul recounted to 

me his astonishment when he referred Arthur Harrison’s 

specimens of this first African podonomine 

from Zimbabwe – the genus Afrochlus – to Lars 

Brundin who was revising the subfamily. Lars’ 

postcard thanking Paul for the material was mailed 

from South Africa where Lars was already seeking 

more material. I am not sure if Freeman’s astonishment 

concerned the ease with which the Head 

of Zoology at the Stockholm Museum could head 

south, or the cost, or both, but Lars was not only 

Head, but also in charge of the Museum’s travel 

budget. 

By this time Freeman had completed his immersion 

in Australasian Chironomidae – having produced 

first the study on New Zealand (Freeman 1959) 

and then his Chironomidae of Australia (Freeman 

1961). These works differ from the African 

studies in several ways: the nomenclatural issues 

were more tractable (less of Kieffer), incompleteness 

of the survey material available to him was 

acknowledged (no Arthur Harrison!), more genera 

were described as new in the works (3 from New 

Zealand, 12 for Australia), and there was a strong 

visibility of some modern biogeographic thinking. 

Although Freeman did not publish on South 

American Chironomidae, he understood Edwards 

(1931) studies, and thus was able to recognise 

Neotropical elements in both New Zealand and 

Australia (e.g. Stictocladius, Riethia). Further, he 

reallocated some African species of Chironomini 

to groupings that he recognised and described as 

new from Australia, notably Conochironomus and 

Skusella. In the short summary in the introduction 

to the Australian work (‘Distribution and affinities 

of the Australian Chironomidae’), there is scarcely 

an incorrect idea. Studies in both countries since 

Freeman’s publications have extend the biogeographic 

ideas, notably through Brundin (1966) 

who encountered a much more diverse Podonominae 

fauna than Freeman had available to him, and 

18 

to myself including with Don Edward (e.g. Cranston 

& Edward 1999), who delved into the ‘little 

black orthoclads’ of the austral continents. Neverthe-less, 

Freeman’s new Antipodean genera hold 

up, including against the molecular data becoming 

available. 

That Paul Freeman’s research on Chironomidae 

slowed down, albeit almost ceased in the late 1960s 

was due to his promotion to lead the Entomology 

Department of the Natural History Museum 

(termed ‘keeper’). His leadership skills were well 

demonstrated in 1964 when he organised the International 

Congress of Entomology, held in London. 

Further, he had a truly hands-on involvement in the 

sorely-missed ‘new’ Insect Gallery of the Museum 

that lured many a child, and perhaps adults too, 

with a celebration of insect diversity long before 

the term became popular. This was all before my 

time – when I interviewed for a lowly technical 

position in the Museum in the ‘summer of mass 

unemployment’ (1971) Freeman already had occupied 

the top floor Keeper’s Office for 3 years. 

However he was the Departmental representative 

on the recruitment panel established for some vacancies, 

including the one that I had applied for: 

assistant scientific officer in the Ornithology section. 

On being told that the vacancy in birds was 

filled already, the Keeper put the ‘soft sell’ on me to 

consider working with insects, leveraging the Museum’s 

generous policy on work release to pursue 

higher education and extolling the pathways that 

an enthusiastic junior member of the staff could 

pursue as a career. My negative experiences with 

entomology teaching during an incomplete undergraduate 

degree were no match for this persuasiveness, 

and so I declined other offers to control yeast 

quality in a brewery or culture cells in a cancer 

research hospital. Obituaries for Paul Freeman in 

the Guardian (http://www.guardian.co.uk/environment/2010/aug/25/paul-freeman-obituary) 

and the 

Telegraph (http://www.telegraph.co.uk/news/obituaries/science-obituaries/7960471/Paul-Freeman. 

html) both point to his support for his younger staff 

– he is quoted as stating “It is important to look after 

the junior staff, as the senior staff can look after 

themselves”. I can affirm that this was especially 

so in my case – shortly after entry I was given the 

position of technical support for the nematocerous 

Diptera families, already a budding career trajectory 

for two assistants that later became Keepers 

– Dick Vane-Wright and Richard Lane. After familiarisation 

with the Diptera families and some 

work with both Tipulidae and Mycetophilidae I 

was encouraged, surely with the guiding hand of 

Paul Freeman, to curate the collections of Tanypodinae 

(which existed as pinned adults) in the 

light of Sepp Fittkau’s Die Tanypodinae (Fittkau,

1962). There was a steep learning curve – it was 

in German (without Google translator to assist), 

dealt with features that could only be seen on good 

slide mounts with high power magnification, and 

described a plethora of genera compared to what 

was currently in use in English language guides. 

The visit to London of Arthur Harrison to review 

the African Tanypodinae against Fittkau’s work assured 

me not only that I was on the right track, but 

that I was not the only one interested in getting the 

subfamily into a modern framework. 

When it came time to undertake a Ph.D., it was natural 

to stay with the Chironomidae, and I chose to 

work with the immature Orthocladiinae with guidance 

from ecologist Alan Hildrew and from Paul 

Freeman. This was before the days when Museums 

and like institutions saw a role for themselves 

in higher education, and certainly I was early into 

the system of having formal approval for Museum 

research to be directed towards the goal of a higher 

degree. As he had promised at outset, Paul Freeman 

was very supportive throughout the study and 

although not very conversant with immature stages 

he knew the broad and specific literature extremely 

well. Perhaps what has stayed with me most was 

his questioning of ‘publishability’ of research, 

long prior to the ‘publish or perish’ days. Simply 

put, he felt that if the taxpayer has paid for the research 

then there was an obligation to complete 

the work by publishing it. With the Keeper coming 

from a background in Diptera, I often heard it said 

that the cluster of staff Dipterists were the recipients 

of some favouritism. Although those were the 

days when budgets seemed to increase each year, 

and the Diptera section surely was blessed with a 

stream of very able technicans and some more senior 

recruits, Paul argued his strong support for the 

group was based on their publishing productivity. 

Amongst these was the multi-collaborative project 

led by Roger Crosskey’s editing of the Catalogue 

of the Diptera of the Afrotropical Region and for 

which Paul and I co-authored the Chironomidae 

contribution. 

My career owes its entirety to that recruitment 

promise made by Paul Freeman, and delivered 

upon – support his junior staff he surely did, by 

deeds and example. 

Peter S. Cranston 

University of California, Davis 

References 

Brundin, L. 1966. Transantarctic relationships and 

their significance, as evidenced by chironomid 

midges with a monograph of the subfamilies 

19 

Podonominae and Aphroteniinae and the austral 

Heptagyiae. Kung. Sv. Vetenskapsakad. 

Handl. 11: 1–472. 

Cranston, P.S. & Edward, D.H.D. 1999. Botryo 

cladius gen. nov.: a new transantarctic genus of 

orthocladiine midge (Diptera: Chironomidae). 

Syst. Ent. 24: 305–33. 

Edwards, F.W. 1926. The phylogeny of nematocerous 

Diptera: a critical review of some recent 

suggestions. pp. 111–130, Internationaler Entomologen-Kongress, 

Zurich, July, 1925. 

Edwards, F.W. 1931. Part II. Fascicle 5. Chironomidae. 

Diptera of Patagonia and South Chile. 

pp. 232–331. 

Fittkau, E.J. 1962. Die Tanypodinae (Diptera: Chironomidae). 

(Die tribus Anatopynyiini, Macropelopiini 

und Pentaneurini). - Abhandl. Lar 

vensyst. Ins. 6: 1-453. 

Freeman, P. 1950. A species of chironomid from 

the Sudan suspected of causing asthma. Proc. 

R. Ent. Soc. Lond. (B) 19: 58–59. 

Freeman, P. 1955. A study of the Chironomidae 

(Diptera) of Africa south of the Sahara. Part I. 

Bull. Brit. Mus. (Nat. Hist.), Ent. 4: 1–67. 


(Diptera) of Africa south of the Sahara. Part II. 



(Diptera) of Africa south of the Sahara. Part III. 



(Diptera) of Africa south of the Sahara. Part IV. 


Freeman, P. 1959. A study of the New Zealand 

Chironomidae (Diptera, Nematocera). Bull. 

Brit. Mus. (Nat. Hist.), Ent. 7: 393–437. 

Freeman, P. 1961. The Chironomidae (Diptera) of 

Australia. Austr. J. Zool. 9: 611–737. 

Freeman, P. 1964a. A new species of Diamesa 

(Diptera, Chironomidae) from Africa south of 

the Sahara. Ann. (K.K.) Naturhist. Mus. Wien 

67: 407–408. 

Freeman, P. 1964b. Notes on Chironomidae (Diptera: 

Nematocera). Proc. R. Ent. Soc. Lond. (B) 

33: 147–150. 

Freeman, P. & Cranston, P.S. 1980. Family Chironomidae. 

In Crosskey, R.W. (ed.), Catalogue 

of the Diptera of the Afrotropical Region, pp. 

175–202. British Museum (Natural History), 

London.

Abstract 

A recent chironomid record of three shallow, high 

altitude lakes in southern Tibet, as well as a short 

palaeolimnological history of one lake, are presented. 

The recent chironomid assemblages consisted 

of 13 taxa; one of the Orthocladiinae taxa 

recorded most likely represents a new species. In 

spite of the low head capsule concentration in the 

sediment core of Lake Karuugema, probably due 

to physical disturbance, redistribution and outwash 

of head capsules, there was a trend from assemblages 

composed of stenothermal/rheophilic 

Current researCh 

Short comment on chironomid aSSemblageS and Stratigraphy of high 

altitude lakeS from tibet 

Ladislav Hamerlík 1,2* , Kirsten S. Christoffersen 1 , Klaus P. Brodersen 1 

1 Freshwater Biological Laboratory, Department of Biology, University of Copenhagen, Helsingørsgade 

51, DK-3400, Hillerød, Denmark 

E-mail: ladislav.hamerlik@savba.sk 

2 Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, SK-84506, Bratislava, Slovakia 

* Corresponding author 

taxa to eurythermal/limnophilic taxa. This shift 

in assemblage structure suggests that changes in 

monsoon precipitation and catchment hydrology 

may have influenced the habitat conditions of the 

chironomids. 

Introduction 

Our knowledge of chironomids of Tibetan Plateau, 

whether recent or subfossil, is fairly limited. In this 

short note, we aim to report the composition of 

recent chironomid assemblages of three shallow, 

high altitude lakes in southern Tibet, and to investigate 

a short palaeolimnological history of one of 

Figure 1. Photography of Lake Karuugema (A) and the sediment core (B) that was used for the chironomid analysis. 

The two other lakes, Sharmar tso (C) and Sijing la tso (D) are also pictured. Photo: K. S. Christoffersen. 

20

Table 1. Basic parameters of the lakes surveyed. 

Parameter Karuugema Sharmar tso Sijing la tso 

Coordinates 28º42.793’N 85º53.441’E 30º02.056’N 90º24.355’E 29º46.613’N 82º22.887’E 

Altitude (m asl) 4,654 4,401 4,925 

Average depth (m) 0.4 4.6 6.5 

Conductivity (µS/cm) 855 220 18 

O2 (mg/L) (top/bottom) 8.2 9.3/7.2 2.3/2.1 

pH 9.1 8.9 7.2 

the lakes as it is situated in an area with extreme 

climate (cold winters and hot summers). 

Material and methods 

Three shallow lakes (Fig. 1A, C, D) located on the 

Tibetan Plateau were surveyed during July 2009. 

