Annls Limnol. 23 (1) 1987 : 27-60

The zoogeographical distribution of (Insecta : Diptera)

P. Ashe' D. A. Murray2 F. Reiss3

Keywords : Chironomidae, distribution, zoogeography, ecology, Diptera.

A review of present information on the zoogeography of Chironomidae at subfamily and generic levels is given. The known distribution, including some previously unpublished records, of all recognisable genera (307) and subgenera (56) is tabulated according to zoogeographical region. The greatest number of genera is known from the Nearctic (202) follo­ wed by the Palaearctic (187), Neotropical (109), Australasian (107), Afrotropical (104) and Oriental (100). The chironomid fauna of each zoogeographical region is discussed and a comparison of the number of known species and percentage repre­ sentation of each subfamily for each region is given. An account of the distribution and ecology of each of the ten subfami­ lies is included. To date, some 3 700 species are described representing only 30 % of the estimated world chironomid fauna. The fossil record of the group is briefly discussed and the Chironomidae are estimated to be at least 200 million years old dating back to the Triassic period.

Répartition biogéographique des Chlronomidés (Insecta : Diptera).

Mots clés : Chironomidés, répartition, biogéographie, écologie, Diptera.

Une révision des connaissances actuelles de la biogéographie mondiale des sous-familles, genres et sous-genres de Chi­ ronomidae est présentée. La répartition de 307 genres et 56 sous-genres dans les différentes régions biogéographiques est établie à partir des données de la littérature et de données originales. La région néarctique compte le plus grand nombre de genres (202), suivie par les régions paléarctique (187), néotropi­ cale (109), australienne (107), afrotropicale (104) et orientale (100). Le nombre d'espèces connues dans chaque région, leur proportion relative dans les 10 sous-familles, sont comparées ; leur répartition et leur écologie sont examinées. A ce jour, quelque 3 700 espèces sont décrites ; elles représenteraient seulement 30 % de la faune totale présumée des Chironomidae du monde. Les découvertes fossiles permettent d'estimer la présence des Chironomidés dès la période tria- sique, il y a au moins 200 millions d'années.

1. Introduction immature stages of most species occur in freshwa­ ter but many terrestrial, marine and brackish water The family Chironomidae is a cosmopolitan group species are known. of dipteran representatives of which occur in all zoogeographical regions of the world, inclu­ Most of the early entomologists lived and worked ding . There are few areas where the Chi­ in western Europe and consequently the fauna of ronomidae can be regarded as being absent. The the Palaerctic, in particular the western Palaearc­ tic, was most intensively studied. It is not surpri­ sing therefore that the chironomid fauna of this 1. Department of Zoology, Trinity College, Dublin 2, Ireland. region has been most thoroughly studied, followed 2. Department of Zoology, University College Dublin, Belfield, Dublin 4, Ireland. by the Nearctic, Afrotropical and Australasian 3. Zoologische Staatssammlung, Munchhausenstrasse 21, regions. The fauna of the Neotropical and Oriental D-8000 Munchen 60, Federal Republic of Germany. regions is particularily poorly known.

Article available at http://www.limnology-journal.org or http://dx.doi.org/10.1051/limn/1987002 I* \SHK. IX A. Ml'KKA Y. I . KIISS

Ashe (1983) recognised 355 valid genera within the Neotropical region, the Buchonomyiinae from the ten chironomid subfamilies. Some of those genera, Palaeartic and Oriental, the Aphroteniinae from the though technically valid, are of uncertain status. A Neotropical, Afrotropical and Australasian regions few of the valid genera have since been synonymi- and the from the Palaearctic, Nearc­ zed with other genera and some new genera have tic, Neotropical and Oriental regions. been recently described. The 307 genera treated in this work are recognisable genera (i.e. identifiable There has been no previous attempt to give an using recent taxonomic works) excluding those account of the global distribution of chironomid genera which are of doubtful status or those which genera. In this work the definition of zoogeographi­ colleagues have indicated are unpublished synonyms. cal regions in general follows the classical divisions A total of 56 subgenera are currently recognised. with the main difference being that all continental Represenatives of six subfamilies, , and oceanic islands are assigned to zoogeographical , , , Orthocla- regions (Fig. 1). The proposed boundary of the Antarc­ diinae and , have been recorded from tic region has been extended to include some of the all zoogeographical regions. The remaning four sub­ subantarctic islands. Further comments on the boun­ families appear to have a more restricted distribu­ daries between some zoogeographical regions are tion. The Chilenomyiinae are known from the given later in the text. (3) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 29

2. Chironomid fauna of the zoogeogra­ Table 1 : The number of genera of each subfamily recor­ phical regions ded from the six zoogeographical regions.

The known zoogeographical distribution of all currently recognised genera and subgenera is listed (p. 10-18) and a summary of the subfamily generic representation in the zoogeographical zones is given in Table 1.