The basic lake parameters were measured using 

a multiprobe (YSI-556) and are presented in 

Table 1. The sediment cores were retrieved from 

the middle part of the lakes with a Kajak corer by 

hand and sectioned on site into 1 cm slices using 

standard equipment. Samples were collected in zip 

lock bags and stored cold for transportation to the 

laboratory. 

To assess the recent chironomid assemblages, the 

uppermost 4 cm layers of the sediment cores were 

used. Furthermore, a 19 cm long sediment core of 

Lake Karuugema was used to carry out a palaeoecological 

reconstruction (Fig. 1A, B). Lake Karuugema 

is situated on a large plateau with small 

moraine hills surrounded by larger mountains. The 

surrounding bedrock consists of carbonate and 

clastic deposits of Triassic-Jurassic age. The area 

is seasonally frozen and the catchment is characterized 

by sparse, high–altitude desert vegetation. 

The lake is permanent but undergoes seasonal 

changes in water level. At the time of visit the max 

depth was approx. 1.2 m and the mean lake depth 

was estimated to be ~40 cm. The bottom consists 

of sandy sediment with very sparse vegetation. 

Calcium precipitation was observed on the plants. 

Given the low lake depth, wind exposure, physical 

disturbance and winter freezing will influence the 

biota directly, but also indirect influence through 

in-lake processes will occur. 

The samples were processed for chironomid analysis 

according to standard methods: freeze-dried 

sub-samples were deflocculated for 10–20 min 

in 10% KOH heated to 75 ºC (Walker & Paterson 

1985). The sediment was passed through a 

90 μm mesh sieve. The chironomid head capsules 

were hand picked under a binocular microscope 

(40×), dehydrated in 99% ethanol and mounted 

in Euparal®. Identification was performed under 

a compound microscope at 400× magnification, 

with reference to Wiederholm (1983), Roback & 

Coffman (1987) and Brooks et al. (2007). Most of 

the specimens were identified to genus level, and, 

in some cases, to species-morphotype level. Fragments 

that consisted of more than half the mentum 

were counted as a whole head capsule, fragments 

that consisted of half the mentum were counted as 

half a head capsule and smaller fragments were 

Table 2. Relative abundances of chironomid taxa recorded in the uppermost 4 cm of the lake sediments. 

Taxon Karuugema (%) Sharmar tso (%) Sijing la tso (%) 

Procladius (H.) sp. 8 - - 

Orthocladiinae indet. 7 - 5 

Paracladius sp. 8 - - 

Psectrocladius sordidellus-type - - 12 

Smittia/Parasmittia sp. - 11 - 

Tvetenia discoloripes gr. 1 - - 

Rheocricotopus sp. - 11 - 

Chironomus plumosus-type 4 - 2 

Chironomus anthracinus-type 3 67 5 

Corynocera oliveri-type 1 - - 

Micropsectra sp. 66 11 29 

Paratanytarsus sp. 1 - 48 

Tanytarsus sp. 2 - - 

21

excluded. β-diversity of the sediment sequence 

was expressed as DCA gradient-length in SD (ter 

Braak & Šmilauer 2002). 

Results and discussion 

Recent chironomid assemblages 

Altogether, 13 taxa were recorded in the uppermost 

4 cm sediment layers of the three lakes sampled 

(Table 2). The most frequent were Chironomus anthracinus 

type and Micropsectra sp. occurring in 

all three lakes, followed by Orthocladiinae indet., 

Chironomus plumosus type and Paratanytarsus 

sp. Taxa of the subfamily Chironominae (such as 

Micropsectra sp., C. anthracinus type and Paratanytarsus 

sp.) dominated numerically in all three 

lakes. 

According to Williams (1991), Chironomus reductus 

(as Tendipes reductus) was the most important 

chironomid and formed a significant part of the total 

benthic biomass of the high altitude saline lakes 

of Quinghai region (China). 

During the investigation of 42 lakes of the Tibetan 

Plateau, species of the Psectrocladius genera (especially 

P. sordidellus type), Cricotopus/Orthocladius 

sp., Paratanytarsus sp., Tanytarsus spp. and 

Chironomus spp. were the dominating taxa (Zhang 

et al. 2007). Surprisingly, species of Cricotopus/ 

Orthocladius genera were not recorded and both 

Tanytarsus sp. and P. sordidellus type were relatively 

rare in our samples. Moreover, the most 

abundant taxon of the present study, Micropsectra 

sp., was not recorded by Zhang et al. (2007). However, 

the genus Micropsectra is the dominant Tanytarsini 

in Nepalese Himalayas, especially above 

2000 m asl (Roback & Coffman 1987). According 

to the head capsule characteristics, (dark brown 

head and first two antennal segments, very short 

spur on the antennal pedestals and short pedicels 

of Lauterborn organs) the morpho-type dominating 

in our sediment cores seems to be the same 

as described in Roback & Coffman (1987) and 

also found widespread in the lakes of the Nepalese 

Himalayas by Manca et al. (1998). According to 

the imagos found by Manca et al. (1998), this larva 

type could belong to a species close to Micropsectra 

nepalensis Säwedal. 

One taxa of the Orthocladiinae subfamily was recorded 

which did not resemble known genera and 

which most likely represents a new species. The 

morpho-type called Orthocladiinae indet. (Fig. 2) 

has also been recorded from other lakes in Tibet 

(Tang, pers. comm.). The head capsule has plumose 

S1 setae, 5-segmented antenna and simple 

premandibles. Mandibles bear 3 inner teeth and 

22 

2 robust dorsal teeth, partially obscuring the inner 

teeth. The mentum is very characteristic with 

3 pale median teeth and markedly darker lateral 

teeth, which are folded back (resembling e.g. Corynocera 

ambigua); there is also a little rounded 

basolateral tooth on the mentum, which is characteristic 

for Limnophyes and relative species. The 

shape of the mandibles and the mentum suggests 

that this species is most likely a collector-gatherer, 

feeding on fine organic particles of the detritus. 

This combination of the head capsule characteristics 

did not allow us to place this type to the existing 

genera with certainty. However, some characteristics 

indicate that it might be a species close 

to Limnophyes or Thienemannia (Cranston, pers. 

comm.). 

Figure 2. Photographs of the mentum (A), inner (B1) 

and dorsal teeth (B2) of the mandible, and the 5-segmented 

antenna (C) of the unidentified specimen (Orthocladiinae 

indet.) found in the sediment core. 

Chironomid stratigraphy of Lake Karuugema 

The age of the 19 cm long sediment core was not 

identified; however, given the high altitude of the 

lake and the very low amount of the fine organic 

accumulation in the sediment, the sedimentation 

rate of the lake is most likely similar to that of 

other arctic and/or high altitude lakes and is consequently 

very low. In our estimation, the sediment 

core may represent several hundred years. There 

was a great amount of fine mineral sediments, such 

as sand, mica and quartz throughout the sediment

Figure 3. Chironomid stratigraphy of Lake Karuugema. Chironomid taxa are given as percentages of the total number 

of head capsules; sum represents the total number of head capsules per sample. Taxa are ordered according to weighed 

average score. 

core. It is probably originated from soils and bedrock 

as well as from the surrounding glaciers and 

is transferred to the lake through its intense inlets. 

In total, 13 chironomid taxa of 3 subfamilies were 

recorded (Fig. 3). Besides the head capsules, also 

the larva of Tvetenia discoloripes group was found 

in the uppermost sediment layer, however, given 

the only occurrence of this taxon, it was excluded 

from the analysis. Generally, the sediment core 

was fairly poor in chironomid head capsules and 

their density varied markedly from layer to layer. 

A maximum of 100 head capsules was counted 

per sample, however, at some layers only 10 head 

capsules were found. From the bottom up to 7 cm, 

the abundance was constantly low, without obvious 

oscillations. From the 7-8 cm layer, however, 

the abundance increased rapidly to 3-4 cm, where 

it reached the maximum and started to decrease to 

the same low abundance level as before. Taking the 

whole core into account, the most abundant taxa 

were Micropsectra sp. (54% of all head capsules), 

Paracladius sp. (9.5%), Paratanytarsus sp (8.7%), 

Chironomus anthracinus-type (8.3%), Procladius 

(H.) sp. (5.9%) and Orthocladiinae indet. (4.9%). 

These taxa were also the most constant along the 

sediment core, occurring at least in the half of all 

samples. Despite of its low total abundance, also 

Brillia sp. had high frequency occurring in 40% 

of all samples. Most of the above taxa have been 

found previously in lakes of the Tibetan Plateau 

(Zhang et al. 2007, Chen et al. 2009). 

The species turn-over was relatively high (DCA 

gradient length 2.5 SD) and there was a gradual 

but obvious shift in taxonomic composition. Most 

of the taxa were represented throughout the sediment 

core, however, some taxa were characteristic 

at the base, while others for its upper, most recent 

portion. Orthocladiinae Brillia sp. and Hydrobaenus 

sp. occurred mainly towards the base, for 

23 

which also very low abundance was characteristic. 

Species of the genus Hydrobaenus are mainly 

cold stenothermal, but ecologically rather diverse, 

dwelling the littoral of oligotrophic lakes, ponds, 

puddles, rivers and streams. Some species aestivate 

during the summer season (Sæther 1976). Species 

of the genus Brillia are considered to be primarily 

rheophilic, but also occur in littoral and hygropetric 

zones of lakes (Wiederholm, 1983). High 

frequency of both taxa mentioned in the bottom 

layers linked with low total abundance could be an 

indicator of strong influence of inlet streams. 

On the contrary, taxa only occurring in the upper 

portion of the core were Chironominae and 

Tanypodinae, such as Corynocera oliveri type, 

Chironomus plumosus type, Tanytarsus sp. and 

Procladius (H.) sp. Moreover, in these layers, the 

proportion of Orthocladiinae was lower then in the 

deeper layers. Appearance of Chironominae (especially 

C. plumosus type) and Tanypodinae preferring 

fine sediments, linked with increasing abundance 

suggests lower intensity of inlet streams, 

higher sedimentation rate, more stable environment, 

supporting higher abundance and occurrence 

of taxa preferring fine sediments. There is likely 

to be a linkage of inflow intensity and the amount 

of monsoon precipitation, found by Morrill et al. 

(2006) in another Central Tibetan lake. They found 

significant differences in monsoon precipitation 

linked with lake depth since early Holocene, with 

recently increased precipitation following a late 

Holocene dry period, which is in accordance with 

our results. 

Conclusion 

The new records of chironomids from the three 

studied lakes extend the taxa-list of the region and 

will be valuable for taxonomists and limnologists 

aiming for future studies in high altitude lakes in

Tibet. In spite of the low head capsule concentration 

in the sediment core of Lake Karuugema, 

probably due to physical disturbance, redistribution 

and outwash of head capsules, there was a 

trend from assemblages composed of stenothermal/rheophilic 

taxa to eurythermal/limnophilic 

taxa. This shift in assemblage structure suggests 

that changes in monsoon precipitation and catchment 

hydrology may have influenced the habitat 

conditions of the chironomids. 

Acknowledgement 

The authors wish to thank Anders Kjær and his 

staff at Innovation Center Denmark in Shanghai 

for the logistic and financial support throughout 

the project. Prof. Z. Renbin from Nanjing Institute 

of Geology and Paleontology, China, kindly 

provided information on the bedrock types. We are 

also grateful to Mr. J. Skafte for assistance during 

field work and to Prof. P. Bitušík from Matthias 

Belius University for his helpful comment to an 

earlier version of the manuscript. 

References 

Brooks, S.J., Langdon P.G & Heiri O. 2007. The 

Identification and Use of Palaearctic Chironomidae 

Larvae in Palaeoecology. QRA Technical 

Guide No. 10. - Quaternary Research Association, 

London, pp. 276. 

Chen, J., Chen, F., Zhang E., Brooks, S., Zhou, A. & 

Zhang J. 2009. A 1000-year chironomid-based 

salinity reconstruction from varved sediments 

of Sugan Lake, Qaidam Basin, arid Northwest 

China, and its palaeoclimatic significance. - 

Chinese Sci. Bull. 54(20): 3749-3759. 

Manca, M., Ruggiu, D., Panzani, P., Asioli, A., 

Mura, G. & Nocentini, A.M. 1998. Report on 

a collection of aquatic organisms from high 

24 

mountain lakes in the Khumbu Valley (Nepalese 

Himalayas). In Lami A. and Giussani G. 

(Eds) Limnology of high altitude lakes in the 

Mt Everest Region (Nepal). - Mem. Ist. Ital. Idrobiol. 

57: 77-98. 

Morrill, C., Overpeck, J.T., Cole, J.E., Liu, K., 

Shen, C. & Tang, L. 2006. Holocene variations 

in the Asian monsoon inferred from the geochemistry 

of lake sediments in central Tibet. 

- Quat. Res. 65: 232-243. 

Roback, S.S. & Coffman, W.P. 1987. Results of the 

Nepal Alpine Zone Research Project, Chironomidae 

(Diptera). - Proc. Acad. Nat. Sci. Phil. 

139: 87-158. 

Sæther, O.A. 1976. Revision of Hydrobaenus, 

Trissocladius, Zalutchia, Paratrissocladius, and 

some related genera (Diptera: Chironomidae). 

- Bull. Fish. Res. Bd Can. 195: 1-287. 

ter Braak, C.J.F. & Šmilauer, P. 2002. CANOCO 

Reference Manual and Users Guide to Canoco 

for Windows. Software for Canonical Community 

Ordination (version 4), Wageningen: Centre 

of Biometry. 

Walker, I.R. & Paterson, C.G. 1985. Efficient separation 

of subfossil Chironomidae from lake 

sediments. - Hydrobiologia 122: 189-192. 

Wiederholm, T. (ed.) 1983. Chironomidae of the 

Holarctic region Keys and diagnoses. Part I 

Larvae. - Ent. Scand., Suppl. 19: 1-457. 

Williams, W.D. 1991. Chinese and Mongolian saline 

lakes: a limnological overview. - Hydrobiologia 

210: 39-66. 

Zhang, E., Jones, R., Bedford, A., Langdon, P. & 

Tang., H. 2007. A chironomid-based salinity 

inference model from lakes on the Tibetan Pateau. 

- J. Paleolimn. 38: 477-491.

Chironomidae (inseCta, diptera) from alto paranapanema Basin, 

Abstract 

We investigated the community of Chironomidae 

from three rivers belonging to the same river basin 

in Southern Brazil. Our objective was to analyze 

if the Chironomidae communities from rivers 

of the same basin were similar and relate this to 

land-use and water quality variables. Samples of 

insects were taken using artificial substrate baskets 

and left 44 days in the field for colonization 

during the dry season in 2002. Study reaches with 

the relevant land-use category present for at least 

500 m along both river banks above and alongside 

the study reach were selected and land-use, terrain 

slope of the river basin and chemical and physical 

variables of the water were analysed. Faunal 

data were analyzed by number of individuals, richness 

of genera and community indices. Statistical 

analyses were performed in order to investigate 

the relationship between abiotic variables and the 

Chironomidae communities. Twenty-two genera 

were identified; Rheotanytarsus (Thienemann & 

Bause) was the most abundant in all assemblages. 

Some genera showed preferences in their distribution, 

and were observed in only one of the rivers. 

Land-use and slope of the terrain were similar for 

all rivers, while the water quality variables were 

different for the Taquari River compared to the two 

other sites. This may explain the differences in the 

Chironomidae community observed for this locality. 

Introduction 

The ever increasing human population has induced 

a need for more agricultural land to provide food 

and sources for bioenergy. The Southeastern Brazilian 

region is now over-exploited as a result of 

recent deforestation (also of the riparian forests) 

and replacement by crop plantations or pastureland 

for cattle (Loureiro 1998). 

The importance of the riparian forest to aquatic 

systems and their biota has been described extensively 

(Casatti et al. 2006; Matthaei et al. 2006). 

southeastern Brazil 

Kathia Sonoda 1* , Juaci Malaquias 1 , Carlos Vettorazzi 2 

1 Embrapa Cerrados, Rod BR 020 km 18, Caixa Postal 08223, CEP 73310-970, Planaltina, DF, Brazil 

E-mails: kathia.sonoda@cpac.embrapa.br, juaci.malaquias@cpac.embrapa.br 

2 ESALQ. Av. Pádua Dias, 11. Caixa Postal 9. CEP 13418-900. Piracicaba, SP, Brazil 

E-mail: cavettor@esalq.usp.br 

*Correspoding author 

25 

The usual conclusions in such studies focus on the 

negative aspect of the conversion of riparian forest 

to agricultural land (Cetra and Petrere 2007; 

Galbraith et al. 2008). Their presence is important 

for the maintenance of temperature equilibrium, 

as continous food supply for aquatic animals, as 

source of organic material and stabilization of 

stream banks (Marinho Filho and Reis 1989; Rodrigues 

1992; Aguiar et al. 2002). 

Because of their widespread distribution and the 

sensitiveness to pollution observed for some species, 

chironomids are used worldwide as biological 

indicators of environmental quality (Bacey and 

Spurlock 2007; Chessman et al. 2007; Smith et 

al. 2007). Several projects around the world demonstrate 

their application as bioindicators (WRC 

2001; MDFRC 2007). 

In Brazil, several studies have been conducted in 

order to analyse the influence of the surrounding 

land-use on the aquatic fauna (Sonoda 2005; 

Corbi and Trivinho-Strixino 2008). The interaction 

of terrestrial and aquatic systems is often analysed 

in small watersheds in an attempt to reduce 

the influence of extraneous environmental factors 

that, consequently, make it more difficult to detect 

the effects of differences in catchment landuse 

(Siqueira and Trivinho-Strixino 2005; Roque 

et al. 2008). Studies conducted on large Brazilian 

aquatic systems are rare in the scientific literature; 

as the velocity of river flow and sudden changes 

in the water level are some of the difficulties encountered. 

As stated by Vinson and Hawkins (2003), streams 

within similar biomes support similar number of 

taxa. This implies that the physical and biological 

environment of streams are a strong selective force 

on insect stream communities. 

The rivers studied here are similar with regard to 

land-use and cover, terrain slope, river channel 

morphology and climate. Our objective was to 

analyze if the Chironomidae communities from

Figure 1. Map of São Paulo State, with the location of the sampling sites (white dots). The insert shows the location 

of São Paulo State in Brazil. 

rivers of the same basin were similar and if possible 

to explain reasons for differences if these were 

found. 

Material and Methods 

Study Area 

Our study was carried out in the Alto Paranapanema 

River basin, State of São Paulo (Brazil) 

(Figure 1), which has a drainage area of 22,550 

km2 . The rivers selected for the study were Paranapanema, 

Apiaí-Guaçu and Taquari; a description 

of the sampling sites is given in Table 1. 

Based on previous methodological studies (Lammert 

and Allan 1999; Solimini et al. 2000; Cuffney 

et al. 2002), the following criterion was used when 

selecting our study reaches: the land-use analyzed 

(pasture or forest) had to be present for at least 500 

m length along both river banks, both above and 

alongside each study reach (Figure 2). 

Sampling procedure 

Six baskets (30 cm × 15 cm × 8 cm, mesh size 2.0 

cm) filled with artificial substrates [clay rocks of 

particle size 16-32 mm, classified as coarse gravel 

after Gordon et al. (1992)] were placed under water 

along the river banks during the dry season. After 

the 44 days of colonization, the baskets were 

removed, placed in 80 % alcohol and carried to the 

laboratory. There, they were washed under flowing 

water over a sieve of mesh size of 0.2 mm. 

The chironomid larvae were sorted and identified 

to genus following the key provided by Trivinho- 

Strixino and Strixino (1995). 

Land-use and water quality variables 

LANDSAT imagery was used to generate digital 

maps from which percentages of land-use distribution 

and terrain slopes upstream of the sampling 

areas were calculated. 

Chemical and physical variables were surveyed by 

Salomão (2004) who analyzed the following water 

variables: fine suspended solids (FSS), coarse suspended 

solids (CSS), total suspended solids (TSS), 

water temperature, pH, conductivity, dissolved 

oxygen, dissolved organic carbon, dissolved inorganic 

carbon, free CO , chloride, NO , NO , NH , 

2 2 3 4 

SO , Na, K, Mg, Ca (Table 2). 

4 

Data analysis 

The chironomids were analyzed as total number 

and percentage of individuals per genus, number 

of genera, taxon richness, Shannon-Wiener’s diversity 

index, Margalef’s richness, evenness index 

(Odum 1984; calculated using 2 as the base for the 

logarithm) and Sørensen’s modified index (Dise- 

Parameter Paranapanema River Apiaí-Guaçu River Taquari River 

Area of sub-basins (km2 Table 1. Basic description of the sampling sites in the Alto Paranapanema River Basin. 

) 710.39 899.74 828.08 

Depth (m) 0.5 1.4 1.8 

Width (m) 17.5 15 17 

Latitude 23º90’79” 23º93’11” 23º96’91” 

Longitude 48º25’96” 48º65’78” 48º94’61” 

26

Figure 2. Schematic drawing of the experimental 

design in each of the three rivers. Colonisation baskets 

were placed along both river margins at each of the 

three sampling points at all study sites. 

rud and Ødegaard 2007) that permits the integrated 

analysis of all sites. 

The X2 test was used to analyze the influence of 

location (rivers) on individual abundance and 

diversity indices. We utilized Pearson’s correlation 

and two-way ANOVA to integrate abiotic 

and biotic data, to analyze the interactions among 

them. All statistical analyses were performed using 

SAS® (version 9.1.2, The SAS Institute, Cary, 

NC) (Wright and Covich 2005). 

Results 

Watershed and water features 

The analysis of land-use and terrain-slope showed 

similar patterns at the sampled rivers within the 

watershed (Table 2); native vegetation was the 

main land-use followed by pasture and areas of 

reforestation. The percentage of the adjacent landuse 

was slightly different for the Apiaí-Guaçu 

River, showing the least reforestation (6.2 % of the 

area) and the most pasture land-use (39 %) when 

compared to the other rivers (Table 2). 

The slopes were divided into seven classes with 

Table 2. Main land-use and main slope-class of the 

river sub-basins upstream of the sampling sites. 

Land-use/ 

terrain-slope 

Paranapanema 

Apiaí- 

Guaçu 

Taquari 

Pasture 22.6% 39% 25.3% 

Native 

vegetation 

56.7% 43.7% 47.2% 

Reforestation 16.2% 6.2% 17.9% 

Paranapanema River and the Apiaí-Guaçu River; 

and 15 genera from the Taquari River (Table 4). 

Chironominae was the sub-family with highest 

number of individuals and also the greatest taxon 

richness; Tanytarsini was the most abundant tribe 

in all rivers. This was due to the high number of 

Rheotanytarsus, the numerically dominant genus, 

always representing more than 59 % of the assemblages 

and reaching 71.4 % in the Taquari River. 