2.1. Palaearctic and Nearctic regions Telmatogetoninae 2 2 2 2 2 2 These two regions, which together constitute the Chilenomyiinae 0 0 1 0 0 0 Holarctic, are best considered together because of Tanypodinae 29 36 15 16 17 19 faunal similarity. There are many monographs and Buchonomyiinae 1 0 0 0 1 0 revisions of various genera available, especially on Podonominae 5 5 5 7 2 2 the Palaeartic fauna. Eight of the ten chironomid Aphroceniinae 0 0 2 3 0 1 subfamilies are found in the Holarctic. The two sub­ Diamesinae 11 10 5 3 6 2 families which do not occur are the Chilenomyiinae Prodiamesinae 3 4 2 0 2 0 and Aphroteniiae although fossil representatives of 74 79 34 36 33 31 the latter have been described from northern Chironojiinae 62 66 43 40 37 47 U.S.S.R. The most important taxonomic work on identifying the genera of the region is the collabo­ TOTAL 187 202 109 107 100 104 rative work « Chironomidae of the Holarctic Region » edited by Wiederholm (1983 el seq.). Volu­ mes dealing with the larvae and pupae are publis­ hed and one further volume dealing with adult males expected to occur in the arctic-subarctic regions of is in preparation. the Nearctic. Three of the four genera of Prodiame­ sinae are recorded from the Holarctic but the fourth In the Palaearctic and Nearctic respectively the , Compteromesa, is only known from the S.E. subfamily Tanypodinae is represented by 29 and 36 Nearctic. genera with 26 genera common to both regions (Table 1). Genera which occur in the Nearctic not The subfamily Orthocladiinae is represented yet recorded from the Palaeartic are : Aiotanypus, in the Holarctic by 91 genera of which 74 are Brundiniella, Cantopelopia, Coelotanypus, Fitt- found in the Palaearctic, 79 occur in the Nearc­ kauimyia, Helopelopia, Hudsonimyia, Meropelopia, tic and 62 genera are common to both. Twelve Pentaneura and Radoianypus. Genera currently uni­ genera are currently unique to the Palaearctic que to the Palaearctic are , Pentaneurella region : Boreosmittia, Dratnalia, , and Telmatopelopia. The majority of these genera Lappokiejferiella, Lindebergia, Propsilocerus, may ultimately be found in both regions with the Semiocladius, Stackelbergina, Tavastia, Tokuna- possible exception of Aiotanypus, Anatopynia, Pen­ gayusurika, and Tsudayusurika. taneura and Telmatopelopia. Ten genera are only found in the Nearctic : Baeoc- tenus, Doithrix, Oreadomyia, Platysmitiia, Plhudso- Among the smaller subfamilies the representation nia, Saetheriella, Sublettiella, Tethymyia, Trichochi- of genera in the Palaearctic and Nearctic is very lus and . similar or identical. The Buchonomyiinae (sole genus : Buchonomyia) within the Holarctic is only In the Holarctic the Chironominae are represen­ known from the Palaearctic (Ireland through Middle ted by 74 genera. The Palaearctic and Nearctic con­ Europe to Iran). The same five genera of Podonomi- tain 62 and 66 genera respectively of which 54 are nae occur in both regions. Only one additional genus common to both regions. Twelve Nearctic genera of Diamesinae, Lappodiamesa, occurs in the (, , Caladomyia, , Goeldichi- Palaearctic. Representatives of this genus have been ronomus, Hyporhygma, Nimbocera, Oschia, Skutzia, found in high northern latitudes and it could be Stelechomyia, Sublettea and Xestochtronomus) do 30 P. ASHE, D.A. MURRAY, F. REISS (4) not occur in the Palaearctic and of these Oschia and regarding some of the genera listed in Table 2 and Stelechomyia are unique to the Nearctic. Eight some are so recently described that it is impossible Palaearctic genera (Baeoiendipes, , Kloosta, to predict the distribution of a particular genus. Lithotanytarsus, Neostempellina, Nilodorum, Thie- Very few truly endemic genera in either the nemanniola and ) are not recorded Palaearctic or the Nearctic are to be expected. from the Nearctic and five of these, Fleuria, Kloo- A total of 1285 valid species is known from the sia, Lithotanytarsus, Neostempellina and Thieneman- Palaearctic (Cranston & Ashe, in press) which pos­ niola, are unique to the Palaearctic. sesses the richest described chironomid fauna of any In summary, the close zoogeographical relations­ region. The Nearctic has 978 valid species (Table 3). hip between the Palaearctic and Nearctic regions is Figures for the Nearctic region have been provided reflected in the fact that of the 307 world genera 227 by Cranston (pers. comm.). When the subfamily (74 %) occur in the Holarctic and 162 of these are representation in each region is compared with common to both regions. There are 40 genera in the regard to numbers of species and percentage of the Nearctic which are not known from the Palaearctic total fauna some major differences in the main sub­ while the latter region has 25 genera which are not families are apparent (Table 3). These figures repre­ known from the Nearctic - a combined difference of sent the stage of taxonomic knowledge of the parti­ 65 genera (Table 2). This discrepancy seems to give cular subfamily within the region rather than any support to the idea of major differences between the real major differences in subfamily representation. two regions. However, a more detailed examination The Tanypodinae possess 139 species in the Nearc­ reveals the fact that slightly over half of these genera tic and 116 species in the Palaearctic reflecting the (33) are currently monolypic and the immature sta­ more intensive work done on this subfamily throug­ ges of 42 are recently described or unknown while hout North America in recent years compared with most of the remaining genera contain only 2-6 spe­ more localised studies in the Palaearctic. In the cies (Table 2). Genera with few species are often res­ Orthocladiinae the differences between the two tricted to very specific habitats and thus less likely regions are not very significant but the Chironomi- to be collected in random sampling. More intensive nae with 348 species in the Nearctic is 192 species collecting in the future may redress this situation. poorer than the Palaearctic with 540 species. This Genera currently endemic to the Palaearctic or the anomaly can be partly explained by the fact that the Nearctic only and which occur in high northern lati­ Tanytarsini are very under-rep resented in the Nearc­ tudes can be expected to occur in the other region tic (Table 4). Very few new species of Tanytarsini (e.g. Pentaneurella, Lappodiamesa) ; however, those have been described in North America over the last genera confined to southern latitudes within the two decades pending a revision of the group by Prof. Palaearctic or Nearctic regions are unlikely to occur J.E. Sublette and when complete the imbalance bet­ in the other region unless those genera also show ween the two regions should be less marked. The a much wider distribution beyond adjoining zoogeo­ lower number of in the Nearctic is due graphical regions. Most of the genera which are to the less active study of the group there. The hig­ found in the southern Nearctic (but not in the her number of in the Nearctic Palaearctic) are essentially Neotropical elements reflects the proximity to the Neotropical region which have spread northwards (e.g. Pentaneura, where this group is abundant with at least 43 spe­ Lopescladius, Beardius, Caladomyia, Goeldichirono- cies reported (Fittkau & Reiss 1979). nius, Nimbocera and Xestochironomus) and if they do not occur at least in the Australasian and Orien­ As the present paper was being written we became tal regions then they cannot be expected from the aware that a paper on the problems of Holarctic bio- Palaearctic. It is probable that some essentially geography was at the same time being prepared Oriental elements will be discovered in the southern (Cranston & Oliver, in press). Although different Palaearctic (especially the lowlands of central China) aspects of the Holarctic generic fauna are discus­ similar to the situation in the southern Nearctic sed in varying detail in this paper and in Cranston which receives some Neotropical elements. Insuffi­ & Oliver (in press) it is interesting to note that simi­ cient data on the distribution of individual genera lar or identical conclusions are reached on topics and species makes it difficult to make statements such as monotypic and endemic genera, origin of certain genera, etc. However, the « Antillean Table 2 : Genera within the Holarctic which are currently only known from either the Palaearctic or the Nearctic (num­ bers in parentheses refers to the number of described species ; * = Palaearctic endemic ; + - Nearctic endemic ; $ = immatures unknown or only recently described, since 1980).