In contrast, Procladiini was the rarest tribe represented 

only by two individuals belonging to Djalmabatista 

at the Apiaí-Guaçu River. 

At the Paranapanema River, Nanocladius and 

Ablabesmyia were frequent too, and at the Apiaí- 

Guaçu River, Nanocladius was the second most 

abundant genus after Rheotanytarsus (Table 4). 

Some genera were rare in all environments: Chironomus 

and Endotribelos were sampled only at 

the Paranapanema River, while Tribelos and Djalmabatista 

were found exclusively at the Apiaí- 

Guaçu River. The Taquari River had no unique 

taxa. 

The communities from the Paranapanema River 

Table 4. Relative distribution (%) of genera in each 

community. 

Genus Paranapanema 

Apiaí- 

Guaçu 

Taquari 

Beardius 2.35 3.71 0.33 

Chironomus 0.56 0.00 0.00 

Endotribelos 0.09 0.00 0.00 

Fissimentum 0.28 0,00 4.10 

Goeldichironomus 0,00 0.15 0.00 

Harnischia 0.85 0.15 1.33 

Lauterborniella 1.97 1.19 0.00 

Parachironomus 0.00 0.15 0.11 

Pol. (Asheum) 3.85 0.59 0.33 

Pol. (Polypedilum) 0.00 1.34 1.99 

Stenochironomus 0.09 0.00 0.11 

Tribelos 0.00 0.30 0.00 

Rheotanytarsus 59.81 59.50 71.43 

Corynoneura 3.38 4.15 3.54 

Lopescladius 0.19 0.00 1.33 

Nanocladius 10.33 16.77 3.32 

Ablabesmyia 11.27 1.48 9.63 

Labrundinia 2.54 8.01 1.11 

Larsia 0.19 0.15 0.22 

Nilotanypus 0.85 0.59 0.00 

Pentaneura 1.41 1.48 1.11 

Djalmabatista 0.00 0.30 0.00 

28 

Table 5. Biotic indices of the communities from the 

three rivers. 

River Diversity Richness Evenness 

Paranapanema 

0.74 1.59 0.18 

Apiaí- 

Guaçu 

0.75 1.70 0.18 

Taquari 0.53 1.43 0.13 

and the Apiaí-Guaçu River had similar values for 

diversity and evenness indices (Table 5), the differences 

seen in the richness index was due to the 

greater number of individuals in the Paranapanema 

assemblage. Taquari’s community showed lower 

values for all indices due to the marked dominance 

of Rheotanytarsus. The Sørensen’s modified 

similarity index indicated a medium level of 

similarity among communities from the three rivers 

(S=0.55). 

Discussion 

Watershed and water features 

As the aquatic ecosystems are tightly coupled with 

their catchments (Maloney et al. 2008, Lamberti 

et al. 2010), the selection of three rivers from the 

same basin permitted us to minimize variation 

in catchment-scale features such as surrounding 

land-use, terrain-slope, climatic conditions, and 

channel morphology. Such variation can confound 

the influence of water quality on the aquatic biota 

and should be eliminated if possible. Some studies 

have emphasised the great importance of land 

cover as the main factor defining the structure of 

the aquatic entomofauna (e.g. Corbi and Trivinho- 

Strixino 2008). Our analysis of land-use upstream 

of the sampling sites (Table 2) showed minimal 

differences among the rivers and led us to believe 

that the dissimilarity in chironomid community 

structure was influenced by other factors. 

Despite similarities in catchment features, water 

quality variables showed great differences between 

the rivers, where the Taquari River showed higher 

values of NO , SO , K, Mg, Na and Ca (Table 

3 4 

3). Conductivity is highly influenced by a range 

of chemical variables. For the Taquari River, sodium, 

magnesium and calcium ions were the ones 

with highest concentrations, however, the statistical 

analysis only returned a positive association 

of sulphate to this river. High values of limestone 

and sulphate are recorded in the soil surrounding 

the Alto Paranapanema River basin (Milléo et al. 

2008; CPRM 2009) and this might explain the 

high values of calcium ions and sulphate measured 

there.

There is restricted urban development in the area, 

thus, the main source for the high levels of ions 

probably is related to local agricultural practice. 

However, additional studies probably should be 

conducted to explore the source of the high ionlevel 

at this site. The high level of TSS at the 

Taquari site probably is not controlled by land-use 

as the watershed showed the same percentage of 

forest cover as the Paranapanema River, where 

the quantity of suspended solids was significantly 

lower (10 % of the TSS recorded at Taquari). 

Chironomidae fauna 

The subfamilies Chironominae, Orthocladiinae 

and Tanypodinae are quite common in Neotropical 

streams (Sanseverino and Nessimian 2008). 

The Chironominae frequently are described as the 

most abundant Chironomidae subfamily in the region 

(Suriano and Fonseca-Gessner 2004; Trivinho-Strixino 

and Strixino 2005; Mendes and Pinho 

2009) while Orthocladiinae is the most common 

subfamily in lotic systems with high frequency in 

rapids of streams and rivers (Coffman and Ferrington 

1984, 1996). 

The genus Rheotanytarsus, showed numerical 

dominance in all three rivers, with a remarkable 

abundance of individuals at the Taquari River; 

however no statistical significant correlation to 

any water variable was made that could explain 

the high abundance. The reason for the high abundance 

is not quite clear, but authors have found a 

positive relationship of the genus to high pollution 

level (Simião-Ferreira et al., 2009) of water 

bodies at Anápolis (GO) in the Brazilian Cerrado 

biome. On the other hand, authors have also established 

a direct correlation of its presence to good 

water-quality environments (Corbi and Trivinho- 

Strixino, 1999). It is described as typical of lotic 

environments and prefers rapid flux water due to 

its filtering habits (Higuti and Takeda, 2002). 

A high abundance of Rheotanytarsus was also recorded 

by Sonoda et al. (2009) in the study of the 

influence of land-use on the chironomid fauna of 

rivers of São Paulo State, Southeastern Brazil. In 

this study, the high abundance of Rheotanytarsus 

showed no correlation to land-use, occurring in 

rivers with both adjacent forests and pasture-land. 

Hepp et al. (2008) also recorded a high proportion 

of Rheotanytarsus in rivers of southern Brazil. In 

contrast, Jorcin and Nogueira (2008) found no 

Rheotanytarsus but a high number of Djamabatista 

in the waterfall of the Paranapanema reservoir. 

A high abundance of Djamabatista larvae was also 

recorded by Trivinho-Strixino and Strixino (2005) 

who analyzed the Chironomidae community in the 

29 

Ribeira River, spatially near the Paranapanema 

River Basin. Djamabatista larvae are good swimmers 

and prefer shallow water (Nessimian and 

Henriques-de-Oliveira 2005). As the rivers here 

studied showed no waterfall and rapids, this might 

explain why larvae of this genus were relatively 

rare in our study where only two individuals were 

sampled in the Apiaí-Guaçu River. 

As seen in our study, once the main landscape 

variables are fixed the observed divergence in the 

chironomid communities can be associated with 

significant differences in water quality variables. 

A genus that exemplifies this relationship well is 

Nilotanypus. The genus is considered to be intolerant 

to pollution (Smith and Cranston 1995) and its 

absence from the Taquari River could be a response 

to the poorer water quality at this site. Some Brazilian 

studies has shown how water quality variables 

can influence the entomofauna. Melo (2009) 

found distinctly different macroinvertebrate communities 

in nearby sites (streams) in Southeastern 

Brazil and concluded that these were a result of 

different levels of conductivity and stream size 

(orders). Futhermore, Roque et al. (2010) analyzed 

the Chironomidae fauna from 61 streams and discussed 

the importance of both local (conductivity) 

and broad (riparian forest cover) scales on the 

aquatic community composition in streams of São 

Paulo State. They observed a negative relationship 

between the percentage of riparian forest and generic 

diversity. Similar results have been reported 

from analyses of land-use and macroinvertebrate 

diversity in southeastern Brazil (Corbi 2006; Corbi 

and Trivinho-Strixino 2006; Sonoda et al. 2009). 

A lower generic richness and a dominance of only 

one taxon was observed at the Taquari River. Such 

characteristics are often observed in impacted systems 

(Stone et al. 2005) and influence the values of 

biotic indices. This is also seen in our results (Table 

5). The use of diversity and richness indices as 

appropriate metrics to evaluate stream conditions 

is defended by Suriano et al. (2010) who analyzed 

49 metrics to assess the conditions of streams of 

São Paulo State. Similarity measures are among 

the most common (and accepted) metrics for comparing 

sites or samples (Diserud and Ødegaard 

2007). The value of the Sørensen index indicated a 

medium degree of difference among the communities 

analysed here. 

In conclusion, this study demonstrates that rivers 

belonging to the same basin, with similar land 

characteristics may not necessarily support similar 

midge communities and that poorer water quality 

can be reflected in the chironomid community.

Acknowledgments 

We gratefully acknowledge the contributions of 

the following institutions or individuals to our 

research project: Dr Marcos Salomão for data on 

water quality variables, Dr Alexandre Silva, Dr 

Silvio Ferraz, Dr. Roberta Valente for landusedata 

and GIS consulting, professor Susana Trivinho-Strixino 

for the identification of Tanypodinae 

specimens, FAPESP (State of São Paulo Research 

Foundation) through the Biota/FAPESP Program 

(Proc. Nº 00/14242-6). We also acknowledge two 

anonymous reviewers and Dr. Torbjorn Ekrem for 

their comments and suggestions that contributed 

substantially to the improvement of this manuscript. 

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30 

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C. G. 2008. Using environmental and 

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Strixino, S. 2010. Untangling associations between 

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de Chironomidae (Diptera) em dois córregos 

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riparian quality using two complemen-

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an acid mine-waste impacted tropical stream 

– the chironomid story. In: Cranston, P.S. 

(Ed.). Chironomids: from genes to ecosystems. 

CSIRO, Australia. Pp. 161-173. 

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Carchini, G. 2000. Performance of different biotic 

indices and sampling methods in assessing 

water quality in the lowland stretch of the Tiber 

River. - Hydrobiologia 422/423:197-208. 

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insetos aquáticos de quatro bacias hidrográficas 

do Estado de São Paulo. PhD Dissertation, 

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Brazil. 124p. 

Sonoda, K. C., Matthaei, C. R. and Trivinho-Strixino, 

S. 2009. Contrasting land uses affect Chironomidae 

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K. W. J. and Reeve, J. D. 2005. Macroinvertebrate 

communities in agriculturally 

impacted Southern Illinois streams: patterns 

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907-917. 

32 

Suriano, M. T. and Fonseca-Gessner, A. A. 2004. 

Chironomidae (Diptera) larvae in streams of 

Parque Estadual de Campos do Jordão, São 

Paulo State, Brazil. - Acta Limnol. Brasil. 

16(2): 129-136. 

Suriano, M. T., Fonseca-Gessner, A. A., Roque, 

F. O. and Froehlich, C. G. 2010. Choice of 

macroinvertebrate metrics to evaluate stream 

conditions in Atlantic Forest, Brazil. - Environ. 

Monit. Ass. DOI 10.1007/s10661-010-1495-3. 

Trivinho-Strixino, S. and Strixino, G. 1995. Larvas 

de Chironomidae (Diptera) do Estado de 

São Paulo: guia de identificação e diagnose 

dos gêneros. PPG-ERN/ UFSCar, São Carlos. 

230p. 