PALAEARCTIC TANYPODINAE NEARCTIC

* Anatopynia (1) Aiotanypus (3) *$ Pentaneurella (1) + Brundiniella CD

* Telmatopelopia (1) S Cantopelopia C3) Co«ïota«i/pwa (13)

Fittkauimyia (2) +$ Helopelopia (2) + Hudsonvnyia C2) +S Meropelopia (2)

Pentaneura (4) +S Radotanypue CD BUCHONOMYIINAE

$ Buchonomyia (2)

*$ Lappodiameea (1) PRODIAMESINAE •S C"cmptâï\3mesa CD 0RTHOCLADIINAE

*S Boreosmittia (2) $ /Inttiîc-eJûtJius C3) *S Dratnalia (1) + Baeoatenus (1)

*$ Euryonemus CD $ Compterosmittia (2) *$ Lappokiefferiella (1) $ Diploemittia (2) *$ Lindebeegia C1) +$ Doithrix (5) * Propsilocerue C4) $ Eretmoptera (2)

S Semiocladius (2) $ LipurometrioanemuB (2)

* Stackelbergina (1) $ topescladtwe (5) *$ Tavastia (1) +S Qreadcmyia (1) * Tokunagayusurika CD +5 (2) *$ TrissocladiuB C2) +$ Plhûdscmia (1) *S TsudayuBurika CD +S SaetAeWeZîa CD +S Suiîettieîîa CD + Tethymyia CD +$ Trtetoefciîus CD +$ tfrwteîïa CD

$ (2) CHIRONOMINAE

* (1) Asheum CD * Fleuria (1) S Beardiue (3) *$ Klooeia (1) $ Caladomyia (8) * Lithotanytarsus (D Gillotia (2) *$ Neostempellina CI) GoeldichironcmtB (9) Nilodorum (6) $ Hyporhygma (L) * Thienemanniola (1) Nimbocera C2) $ VirgatanytarauB (7) +$ Oeahia (1) S Skutzia (2) +$ StelecAomyta (1)

Sublettea (1) $ JfeatocftiroMOTTua (12) P. ASHE, DA. MURRAY, F. REISS (6)

Table 3 : The number of described species and the percentage representation of each subfamily in the six zoogeographical regions (figures given for the Palaearctic, Nearctic and Australasian regions, are based on unpublished manuscripts, and may differ slightly from those which appear in forthcoming published catalogues).

PALAEARCTIC NEARCTIC NEOTROPICAL AUSTRALASIAN ORIENTAL AFROTROPICAL

No. X No. % No. % No. % No. % No. %

Telmatogetoninae 5 0.4 7 0.7 10 2.7 16 4.0 3 0.9 3 0.7 Ch i1enomyi inae - - - - 1 0.3 - - - - - Tanypodinae 116 9.0 139 14.2 51 13.8 43 10.7 67 19.2 65 15.3 Buchonomyiinae 1 0.1 - - - - 1 0.3 - Podonominae 13 1.0 13 1.3 83 22.5 33 8.2 3 0.9 3 0.7 Aphroteniinae - - - - 2 0.5 4 1.0 - - 2 0.4 Diamesinae 99 7.7 55 5.6 12 3.3 8 2.0 15 4.3 4 0.9 Prodiamesinae 11 0.9 10 1.0 1 0.3 ------Orthoclàdiinae 500 38.9 406 41.5 86 23.3 109 27.0 78 22.4 Ill 26.1 Chironominae 540 42.0 348 35.6 123 33.3 190 47.1 181 52.0 238 55.9

TOTAL 1285 978 369 403 348 426

Table 4 : Comparison of Chironominae species numbers 2.2. Neotropical region within tribes in the Palaearctic and Nearctic regions. The available monographs on Chironomidae of this region only cover a small part of the total fauna. Palaearctic Nearctic Edwards (1931) deals with the fauna of Patagonia and South Chile and the species described only 261 Chironomini 333 represent a small part of the chironomid fauna of 13 Pseudochironomini 1 this region of South America. However, the generic of Edwards (op. cit.) is outdated. In his Tanytarsini 206 74 excellent monograph on fransantarctic relationships Brundin (1966) deals extensively with the subfami­ TOTAL 540 lies Podonominae, Aphroteniinae and Diamesinae (except ) of this region. Fittkau and Reiss in several publications have dealt with the subfami­ lies Tanypodinae and Chironominae in the Amazon element » as discussed by Cranston & Oliver (in basin but as yet only a small fraction of the known press) appears to be a Neotropical component which species have been described. Reiss ( 1972) has repor­ has migrated northwards (supported by the recent ted on the Tanytarsini fauna of Patagonia and South discovery of Diplosmittia from mainland South Ame­ Chile. The subfamily Orthoclàdiinae is the most rica - see chapter 5) rather than an endemic Nearc­ poorly known although it probably has a rich and tic or Antillean component. varied fauna in the high mountain and cold tempe­ rate regions. The total area of the Palaearctic region (approx. 48 million sq. km.) is over twice that of the Nearc­ A recent catalogue of Chironomidae in the Neo- tic region (approx. 22 million sq. km.) and on the tropics is not available. The number of recognisa­ basis of land area alone the Palaearctic requires ble species presented here was compiled from a twice the effort and resources to obtain the same wide range of published literature and cross­ level of knowledge as in the Nearctic. checked against « Zoological Record ». In this region (7) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 33