Trivinho-Strixino, S. and Strixino, G. 2005. Chironomidae 

(Diptera) do Rio Ribeira (divisa dos 

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253. 

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geographical patterns in local stream 

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767. 

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Aberration in Chironomus pupa 

Short CommuniCationS 

Peter H. Langton 

5 Kylebeg Avenue, Mountsandel, Coleraine, Co. Londonderry, Northern Ireland, BT52 1JN, UK. 

E-mail: PHLangton@kylebegave.fsnet.co.uk 

I have long been interested in chironomid pupal aberrations 

(Langton, 1989) as indicators of how new 

forms can arise in an animal which is unable to compete 

in Darwinian terms. The photograph shows a 

Chironomus nudiventris pupal exuviae with four spurs 

on segment VIII; two dorsal, two ventral. The dark 

spurs are the usual ventral spurs for the genus and 

the light coloured ones are dorsal, the spurs diverging 

from each other by about 90° in the unmounted specimen. 

I have also recently seen a few Chironomus 

plumosus male exuviae from one site without any 

trace of genital sacs. The adults obviously eclosed 

successfully, but what had they got (or not got) at the 

back end? 

Reference 

Langton, P.H. 1989. Functional and phylogenetic interpretation of chironomid pupal structure. - Acta Biol. 

Debr. Oecol. Hung. 2:247-252. 

Cricotopus annulator with blue wing tips 

Peter H. Langton 

5 Kylebeg Avenue, Mountsandel, Coleraine, Co. Londonderry, Northern Ireland, BT52 1JN, UK. 

E-mail: PHLangton@kylebegave.fsnet.co.uk 

Daniel Mengella took a number of photographs of 

a chironomid in mid-October 2009 by his pond in 

Yorkshire, England, showing distinct blue tips to 

the wings. He mailed me for an identification and 

received the standard reply: please send a specimen. 

On April 28, 2010 he again noticed chironomids with 

blue tips to the wings, photographed them, and sent 

specimens for identification. The species is Cricotopus 

(C.) annulator, despite the strange appearance of 

the thoracic dorsum in the photograph. In alcohol 

and when mounted in Euparal, the blue disappears. 

Presumably it is a diffraction colour. I have never observed 

this phenomenon and wonder how widespread 

it is in the Chironomidae. CHIRONOMUS readers 

are encouraged to look out for it and report their findings 

to the journal. 

(The camera used was a Nikon D90 with a 80-200mm 

lens, which includes optical stabilisation) 

33

Trends in Chironomid Research 

Leena Thorat and Bimalendu Nath 

Department of Zoology, University of Pune, Pune-411007, India. 

E-mails: leenathorat@gmail.com, bbnath@unipune.ac.in 

Each model organism is favoured for its own forte, for instance, the Zebra fish, the Chick, Xenopus and 

Aplysia, all have been attracting a larger community of biologists for developmental biology, neurobiology 

and behavioural studies. Whereas, Drosophila could succeed in keeping researchers glued to vast areas of 

biological research apart from genetics for more than a century. On the other hand, chironomid midges have 

been witnessing a fluctuating research history. During their early days they were employed as pioneering 

models in developmental biology and chromosome studies. However, later the attention shifted towards 

studies on taxonomy and systematics and further the course of flow of chironomid research changed its 

trend to ecology, palaeolimnology and ecotoxicological research. In taking stock of the literature highlighting 

the use of Chironomus sp. as a model, there seems to be no room for divergent opinions that chironomid 

midges could never bag a top rank involving research driven by ‘model-oriented’ agenda. As evident from 

the numerical count of publications on midge-centered research topics, as models, they have always been 

restricted to only few groups of workers. 

In an attempt to look at the trend of research using chironomid midges as model organisms, we have carried 

out a thorough sampling of literature with the help of the bibliography listed by Hoffrichter (2000-2009) 

that appeared in the issues of Chironomus Newsletters as a standard for the survey. In the interest of brevity, 

we have considered the past one decade i.e. between 1999 and 2008 as a reference for the evaluation of the 

number of publications encompassing the diverse areas of chironomid research. 

Figure 1. Percentages of covered topics in Chironomidae literature 

The present survey reveals the dominance of mainly three distinct, inter-dependent interests in chironomid 

models which is reflected through their number of publications (Fig. 1). Throughout the decade under the 

present consideration, unequivocally, studies focusing on the ecological role of chironomid midges in intra-faunal 

and faunal-floral interactions and palaeolimnology have topped the rank. Being a major group of 

macro invertebrate fauna, reports addressing their role in different aquatic ecosystems have been evolving 

even prior to 1999 and have always had a major contribution in the community of Chironomidologists. 

Concomitantly, investigations concerning climate change and other environmental issues stimulated a large 

group of palaeolimnologists to adopt midges as test animals for studies predicting the past and future palaeoclimatic 

changes. The second largest field in the list is environmental toxicology that has grown along 

with the growing concern for ecological issues. Being one of the potent bioindicators of the aquatic biota, 

34

midges have become one among the few widely studied models for the examination of cause and effect 

studies in response to different kinds of toxic stressors circulating in the habitat. The third interest of Chironomidologists 

that has modestly frequented the score is the branch of systematics and taxonomy. With 

more and more new species of Chironomus being explored, chironomid taxonomy too saw its rise. However, 

it must be emphasized that in the present scenario, the list of chironomid taxonomists is fewer than 

what may be actually needed by the Chironomus research community. In most cases, prior knowledge of 

taxonomy is a pre-requisite before proceeding with investigations at the molecular level that involve DNA 

barcoding, molecular phylogeny and the like. This fact rings an alarm that in the near future, a huge number 

of species of Chironomus may become extinct before exploration, thus calling for the need of well trained 

taxonomists. The other branches like cell and molecular biology, genetics and chromosome studies, morphology 

and anatomy, developmental and reproductive biology display a uniform trend with a moderate 

score throughout the past ten years. In contrast, publications on immunology, parasitology and behavioural 

studies have been extremely scanty. 

This article that aims to highlight the quantitative estimation of the progress of chironomid research indicates 

that although possessing versatile potential, the chironomid midge has failed to embrace all the 

major areas of biological research. The technique-driven age of the 21st century demands the broadening of 

research horizons that will in turn popularize the chironomid midge as a model organism. Nevertheless, the 

interdependent nature of ecology, systematics, paleolimnology and environmental toxicology that causes 

them to intersect, points out a promising continuum and growth in these studies using midges, which is 

indeed a boon for the understanding of implications of the changing ecological scenario of the globe as a 

whole. 

Reference 

Hoffrichter O. 2000-2009. Current Bibliography. - CHIRONOMUS Newsletter on Chironomidae Research 

13 -22. 

News from chironomid research in India, University of Burdwan 

Dr. Abhijit Mazumdar 

Department of Zoology, University of Burdwan, Burdwan, 713 104, India 

E-mail: abhijitau@rediffmail.com 

The research project, TAXONOMY OF DIPTERA under the aegis of the “All India 

Coordinated project on Insect Taxonomy” funded by the Ministry of Environment & Forests, Govt. of India 

under Prof. P.K. Chaudhuri & Dr. A. Mazumdar implemented in 2002 is in progress in its second phase. 

New habitats have been explored and several new species have been identified that will be published in due 

course. Assistance in the form of material and literature is solicited for its successful execution. 

M.Phil. thesis: Effect of temperature and food on the developmental period of Glyptotendipes barbipes 

(Stæger), an abundant pond-dwelling chironomid. 

Mrs. Sharamita De (Chakravarti) 

Supervisor: Dr. A. Mazumdar, 

Summary: The midges were grown under three temperature regimes (22˚C, 26˚C, 30˚C) and food ratios 

[Ratio (I) 25 mg fish food and 2.5 mg Baker’s Yeast in 12 ml (IX) Martin Solution and Ratio, (II) 50 mg 

fish food and 5 mg Baker’s Yeast in 12 ml (IX) Martin Solution]. Photoperiod was maintained as 14:10 (L: 

D) within the rearing chamber. Correlating with studied parameters indicate that the maximum growth occurred 

at 30˚C and with food regime (i). With food regime (ii) results were not very encouraging. 

New Chironomidologist: 

Dr. Uttaran Majumdar 

Assistant Professor, Department of Zoology 

Hooghly Women’s College, Hooghly, West Bengal 712103, INDIA 

E-mail: uttaran_majumdar@yahoo.com 

Area of research: Systematics and phylogeny of high altitude chironomids. 

35

Implementing interactive identification keys is important for accurate and effective 

chironomid identification 

Andreas Plank 

Holsteinische Straße 23, 10717 Berlin, Germany 

E-mail: andreas.plank@naturwiki.net 

Computer based interactive identification programs have been developed recently, for instance the CD-key 

of Klink and Moller-Pillot (2003) and other identification programs developed using Lucid key software 

available on http://www.lucidcentral.com. They all assist the identification of chironomids in a flexible and 

interactive manner by means of classical dichotomous or multiple access keys. This helps a lot, but unfortunately 

those programs don’t allow the easy extension of their content. An attempt to do so was made in 

Plank (2010) with the development of the Chironomidae Identification Program CHIP, whose content pages 

can be altered in a similar manner to Wikipedia. But CHIP doesn’t have the facility to combine work online 

and, for instance, share photographs with other authors. 

An EU-Project “Key to nature” funded three years ago focused on paper-free identification tools for use 

within schools and universities across Europe, in which a few new alternatives were developed for determinations 

that can be shared and improved online. Based on contributions having a creative common license 

on the one hand, and a Wiki-platform with shared image repositories on the other, authors can contribute 

either keys restricted to editing or with open collaboration. Currently most available keys on that platform 

are just for plants, but hopefully over time more scientists will consider contributing identification keys for 

chironomids, making chironomid keys accessible also to expert assessment or monitoring. 

References 

Klink, A. G. and Moller-Pillot, H. K. M. 2003: Chironomidae Larvae―Key to the higher taxa and species 

of the lowlands of Northwestern Europe. Version: 1.0. CD-Key. ISBN: 90-75000-58-8. 

Plank, A. 2010. Chironomid-based inference models for Tibetan lakes aided by a newly developed chironomid 

identification key. Thesis, Freie Universität Berlin. Available from: http://www.diss.fu-berlin.de/diss/ 

receive/FUDISS_thesis_000000017941. 

Screen shot of a classical determination key on www.keytonature.eu with the possibility of step-by-step 

identification (not shown). 

36

Announcement and Invitation to the 

Dear colleagues, 

The NTNU Museum of Natural History and Archaeology would like to invite you to the 18th International 

Symposium on Chironomidae. 

The Symposium will take place at the Norwegian University of Science and Technology in Trondheim, July 

4-6, 2011, with a post-conference tour on July 7. 

We attempt to bring scientists and students from all over the world to Trondheim and hope many of you will 

consider this a great opportunity to present and discuss recent developments in Chironomidae research. 

Read more about the conference and register your interest by using the preregistration form at the symposium 

website (http://www.ntnu.no/vitenskapsmuseet/chironomidae-symposium). Please visit this site for 

regular updates on the available scientific and social programs. Preregistration will be open until the end of 

2010 while formal registration will start in January 2011. As we depend on the preregistration to estimate 

the approximate number of delegates, we kindly ask you to preregister as soon as possible. This will also 

give us an opportunity to inform you directly about updates in the symposium program per e-mail. 

Looking forward to see you in Trondheim! 

For inquiries, please contact the symposium committee: Chiro2011@vm.ntnu.no 

The Symposium Committee, 

Elisabeth Stur, Torbjørn Ekrem & Kaare Aagaard. 