109 genera have been reported of which 15 are ende­ Table 5 : The number of known species and the percentage mic and curently there are 369 recognisable spe­ representation of each subfamily in the tropical low­ cies. All subfamilies except Buchonomyiinae have lands of South America (modified from Fittkau & Reiss 1979) compared to the fauna of Andean-Patagonian flo­ been recorded from the region. The subfamily Chi­ wing waters (estimated from Brundin 1966). ronominae is dominant with 123 species (33.3 %) fol­ lowed by the Orthoclàdiinae and Podonominae with 86 (23.3 %) and 83 (22.5 %) species respectively and the Tanypodinae with 51 (13.8 %) species (Table 3). The balance of 24 (7.1 %) species represent the five remaining subfamilies. Telmatogetor Chi.lenomyi.ir Some additional information is available for the Tanypodinae Neotropical region which shows the contrast in the Podonominae chironomid fauna, at the subfamily level, between Aphroteniinj the tropical-subtropical lowlands (below 500 m) and Prodiameaina the temperate mountain fauna of flowing waters in the Andean-Patagonian region (Table 5). Figures for the tropical-subtropical lowlands are from Fittkau & Reiss (1979). However, the single Diamesinae spe­ cies mentioned (op. cit. : 276) in their table belongs to Prodiamesa which is now included in the Prodia- mesinae and was collected as a in the foothills of the Peruvian Andes between 100-200 metres Chironominae would probably constitute about (Roback 1966)- this subfamily cannot be regarded 10-12 %, the Tanypodinae about 6-8 % and the Pro- as typical of the tropical lowlands. It is evident that diamesinae about 1 % or less. The subfamily Chile­ the Chironominae are by far the most important nomyiinae is only recently described (Brundin group with nearly 80 % of the species, followed by 1983a), probably occurs in flowing water, and is the Tanypodinae with 13.1 % and the Orthoclàdii­ included in the table for the sake of completeness. nae with only 8.2 %. Neotropical Telmatogeloninae, Significant, from these figures for the Andean- which are only found on the coast can be expected Patagonian flowing waters, compared with those of on rocky shores throughout the region. The Prodia- the tropical-subtropical lowlands, is the presence of mesinae, which in the Neotropical region may occur the Podonominae, Diamesinae, Aphroteniinae and in peripheral areas bordering on the tropical low­ Chilenomyiinae. In addition the Podonominae and lands, can be regarded as more typical of temperate Orthoclàdiinae, each with about 38 % of the fauna, latitudes and cool high mountain regions. Signifi­ together constitute 75 % of the total fauna and are cantly, the Podonominae, Diamesinae, Aphrotenii­ the dominant groups. nae and Chilenomyiinae have not been reported from the tropical-subtropical lowlands. The figures 2.3. Australasian region for the Andean-Patagonian flowing waters are esti­ mated from Brundin (1966 : 440-441) : « podonomine The principal taxonomic works on this region fauna... makes up no less than 38 % of the chirono­ include those of Freeman (1959, 1961) dealing with mid species of the running waters... Diamesinae + New Zealand and Australia respectively ; Brundin Orthoclàdiinae (42 °/o) ». Brundin's (he. cit.) 38 % (1966) on the representatives of the subfamilies Aph­ Podonominae figure is equivalent to 87 species and roteniinae, Podonominae and Diamesinae in Austra­ by taking account of the described Diamesinae (11 lia and New Zealand ; Glover (1973) on with the species) the percentage and number of Orthoclàdii­ Tanytarsini of Australia. The fauna of New Zea­ nae can be estimated. The figures given here (Table land's subantarctic islands is treated by Sublette & 5) are estimates but are probably reasonably accu­ Wirth (1980). The only recent monograph on the rate. No figures are given for the Tanypodinae, Pro- Micronesian fauna is by Tokunaga (1964) although diamesinae or Chironominae but all three together the taxonomy at the generic level is outdated. The only make up 18.5 % of the fauna. The generic placement of the Australian Tanypodinae 34 P ASHE, DA. MURRAY. F. REISS (8) dealt with by Freeman (1961) has been updated by Table 6 : Genera and one Subgenus unique to the Roback (1982b, c). A catalogue of the fauna of the Andean/Patagonian region and S.E. Australia/New region is in preparation (Cranston, pers. comm.) and Zealand. the previously uncertain generic placement of many species has been resolved • figures given in Table 3 Podoahlua Austroùladiue reflect this information. Podonomopais RhinocladiuR The Chilenomyiinae, Buchonomyiinae and Prodia- Podonomus Stiatocladius mesinae are the only subfamilies not reported from Rheoohlua Symbiocladius (Aaletius) the Australasian region. The fauna of this region Aphroteniella Megaeentron consists of 107 genera, of which 25 are currently Paraphrotenia Riethia endemic, and 403 valid species. The number of spe­ cies and the percentage representation of each sub­ Paraheptagyia family is given in Table 3. The Chironominae (47.1%), Orthoclàdiinae (27.0%), Tanypodinae (10.7 %) and the Podonominae(8.2 %)arethe domi­ of the species described from this region by nant subfamilies with the Telmatogetoninae, Diame­ Kieffer, now regarded as nomina dubia, require sinae and Aphroteniinae making up the remainder. re-examination and re-description in line with The immature stages of most of the endemic genera currently accepted diagnostic and illustrative are unknown as are the immatures of most of the methods. In recent years Chaudhuri and co-workers described species. The strong relationship between have begun working on the taxonomy of the fauna the fauna of S.E. Australia + New Zealand and the of the Indian subcontinent. The dividing line between Chile/Patagonian fauna of the Neotropics is seen in the Oriental and Australasian Region's, as given in the fact that 12 genera and one subenus are unique the « Catalogue of the Diptera of the Oriental to the two areas (Table 6). The discovery of the sub­ Region », is Weber's Line but a detailed study on the family Chilenomyiiae, which so far is only known island faunas in the area is required to determine if from Southern Chile, would not be unexpected in this represents the true boundary. At the boundary S.E. Australia and New Zealand. As our knowledge between the Oriental and Palaearctic regions Reiss of the fauna of these areas increases it is likely that (1971) regards those taxa in the Himalayan range this trend will be more conclusively demonstrated. within the Oriental region occurring above 2,000 The chironomid fauna of New Zealand is generally metres to be essentially of Palaearctic origin whereas regarded as being impoverished and at present con­ he (Reiss op. cit.) considered those occurring below sists of 86 species or 21 % of the total fauna of the this altitude to be Oriental elements. Australasian region of which only four species are reported to occur in both New Zealand and Austra­ lia. According to Brundin (1966 : 449) the fauna of Only two of the ten chironomid subfamilies (Chi­ New Zealand shows closer relationship to that of lenomyiinae and Aphroteniinae) have not been recor­ Chile/Patagonia than to the continent of Australia. ded from the Oriental region. A total of 100 genera are now known from the region and there are, at pre­ sent, only four genera - Neopodonomus, Asclerina, 2.4. Oriental region Eusmittia and Trichotendipes which are unique to this region. The larvae and pupae of all four ende­ The chironomid fauna of the Oriental region is the mic genera are unknown. Neopodonomus and Ascle­ least well known of all the zoogeographical regions. rina are known from high altitude (over 2,000 m) in The adults of many described species are poorly and Bhutan and Nepal respectively and are probably incompletely illustrated and described so that reco­ really Palaearctic elements. The genus Eusmittia is gnition of many described or even new species is only known from the Batu Caves in Malaysia (Free­ very difficult. The immature stages have been lar­ man 1962) and the recently described Trichotendipes gely ignored and are mostly unknown or undescri- occurs on the Andaman Islands, Bay of Bengal (Guha bed. A catalogue of the Chironomidae of the Orien­ & al. 1985). The region possesses a fauna of 348 valid tal region has been published (Sublette & Sublette species (compiled from the catalogue of Sublette & 1973). It is evident from this catalogue that many Sublette 1973, Zoological Record and various (9) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 35