A World Catalogue Of Chironomidae (Diptera). Part 2. Orthocladiinae 

Patrick Ashe 1 & James P. O’Connor 2 

1 33 Shelton Drive, Terenure, Dublin 12, Ireland, patrick.ashe@upcmail.ie 

2 National Museum of Ireland, Kildare Street, Dublin 2, Ireland 

Part 2 of A World Catalogue of Chironomidae (Diptera) is expected to be published in early 2011 (probably 

April or May). The number of pages will be approximately 1,000 and due to the large size will be printed 

in two sections (A & B) of about 500 pages each. The two sections will each weigh about 1.3 to 1.4 kg 

and to save on postage costs each section will be posted separately. A quote from the publisher has not yet 

been requested but the price for Part 2 copy is likely to be about Euro 84 to which the relevant amount of 

postage must be added. It is possible therefore to estimate the postage cost (Airmail only) to anywhere in 

the world. The estimated total cost of a copy of Part 2 plus postage and packing can be determined from 

the table given below. 

Ireland 

(incl. Northern Ireland) 

Britain Europe Rest of the world 

Postage Euro 16.00 Euro 26.00 Euro 26.00 Euro 36.00 

Catalogue Part 2 Euro 84.00 Euro 84.00 Euro 84.00 Euro 84.00 

TOTAL Euro 100.00 Euro 110.00 Euro 110.00 Euro 120.00 

The Catalogue is being published by The Irish Biogeographical Society in association with The National 

Museum of Ireland. Once Part 2 of the Catalogue is published and available for sale it will be announced on 

the website of the Irish Biogeographical Society: http://www.irishbiogeographicalsociety.com/ 

The International Electronic Payment System from your Bank account to our Bank account (The Irish 

Biogeographical Society account) worked very well for Part 1 and will be used for Part 2. Money received 

from the sale of Parts 1 and 2 will be used to help pay some of the cost of Parts 3 and 4. 

Anyone wishing to order a copy of either Part 1 or Part 2 can contact the senior author, Dr. Patrick Ashe by 

e-mail: patrick.ashe@upcmail.ie 

37

New book 

Illustrated guide to the Chironomidae of Japan 

This new comprehensive book on Chironomidae contains detailed descriptions of morphology and 

methodology (collection, rearing, mounting, SEM, molecular taxonomy) as well as notes on habitats, 

ecology and the use of chironomids as biological indicators. The book also includes keys to subfamilies, 

genera and main species, diagnoses and numerous detailed drawings and photographs of adult males, 

larvae and pupae. 

Front and sample page of Illustrated guide to the Chironomidae of Japan. 

Language: Japanese 

Editor: Japanese Association of Chironomidae Studies (JACS) 

Editorial board: S. Kondo, M. Yamamoto, T. Kobayashi, K. Hirabayashi, and K. Kawai 

Book size: A5 (W 148mm, H 210mm), 356 pp. + color plate 48 pp. 

ISBN 978-4-8599-1172-3 

Price: ¥ 12,600 (including sales tax 5%) 

Publisher: Bun’ichi-sôgô-shuppan Ltd. 

2-5 Nishi-gokenmachi, Shinjuku-ku, Tokyo, 162-0812 Japan 

Phone: +3-3235-7341 

Fax: +3-3269-1402 

How to order: 

The book can be ordered through the Japanese Amazon web site: http://www.amazon.co.jp/In-English/. 

38

CURRENT BIBLIOGRAPHY: 21 OCT. 2009 - 15 NOV. 2010 

Odwin Hoffrichter 

Institut für Biologie I, Albert-Ludwigs-Universität Freiburg, Hauptstr. 1, D-79104 Freiburg, Germany 

Email: odwin.hoffrichter@biologie.uni-freiburg.de 

In addition to supplements to the last two preceding years the publications of the present year are shown: 

almost one citation per day, which gives the average speed of publishing with regard to chironomids. The 

compilation was achieved, as usual, from many sources: databases, tables of contents of journals, 

references and citations of papers, inspection of many periodicals, lists and pdf's provided by authors 

(thanks to you!). In particular, publisher issued search alerts proved to be rich in results. As a rule, only 

printed titles are reported here with the occasional, but obviously increasing, exception of online-only 

journals (PLoS or BioMed Central journals, e.g.). Titles announced "in press", even with available DOI 

numbers, are not considered before printing. In general, online information should be retrieved elsewhere; 

best check the chironomid home page for eventual references regularly, or use individual websites with a 

host of chironomid-related data. Publications using chironomids as prey or food for animals are not 

treated comprehensively; in particular, studies with frozen midge larvae only for use to feed experimental 

animals are totally disregarded. 

Supplements to current bibliography 2008 

Gavrilă, L., Burlibaşa, L., Uşurelu, M. D., Radu, 

I., Magdalena, L. M., Ardelean, A. and 

Cărăbaş, M. 2008a. Chromosomal rearrangements 

in Chironomus sp. as genosensors for 

monitoring environmental pollution. - XX. Int. 

Congr. Genet., Berlin, Abstr. Book: 32. 

Massaferro, J., Ashworth, A. and Brooks, S. 

2008a. Quaternary fossil insects from South 

America. - In: Rabassa, J. (ed.): The Late 

Cenozoic of Patagonia and Tierra del Fuego. 

Devls Quatern. Sci. 11: 393-409. 

Medina, A. I., Schreiber, E. E. y Paggi, A. C. 

2008a. Distribución de Chironomidae 

(Diptera) en dos sistemas fluviales ritrónicos 

(Andino-serrano) de Argentina. - Revta Soc. 

ent. argent. 67: 69-79. 

Ostendorp, W., Gretler, T., Mainberger, M., 

Peintinger, M. and Schmieder, K. 2008a. 

Effects of mooring management on 

submerged vegetation, sediments and macroinvertebrates 

in Lake Constance, Germany. - 

Wetlands Ecol. Mgmt 17: 525-541. 

Rozdina, D., Raikova-Petrova, G., Marinova, R. 

and Uzunova, E. 2008a. Food spectrum and 

feeding of Barbus cyclolepis Heckel from the 

middle stream of Maritza River (Bulgaria). - 

Bulg. J. agric. Sci. 14: 209-213. 

Scheibler, E. E., Pozo, V. y Paggi, A. C. 2008a. 

Distribución espacio-temporal de larvas de 

Chironomidae (Diptera) en un arroyo andino 

(Uspallata, Mendoza, Argentina). - Revta Soc. 

ent. argent. 67: 45-58. 

39 

Siri, A., Donato, M. and Paggi, A. C. 2008a. New 

phytotelmic habitat of Metriocnemus 

eryngiotelmatus (Diptera: Chironomidae). - 

Revta Soc.ent. argent. 67: 113-115. 

Ruginis, T. 2008a. Diet and prey selectivity by 

age-0 brown trout (Salmo trutta L.) in 

different lowland streams of Lithuania. - Acta 

zool. lituan. 18: 139-146. 

Syväranta, J., Vesala, S., Rask, M., Ruuhijärvi, J. 

and Jones, R. I. 2008a. Evaluating the utility 

of stable isotope analyses of archived 

freshwater sample materials. - Hydrobiologia 

600: 121-130. 

Tarkowska-Kukuryk, M. 2008a. Influence of 

submerged vegetation on the diet of roach 

(Rutilus rutilus L.) in shallow Polesie lakes. - 

Teka Kom. Ochr. Kstalt. Środ. Przyr. OL PAN 

5A: 145-152. 

Supplements to current bibliography 2009 

Abbott, I. M., Sleeman, D. P. and Harrison, S. 

2009a. Bat activity affected by sewage 

effluent in Irish rivers. - Biol. Conserv. 142: 

2904-2914. 

Akbulut, M., Çelik, E. S., Odabaşi, D. A., Kaya, 

H., Selvi, K., Arslan, N. and Odabaşi, S. S. 

2009a. Seasonal distribution and composition 

of benthic macroinvertebrate communities in 

Menderes Creek, Çanakkale, Turkey. - 

Fresenius envir. Bull. 18: 2136-2145. 

Alahuta, J., Vuori, K.-M., Hellsten, S., Järvinen, 

M., Olin, M., Rask, M. and Palomäki, A. 

2009a. Defining the ecological status of small 

forest lakes using multiple biological quality

elements and palaeolimnological analysis. - 

Fundam. appl. Limnol. 175: 203-216. 

Al-Shami, S. A., Siti Azizah, M. N. Che Salmah, 

M. R. and Abu Hassan, A. 2009a. Preliminary 

study of phylogenetic relationship of rice field 

Chironomidae (Diptera) inferred from DNA 

sequences of mitochondrial cytochrome 

oxidase subunit I. - Am. J. appl. Sci. 6: 1004- 

1009. 

Álvarez-Cabria, M., Barquín, J. and Juanes, J. A. 

2009a. Spatial and seasonal variability of 

macroinvertebrate metrics: Do macroinvertebrate 

communities track river health? - 

Ecol. Indicators 10: 370-379. 

Amundsen, P.-A. and Knudsen, R. 2009a. Winter 

ecology of Arctic charr (Salvelinus alpinus) 

and brown trout (Salmo trutta) in a subarctic 

lake, Norway. - Aquat. Ecol. 43: 765-775. 

Anderson, N. J., Brodersen, K. P., Ryves, D. B., 

McGowan, S., Johansson, L. S., Jeppesen, E. 

and Leng, M. J. 2009a. Climate versus in-lake 

processes as controls on the development of 

community structure in a low-arctic lake 

(South-West Greenland). - Ecosystems 11: 

107-124. 

Andrič, M., Massaferro, J., Eicher, U., Ammann, 

B., Leuenberger, M. C., Martinčič, A., 

Marinova, E. and Brancelj, A. 2009a. A multiproxy 

Late-glacial palaeoenvironmental 

record from Lake Bled, Slovenia. - 

Hydrobiologia 631: 121-141. 

Axford, Y., Briner, J. P., Cooke, C. A., Francis, 

D. R., Michelutti, N., Miller, G. H., Smol, J. 

P., Thomas, E. K., Wilson, C. R. and Wolfe, 

A. P. 2009a. Recent changes in a remote 

Arctic lake are unique within the past 200,000 

years. - Proc. natn. Acad. Sci. U. S. A. 106: 

18443-18446 (+ suppl. inf.). 

Bae, M.-J. and Park, Y.-S. 2009a. Changes in 

benthic macroinvertebrate communities in 

response to natural disturbances in a stream. - 

J. Ecol. Field Biol. 32: 197-206. 

Berezina, N. A., Golubkov, S. M. i Gubelit, Yu. I. 

2009a. Struktura litoral'nykh zootsenosov v 

zone nitchatykh vodoroslei estuariya reki 

Nevy. Biol. vnutr. Vod 2009, 4: 48-56. [also 

published as: Structure of littoral zoocenoses 

in the macroalgae zones of the Neva River 

Estuary. - Inland Wat. Biol. 2: 340-347.] 

Campbell, L. M., Thacker, R., Barton, D., Muir, 

D. C. G., Greenwood, D. and Hecky, R. E. 

2009a. Re-engineering the eastern Lake Erie 

littoral food web: The trophic function of non- 

40 

indigenous Ponto-Caspian species. - J. Gt 

Lakes Res. 35: 224-231. 