individual taxonomic publications). The figures avai­ Diamesinae. The Diamesinae are poorly represented lable for the number of species and percentage repre­ in the Afrotropical fauna with only four known spe­ sentation of each subfamily {Table 3) shows that the cies of which three belong to Diamesa and one spe­ Chironominae, Orthoclàdiinae and Tanypodinae are cies in the endemic genus Harrisonina. The possible the dominant groups with 181 (52.0 %), 78 (22.4 %) origin of Afrotropical Diamesa is discussed by Wil­ and 67 (19.2 %) species respectively. The balance of lassen & Cranston (1986) who suggest that they are 22 (6.4 %) species represent the remaining five sub­ derived from species that existed in northern latitu­ families. The Diamesinae is well represented, mainly des and that the sister-group relation­ concentrated in the Himalayas, and it is likely that ship of the three Afrotropical species is with species the species inventory of this subfamily and other cold or species-groups from the high mountain ranges in adapted groups will increase with more intensive the Himalayas and the southern Palaearctic. Diamesa work on the high mountain fauna. is not known to occur further south than Mount Kenya and according to Cranston (pers. comm. to P. 2.5. Afrotropical Region Ashe) the genus has not been discovered in any of the extensive samples examined from rivers in the Dra- After the Palaearctic and Nearctic the chironomid kensberg Range in Southern where ideal con­ fauna of the Afrotropical region is the next best ditions for Diamesa exist. The Prodiamesinae and known of all the regions. Our present understanding Buchonomyiinae are unknown from the region, of the fauna of this region is mainly due to the work although their presence cannot be ruled out, and if of Freeman (1955, 1956, 1957, 1958) who resolved they occur the most likely area would be the East most of the problems with Kieffer's inadequate des­ African Highlands. criptions, redescribed all the known species as well as describing a number of new species. However, the generic taxonomy of Freeman's early papers is out­ 2.6. Antarctic region dated and when identifying species the works men­ tioned above should be used in conjunction with the The Antarctic region as defined here includes the recently published Afrotropical catalogue (Freeman land-mass of Antarctica as well as the islands or & Cranston 1980) in which the species are placed into island groups of : South Shetlands, South Orkneys, their modern generic concept. Recently, Lehmann South Georgia, South Sandwich, Bouvet, Prince (1979, 1981) studied the fauna of two streams in Zaire Edward, Crozet, Kerguelen, Heard and Balleny (lie adding more new species to the faunal inventory. Saint Paul and Ile d'Amsterdam are regarded as belonging to the Afrotropical region). Due to the fact Seven subfamilies are known from the region and that so few chironomid species (or other insects) are those which have not been recorded are the Chile­ known or can be expected from the region it nomyiinae, Buchonomyiinae and Prodiamesinae. To seems best not to assign the region a status similar date, 104 genera are known and 18 genera are cur­ to that of the major zoogeographical regions and rently endemic (List, Table 1). The region has a des­ for this reason a separate column for the Antarctic cribed fauna of 426 valid species (compiled from region is not included in the various tables. Freeman & Cranston 1980 ; Lehmann 1979, 1981 ; The known chironomid fauna of the Antarctic region, Willassen & Cranston 1986). The species and percen­ as defined here, consists of nine species (Table 7) and tage representation of each subfamily is given in the total chironomid fauna will probably consist of Table 3. Interestingly the Chironominae, with 238 not more than 20-30 species. Two genera, Microzetia species, represents 55.9 % of the total chironomid and , are endemic to the region. Belgica con­ fauna whereas the Orthoclàdiinae with 111 species tains two described species, B. antarctica which is the and 26.1 % of the fauna is poorly represented when only chironomid known from mainland Antarctica as compared with the Palaearctic or Nearctic faunas. well as being the most southerly known free-living The Tanypodinae with 65 species (15.3 %) has a rela­ holometabolous (Usher & Edwards 1984). The tively rich fauna. The Chironominae, Orthoclàdiinae second species, B. albipes, is only known from the and Tanypodinae combined make up 97.3 % of the Crozet Islands as is the monotypic genus Microzetia. fauna or 414 species. The balance of 2.7 % and ^spe­ cies represents the remaining four subfamilies i.e. The fauna of Antarctica, particularly in the Gra­ Podonominae, Aphroteniinae, Telmatogetoninae and ham Land area, may in reality be merely an 36 P. ASHE, D.A. MURRAY. F. REISS (10)