Carver, S., Storey, A., Spafford, H., Lynas, J., 

Chandler, L. and Weinstein, P. 2009a. Salinity 

as a driver of aquatic invertebrate colonisation 

behaviour and distribution in the wheatbelt of 

Western Australia. - Hydrobiologia 617: 75- 

90. 

Catalan, J., Barbieri, M. G., Bartumeus, F., 

Bitušík, P., Botev, I., Brancelj, A., 

Cogălniceanu, D., Manca, M., Marchetto, A., 

Ognjanova-Rumenova, N., Pla, S., 

Rieradevall, M., Sorvari, S., Štefková, E., 

Stuchlík, E. and Ventura, M. 2009a. 

Ecological thresholds in European alpine 

lakes. - Freshwat. Biol. 54: 2494-2517. 

Chen, J. H, Chen, F. H, Zhang, E. L, Brooks, S. 

J., Zhou, A. F. and Zhang, J. W. 2009a. A 

1000-year chironomid-based salinity reconstruction 

from varved sediments of Sugan 

Lake, Qaidam Basin, arid Northwest China, 

and its palaeoclimatic significance. - Chin. 

Sci. Bull. 54: 3749-3759. 

Chibunda, R. T. 2009a. Chronic toxicity of 

mercury (HgCl2) to the benthic midge 

Chironomus riparius. - Int. J. envir. Res. 3: 

455-462. 

Coe, H. J., Kiffney, P. M., Pess, G. R., Kloehn, K. 

K. and McHenry, M. L. 2009a. Periphyton 

and invertebrate response to wood placement 

in large Pacific Coastal rivers. - River Res. 

Applics 25: 1025-1035. 

Contente, R. F., Stefanoni, M. F. and Spach, H. L. 

2009a. Size-related changes in diet of the 

slipper sole Trinectes paulistanus 

(Actinopterygii, Achiridae) juveniles in a 

subtropical Brazilian estuary. - Pan-Am. J. 

aquat. Sci. 4: 63-69. 

Córdova, S., Gaete, H., Aránguiz, F. y Figueroa, 

R. 2009a. Evaluación de la calidad de las 

aguas del estero Limache (Chile central), 

mediante bioindicadores y bioensayos. - Latin 

Am. J. aquat. Res. 37: 199-209. 

Cristina, T.-D., Camino, F.-A. and Francisco, G.- 

C. 2009a. Habitat selection and sampling 

design for ecological assessment of 

heterogeneous ponds using macroinvertebrates. 

- Aquat. Conserv. mar. Freshw. 

Ecosyst. 19: 786-796. 

Cross, F. R., Jackson, R. R. and Pollard, S. D. 

2009a. How blood-derived odor influences 

mate-choice decisions by a mosquito-eating

predator. - Proc. natn. Acad. Sci. U. S. A. 106: 

19416-19419. 

Czarnecka, M., Poznańska, M., Kobak, J. and 

Wolnomiejski, N. 2009a. The role of solid 

waste materials as habitats for macroinvertebrates 

in a lowland dam reservoir. - 


Deines, P., Wooller, M. J. and Grey, J. 2009a. 

Unravelling complexities in benthic food 

webs using a dual stable isotope (hydrogen 

and carbon) approach. - Freshwat. Biol. 54: 

2243-2251. 

Dessborn, L., Elmberg, J., Nummi, P., Pöysä, H. 

and Sjöberg, K. 2009a. Hatching in dabbling 

ducks and emergence in chironomids: a case 

of predator-prey synchrony? - Hydrobiologia 

636: 319-329. 

Diomandé, D., Bony, Y. K., Edia, E. O., Konan, 

K. F. et Gourène, G. 2009a. Diversité des 

macroinvertébrés benthiques de la rivière 

Agnéby (Côte d’Ivoire; Afrique de l’Ouest). - 

Eur. J. scient. Res. 35: 368-377. 

Donato, M., Massaferro, J. y Brooks, S. J. 2009a. 

Estado del conocimiento taxonómico de la 

fauna de Chironomidae (Diptera: Nematocera) 

de la Patagonia. - Revta Soc. ent. argent. 68: 

187-192. 

Duan, X., Wang, Z., Xu, M. and Zhang, K. 

2009a. Effect of streambed sediment on 

benthic ecology. - Int. J. Sediment Res. 24: 

325-338. 

Dulić, Z., Poleksić, V., Rašković, B., Lakić, N., 

Marković, Z., Živić, I. and Stanković, M. 

2009a. Assessment of the water quality of 

aquatic resources using biological methods. - 

Desalination Wat. Treat. 11: 264-274. 

Elnitsky, M. A., Benoit, J. B., Lopez-Martinez, 

G., Denlinger, D. L. and Lee Jr., R. E. 2009a. 

Osmoregulation and salinity tolerance in the 

Antarctic midge, Belgica antarctica: seawater 

exposure confers enhanced tolerance to 

freezing and dehydration. - J. exp. Biol. 212: 

2864-2871. 

Ersan, E., Altindağ, A., Ahiska, S. and Alaş, A. 

2009a. Zoobenthic fauna and seasonal 

changes of Mamasin Dam lake (central part of 

Turkey). - Afr. J. Biotechnol. 8: 4702-4707. 

Fedorova, I. A., Polukonova, N. V. i Petrov, N. 

V. 2009a. Tsitogeneticheskie effekty 

kholinotropnykh preparatov pri kombinorovannom 

deistvii na lichinok Chironomus 

plumosus (Diptera) in vivo. (Cytogenetic 

effects of cholinotropic preparations mixture 

41 

on Chironomus plumosus (Diptera) larvae in 

vivo.) - Tsitologiya 51: 849-855. 

Finn, M. A., Boulton, A. J. and Chessman, B. C. 

2009a. Ecological responses to artificial 

drought in two Australian rivers with differing 

water extraction. - Fundam. appl. Limnol. 

175: 231-248. 

Fjellheim, A., Raddum, G. G., Vandvik, V., 

Cogălniceanu, D., Boggero, A., Brancelj, A., 

Galas, J., Sporka, F., Vidinova, Y., Bitusik, P., 

Dumnicka, E., Găldean, N., Kownacki, A., 

Krno, I., Preda, E., Rişnoveanu, G. and 

Stuchlik, E. 2009a. Diversity and distribution 

patterns of benthic invertebrates along alpine 

gradients. A study of remote European 

freshwater lakes. - Fundam. appl. Limnol., 

spec. Issue 62: 167-190. 

Florencio, M., Serrano, L., Gómez-Rodríguez, C., 

Millán, A. and Díaz-Paniagua, C. 2009a. 

Inter- and intra-annual variations of 

macroinvertebrate assemblages are related to 

the hydroperiod in Mediterranean temporary 

ponds. - Hydrobiologia 634: 167-183. 

Free, G., Solimini, A. G., Rossaro, B., Marziali, 

L., Giacchini, R., Paracchini, B., Ghiani, M., 

Vaccaro, S., Gawlik, B. M., Fresner, R., 

Santner, G., Schönhuber, M. and Cardoso, A. 

C. 2009a. Modelling lake macroinvertebrate 

species in the shallow sublittoral: relative 

roles of habitat, lake morphology, aquatic 

chemistry and sediment composition. - 

Hydrobiologia 633: 123-136. 

Friberg, N., Dybkjær, J. B., Olafsson, J. S., 

Gislason, G. M., Larsen, S. E. and , T. L. 

2009a. Relationships between structure and 

function in streams contrasting in temperature. 

- Freshwat. Biol. 54: 2051-2068. 

Fu, Y., Sæther, O. A. and Wang, X. 2009a. 

Corynoneura Winnertz from East Asia, with a 

systematic review of the genus (Diptera: 

Chironomidae: Orthocladiinae). - Zootaxa 

2287: 1-44. 

Gantner, N., Hintelmann, H., Zheng, W. and 

Muir, D. C. 2009a. Variations in stable 

isotope fractionation of Hg in food webs of 

Arctic lakes. - Envir. Sci. Technol. 43: 9148- 

9154. 

Gavrila, L., Barbacar, N., Radu, I., Burlibasa, L., 

Magdalena, L.M., Ardelean, A., Cimponeriu, 

G. D., Popovici, E., Usurelu, M. D., Macovei, 

I., Morănescu, F. and Carabas, M. 2009a. 

New aspects concerning the study of dipteran 

polytene chromosomal phenotype elements in

Chironomus plumosus. - Roman. biotechnol. 

Lett. 14: 4625-4634. 

Guilizzoni, P., Massaferro, J., Lami, A., Piovano, 

E. L., Guevara, S. R., Formica, S. M., Daga, 

R., Rizzo, A. and Gerli, S. 2009a. 

Palaeolimnology of Lake Hess (Patagonia, 

Argentina): multi-proxy analyses of short 

sediment cores. - Hydrobiologia 631: 289- 

302. 

Hågvar, S. and Klanderud, K. 2009a. Effect of 

simulated environmental change on alpine soil 

arthropods. - Global Change Biol. 15: 2972- 

2980. 

Halpern, M., Shakèd, T. and Schumann, P. 2009a. 

Brachymonas chironomi sp. nov., isolated 

from a chironomid egg mass, and emended 

description of the genus Brachymonas. - Int. 

J. syst. evolut. Microbiol. 59: 3025-3029. 

Hamerlík. L. and Bitušík, P. 2009a. The 

distribution of littoral chironomids along an 

altitudinal gradient in High Tatra Mountain 

lakes: Could they be used as indicators of 

climate change? - Annls Limnol. 45: 145-156. 

Hayford, B. 2009a. First records of Podonominae 

(Diptera: Chironomidae) from Outer 

Mongolia, with notes on ecology and 

biogeographic distribution. - J. Kans. ent. Soc. 

82: 305-310. 

Heino, J., Tolonen, K. T., Kotanen, J. and 

Paasivirta, L. 2009a. Indicator groups and 

congruence of assemblage similatrity, species 

richness and environmental relationships in 

littoral macroinvertebrates. - Biodivers. 

Conserv. 18: 3085-3098. 

Johnson, A. A. and Kleve, M. G. 2009a. A new 

genus and species of mermithid (Nematoda) 

emerging from chironomid imagos eclosing 

from the Mississippi River headwaters area in 

northern Minnesota. - J. Parasite. 95: 1493- 

1495. 

Johnson, J. H., McKenna, J. E., Chalupnicki, M. 

A., Wallbridge, T. and Chiavelli, R. 2009a. 

Feeding ecology of lake whitefish larvae in 

eastern Lake Ontario. - J. Gt Lakes Res. 35: 

603-607. 

Johnson, J. H., Nack, C. C. and Chalupnicki, M. 

A. 2009a. Predation by fallfish (Semotilus 

corporalis) on Pacific salmon eggs in the 

Salmon River, New York. - J. Gt Lakes Res. 

35: 630-633. 

Kara, T. ve Tellioğlu, A. 2009a. Büyük Çay 

(Pelte, Elazığ)'da Bulunan Chironomidae 

Larvalarının Taksonomik Yönden 

42 

İncelenmesi. (Taxonomical investigation of 

Chironomidae larvae at Büyük Stream ((Pelte, 

Elazığ).) - Firat Üniv. Fen Bilimleri Derg. 21: 

117-121. 

Kato, Y., Takemon, Y. and Hori, M. 2009a. 

Invertebrate assemblages in relation to habitat 

types on a floating mat in Mizorogaike Pond, 

Kyoto, Japan. - Limnology 10: 167-176. 

Kebede, Y. K., Assefa, F. and Amsalu, A. 2009a. 