List of chironomid genera and subgenera (indented) and their zoogeographical distribution (* " worldwide di­ stribution; + " published record; U • previously unpublished record (see Chapter 5); S • refers to a taxonomic note in Chapter 5; numbers in the right hand column are the sources for published records, see references).

TELMAT0GET0NINAE (2) References * • + + * + + 23 * Thalaesomya + * + + + + 23, 31

CHILENOMYIINAE (l)

Chilenomyia * 6

TANYPODINAE (44)

* Ablabeamyia + + + + + + 23 * Ablabeamyia + + + + + + 23 Asayia * * 60 Karelia + + + + • 13, 38, 55, Sartaia + 59 Alotanypua * * 24 Anatopynia + 24 § Apeectrotanypus + + + + 24, 28, 56 Arctopelopia * + 24 Brundinielia + 24 Cantopelopia * ' * 24, 28 Ckrysopelopia + 28 * ClinotanypuB + + • + + + 24 Aponteua * 55 * ClinotanypuB + + +• + + + 24 Coelopynia * 26 Coeîotanypwa + * + * 24 _ * Conchape lopia * * * V * * 24 Devotanypua + + 24 S Djalmabatieta + + + • 24 Fittkauimyia *• + *** 24» 56 Gressittiua + 85 Guttipelopia * * 24 Bayesomyia * + 43 Helopelopia * 24 Hudsonimyia * 24 Krenopelopia + * +24 S Laii-uttdinta + + + • 23 (11) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 37 „ « Id S 'a u o 3 ° c — u4 -S s. Z o iS » Ï < o References * Laraia + + + • + * 24, 28, 56 Lepidopelopia + 28

* Macropelopia • * + • 24 Meropelopia • 24 Monopelopia • + • 24, 28 NaelotanypitB + 58 Nataraia * 24 * Nilotanypue + * U u U • 24 Paramerina • • • 24, 28 Pentaneura + * 24 PentaneureI la + 24 • • * 24 i* Proaladius Holotanypua U V 56 Proaladiua u 24 Peilotanypus * + u * 22, 28, 55, 57 Paeotrotanypua • • 23 RadotanypuB • 21 Rheopelopia + * u 24 * Tanypus + * * • • 24 Apelopia • 55 * Tanypus + + • • + 24 TeImatopelopia + 24 Telopelopia + • 24 Thienemannimyia + • • + 24, 28, 48 Triasopelopia + • + 24, 28 Xenopelopia + • 24

2avrelimyia • * * 24

BUCHONOMYIINAE CD Buohonomyia + 40

PODONOMINAE (14)

Afrochtua • 4 i ArchaeochluB • 4, 20 Boreoohlue • + • 7 Laaiodiameaa + • 7 Microzetia Croi set Islands 77

Neopodonomua 9 Paraboreochlua • • 7 Parochlua • • » • 7 Podochlus + 4 i Podonomopaia + + 4 Podcnomua • 4 RheoohluB + + 4 P. ASHE, D.A. MURRAY. F. REISS

Triahotanypua + • 7 ZtlandochîuB • 4

APHROTENIINAE (4) Anaphrotenia • 8 Aphro tenia • 4 Aphroteniella + + 4 Paraphrotenûz + • 4

DIAMESINAE (18) Ar-ctodiameea * u 39 Boreoheptagyia • + + 45, 79 Diamesa + + + • + 4, 45 fiarrisonina * 4 Heptagyia +- 4 Lappodiameea + 43 Limaya + 4 Lobodiameea • 4 Maoridiameea • 4 Pagaatia * * 45 Paraheptagyia * * 4 ftotttoatta + + • 4, 45 Pvotanypue + + + 4, 45 Paeudodiameea + + • 45 Pachydiameaa * 44 Peeucfocfiameea + + +45 Paeudokiefferiella + + 45 Reieeùi * 5 Sj/mpottfezstùz • + «-45 Syndiameaa * * 45

PRODIAHESINAE (4) Compterimeea + 84 MonodiameBa + + * 0 64 Octoniomesa + + • 4, 64 Prod iameea + + • 23

ORTHOCLADIINAE (118) Abiokomyia + + 18 Acarnptoaladiue • * 18 AcrttTofcopwB + • • 18 AllocladiuB + 28 AllometriocnemuB * 26 Allotriaaocladiua * • 12, 27 (13) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 39