Environmental impact of coffee processing 

effluent on the ecological integrity of rivers 

found in Gomma Woreda of Jimma Zone, 

Ethiopia. - Proc. Wld envir. Wat. Resourc. 

Congr. 2009: Great Rivers 342: 2462-2471. 

Kernan, M., Brancelj, A., Clarke, G., Lami, A., 

Raddum, G., Straškrábová, V., Stuchlík, E., 

Velle, G. and Ventura, M. 2009a. 

Environmental and biological characteristics 

of high altitude lochs in Scotland. - Fundam. 

appl. Limnol., spec. Issue 62: 379-417. 

Kernan, M., Catalan, J., Ventura, M. and Curtis, 

C. J. 2009a. A biological survey of high 

mountain and high latitude lakes across 

Europe: aims, sampling strategy, methods and 

main achievements. - Fundam. appl. Limnol., 

spec. Issue 62: 3-16. 

Kernan, M., Ventura, M., Bitušík, P., Brancelj, 

A., Clarke, G., Velle, G., Raddum, G. G., 

Stuchlík, E. and Catalan, J. 2009a. 

Regionalisation of remote European mountain 

lake ecosystems according to their biota: 

environmental versus geographical patterns. - 

Freshwat. Biol. 54: 2470-2493. 

Khangarot, B. S. and Das, S. 2009a. Acute 

toxicity of metals and reference toxicants to a 

freshwater ostracod, Cypris subglobosa 

Sowerby, 1840 and correlation to EC50 values 

of other test models. - J. hazard. Mat. 172: 

641-649. 

Ki, J.-S., Kim, I.-C. and Lee, J.-S. 2009a. 

Comparative analysis of nuclear ribosomal 

DNA from the moon jelly Aurelia sp.1 

(Cnidaria: Scyphozoa) with characterizations 

of the 18S, 28S genes, and the intergenic 

spacer (IGS). - Hydrobiologia 616: 229-239. 

Kilroy, C., Larned, S. T. and Biggs, B. J. F. 

2009a. The non-indigenous diatom 

Didymosphenia geminata alters benthic 

communities in New Zealand rivers. - 

Freshwat.. Biol. 54: 1990-2002. 

Krasheninnikov, A. B. i Makarchenko, E. A. 

2009b. K faune khironomid podsemeistv 

Podonominae, Diamesinae i Orthocladiinae

(Diptera, Chironomidae) zapovednika 

Visherskii i prilegayushchikh territorii 

(Severnyi Ural). (On the chironomid fauna of 

subfamilies Podonominae, Diamesinae and 

Orthocladiinae (Diptera, Chironomidae) of the 

Vishersky Nature Reserve and bordering 

territories (North Urals).) - Evraziat. ent. Zh. 

(= Euroas. ent. J.) 8: 335-340. 

Krestian, B. J., Kosnicki, E., Spindler, P. H., 

Stringer, S. and Epler, J. H. 2009a. First 

nearctic records of Oliveiriella Wiedenbrug 

and Fittkau, with new distributional records 

for two other new world species of 

Orthocladiinae (Diptera: Chironomidae). - 

Ent. News 120: 349-362. 

Kullman, M. A., Kidd, K. A., Podemski, C. L., 

Paterson, M. J. and Blanchfield, P. J. 2009a. 

Assimilation of freshwater salmonid 

aquaculture waste by native aquatic biota. - 

Can. J. Fish. aquat. Sci. 66: 1965-1975. 

Lamentowicz, M., Balwierz, Z., Forysiak, J., 

Płóciennik, M., Kittel, P., Kloss, M., Twardy, 

J., Żurek, S. and Pawlyta, J. 2009a. 

Multiproxy study of anthropogenic and 

climate changes in the last two millennia from 

a small mire in central Poland. - 

Hydrobiologia 631: 213-230. 

Larocque-Tobler, I., Grosjean, M., Heiri, O. and 

Trachsel, M. 2009a. High-resolution 

chironomid-inferred temperature history since 

ad 1580 from varved Lake Silvaplana, 

Switzerland: comparison with local and 

regional reconstructions. - Holocene 19: 1201- 

1212. 

Li, X., Zhang, Xin, Zhang, J., Zhang, Xing, 

Starkey, S. R. and Zhu, K. Y. 2009a. 

Identification and characterization of eleven 

glutathione S-transferase genes from the 

aquatic midge Chironomus tentans (Diptera: 

Chironomidae). - Insect Biochem. molec. Biol. 

39: 745-754. 

Luoto, T. P., Sarmaja-Korjonen, K., Nevalainen, 

L. and Kauppila, T. 2009a. A 700 year record 

of temperature and nutrient changes in a small 

eutrophied Lake in southern Finland. - 

Holocene 19: 1063-1072. 

Makarchenko, E. A. and Makarchenko, M. A. 

2009f. Novye nakhodki khironomid (Diptera, 

Chironomidae) na Dal'nem Vostoke i 

sopredel'nykh territoriyakh. V(Diptera, 

Orthocladiinae) from the Far East and 

bordwering territories. III. Podsemeistvo 

Orthocladiinae. (New records of chironomids 

(Diptera, Chironomidae) from the Far East 

43 

and bordering territories. VIII. Subfamily 

Orthocladiinae.) - Evraziat. ent. Zh. (= 

Euroas. ent. J.) 8: 326-334. 

Marchetto, A., Rogora, M., Boggero, A., 

Musazzi, S., Lami, A., Lotter, A. F., Tolotti, 

M., Thies, H., Psenner, R., Massaferro, J. and 

Barbieri, A. 2009a. Response of Alpine lakes 

to major environmental gradients, as detected 

through planktonic, benthic and sedimentary 

assemblages. - Fundam. appl. Limnol., spec. 

Issue 62: 419-440. 

Markiyanova, M. F. 2009a. Tsitogeneticheskaya 

struktura populyatsii Chironomus plumosus 

(L.) (Diptera, Chironomidae) Kurshskogo 

zaliva Baltiiskogo morya v svyazi s 

izmeneniem ego trioficheskogo statusa. - Biol. 

vnutr. Vod 2009, 4: 44-47. [also published as: 

Cytogenetic structure of Chironomus 

plumosus (L.) (Diptera, Chironomidae) 

population in regard to the trophic state of the 

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2010a. Seasonal abundance, diet, and energy 

consumption of round gobies (Neogobius 

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- Ecol. Freshwat. Fish 19: 206-215. 

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Planelló, R., Martínez-Guitarte, J. L. and 

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Poulton, B. C., Allert, A. L., Besser, J. M., 

Schmitt, C. J., Brumbaugh, W. G. and 

Fairchild, J. F. 2010a. A macroinvertebrate 

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Fundam. appl. Limnol. 176: 61-73. 

Ravinet, M., Syväranta, J., Jones, R. I. and Grey, 

J. 2010a. A trophic pathway from biogenic 

methane supports fish biomass in a temperate 

lake ecosystem. - Oikos 119: 409-416. 

Raz, N., Danin-Poleg, Y., Broza, Y. Y., Arakawa, 

E., Ramakrishna, B. S., Broza, M.. and Kashi, 

Y. 2010a. Environmental monitoring of Vibrio 

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Rees, A. B. H. and Cwynar, L. C. 2010a. 

Evidence for early postglacial warming in 

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Quatern. Sci. Rev. 29: 443-454. 

Reynaga, M. C. and Martín, P. R. 2010a. Trophic 

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55 

Roque, F. O., Siqueira, T., Bini, L. M., Ribeiro, 

M. C., Tambosi, L. R., Ciocheti, G. and 

Trivinho-Strixino, S. 2010a. Untangling 

associations between chironomid taxa in 

Neotropical streams using local and landscape 

filters. - Freshwat. Biol. 55: 847-865. 

Rosa, R., Moreira-Santos, M., Lopes, I., Silva, L., 

Rebola, J., Mendonça, E., Picado, A. and 

Ribeiro, R. 2010a. Comparison of a test 

battery for assessing the toxicity of a 

bleached-kraft pulp mill effluent before and 

after secondary treatment implementation. - 

Envir. Monit. Assess. 161: 439-451. 

Roskosch, A., Lewandowski, J., Bergmann, R., 

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Identification of transport processes in 

bioirrigated muddy sediments by 

[18F]fluoride PET (Positron Emission 

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1094-1097. 

Roskosch, A., Morad, M. R., Khalili, A. and 

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on lakes using the chironomid pupal exuvial 

technique. - Ecol. Indicators 10: 594-601. 

Rybak, J. and Sadłek, W. 2010a. Ecological 

impact of a dam on benthic macroinvertebrates 

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from Manitoba, Canada (Diptera: Chironomidae), 

- Aquat. Insects 32: 167-190. 

Sæther, O. A., Andersen, T., Pinho, L. C. and 

Mendes, H. F. 2010a. The problem with 

Polypedilum Kieffer (Diptera: Chironomidae), 

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Sanseverino, A. M., Trivinho-Strixino, S. and 

Nessimian, J. L. 2010a. Taxonomic status of 

Nimbocera Reiss, 19762, a junior synonym of 

Tanytarsus van der Wulp, 1874 (Diptera: 

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Saulnier-Talbot, É. and Pienitz, R. 2010a. 

Postglacial chironomid assemblage succession 

in northernmost Ungava Peninsula, Canada. - 

J. Quatern. Sci. 25: 203-213. 

Scharold, J. V., Corry, T. D., Bolgrien, D. W. and 

Angradi, T. R. 2010a. Spatial variation in the 

invertebrate macrobenthos of three large 

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Schlekat, C. E., Van Genderen, E., De 

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Schoenly, K. G., Cohen, J. E., Heong, K. L., 

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dung on a water–sediment system with the 

aquatic invertebrates Daphnia magna and 

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Senderovich, Y., Izhaki, I. and Halpern, M. 

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Inversionnyi polimorfizm Glyptotendipes 

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of Glyptotendipes glaucus Mg. 

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artificial subtropical ponds of Taiwan. - 


Shiels, D. R. 2010a. Implementing landscape 

indices to predict stream water quality in an 

agricultural setting: An assessment of the 

56 

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Stoaks, R. 2010a. Rare and unusual arthropod 

ecology, behavior and occurrence at 

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Tang, H., Song, M.-Y., Cho, W.-S., Park, Y.-S. 

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87. [also published as: Food spectra of the 

farm juveniles of the chum salmon 

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molecular phylogeny of Anisofilariata 

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Tolonen, K.T, and Hämäläinen, H. 2010a. 

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reconstruction derived from a combined 

biogenic Si and chironomid record from 

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Tremblay, V., Larocque-Tobler, I. and Sirois, P. 

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Chemosphere 80: 53-59. 

57 

Tronstad, L. M., Tronstad, B. P. and Benke, A. C. 

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2010a. Is Nuphar lutea (L.) Sm. a structuring 

factor for macrozoobenthos and selected 

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species of Hydrobaenus Fries, 1830 (Diptera, 

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37-43. 

Zhang, E., Liu, E., Jones, R., Langdon, P., Yang, 

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recent changes in human activity and 

eutrophication of Lake Wushan from the 

middle reach of the Yangze River, China. - J. 

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Zhang, Y., Dudgeon, D., Cheng, D., Thoe, W., 

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macroinvertebrate communities in the Pearl 

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652: 71-88. 

Živić, I., Marković, Z., Filipović-Rojka, Z. and 

Živić, M. 2010a. Influence of a trout farm on 

water quality and macrozoobenthos 

communities of the receiving stream (Trešnica 

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687.

Chironomus 23 (Nov 2010) - Insect Division

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