References Ancy loc ladius + 84 Antilloc ladius + + IS Apometriocnemue U + 68 Aeclerina + 49 •Auatrobrillia * 26 Austrooladius * + 26 BaeoatenuB * 18 ; Crozet Islands 78 Boreoamittia + 92 * * + 18 Bryophaenoo ladius • + + + + 18 CamptocXadiua + + + 18 * Cardioclodiua + + + + + + 23 Chae toc ladiu a + + + + 10,18 Chaamatonotus + + 18 * Clunto + + + + + + 23 Comptevomittia + + 18 * + + + +. + + 23, 86 * CriootopUB + + + + + + 18 * CriootopUB + + + + + + 18 laocladiuB • + + + 33, 37, 80, 86 Hauriua + 38 NoatococladiuB * * 1, 80 * * 18 % Diploamittia + • 72 Doithrix * 18 DoloplastUB 26 Doncricotopue + + 18 + 18 £po£eocl(2tiiua + + U 18 Eretmoptera + Antarctica 18 * ++++++ 18 § Euryanemug + 41 ffwyftapaia + + 18 Eusmittia + 83 Freemaniella + 28 GeorthoaladiuB + + 18 Atelopodella + 18 Georthooladius • + 18 G^rnnometr£c'C7i«zn7KB • + + +18 Gymnornetriocnemus + + • +18 Baphidooladius + + 65 Gynnidoo ladius + 85 Halooladius * * 18 P. ASHE, D.A. MURRAY, F. REISS

Haloclaâiua + + 18 PaarmooladiuB + 1g + + +18 HeterotanytareuB + + 18 HeterotriBBOcladiue • • + 18, 83 Hevelius + 85 Hydcobaenua + + 18 + 23 Jrïeofcrt'llta + 48 Kiefferophyee « 2* fciepperra • 28 Kreiosmittia + + D • 18, 28 KuBchbliue • 85 Lappokiefferiella + 92 LappoBmittia * • 15, 22 + • + + + + 16 Lindebergia + 89 Ct'pur orne i riocnemu e + + 18 Lopesaladiua + + 18 Cordiella + + 16 Lopeoaladiua + + 67 Mzryella + 85 Mecaorue * 65 Mesocricotopua + + 18 MesosmtÉtia + + + + 18, 37, 71 Metrt'ocMemua + + + + + + 18 Nakataia + 85 /toi0cZac?tus ++ + + + + 18, 86, 88 NanocladiuB + + + + + + 18, 86, 88 PleaopteracoluthuB + 18 Naeuticladiue * • 26, 81 Nesiocladiua • . " 85 0£tuerï<2i<3 + + 18 Oreadomyia + 18 QrthocladiuB + + + + + + 18 EudactyloaladiuB + + + U 19, 22, 83 EuorthocladiuB + + 22, 63 OrthooladiuB + + + + + + 18 Pogonocladiue + + 19, 22 ParachaetocladiuB + + 18 Raraoïaiiue • + 18 Paracriaotopus + + U 18 + + + + + 3, 18 ParalimnophyeB * V 18 THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE

* Parametriocnemua Faraphaenoclaâiue 18, 61 Parasmittia * Paratriahoaladius 18, 34 ParatrisBocladius 18 18 Fetalocladius 84 Platyamittia 18 Plhudaonia PropsiloaeruB FseatroaladiuB AIlopaeotrootadiuB Mesopsectrocladius Monopeectrocladiua Fsectrocladius Poeudorthocladius Lordella

Pseudorthoc ladius Faeudoamittia 18, 26, 28, 61 pailometriocnemus 18, 91 PteroBis §* RheocricotopuB PsilocriaotopuB Rhinocladius Saeth^iella SsmioaladiuB * StackeIbergina I StictoaladiuB Stilocladius Sublettiella 65 Symbiocladiua 15, 22, 35, 54 Acletius 35, 54 SymbiocladiuB 15, 22 Symposiocladius 18 SynorthocladiuB Tavastia Tethymyia ThalaBBOsmittia Thicnemannia * Thienenanniella 42 P. ASHE DA MURRAY. F. REISS (16)

References

Tokunagayusurika • 18 Trichoahitua + 70 TrieeooladiuB • 18 Taudayuaurika • 76 • + 18 Umiella • 16 Xylotopua • + IS Zaluteohia * * IS

CHIRONOMINAE (101) Aoaicarella Aedokritue «23 ABheum • • 47, 53 Axarua • 47 Baeotendipea • + 47 Beardiua • S3 Beokidia • 47 Caladomyia * • 73 CanpoBimyia 75 Chernovakiia • • U 47 * ChironomuB * * • * * 47 CamptochironomuB • * • 22, 36 Chaetolabia 93 * ChironomuB • • • • • 47 LoboohironomuB • 62 * * • • • * • 23, 87 * cladotanytaraua • • • 75 Collartomyia 28 ConoohironotmB • + 26 • • 47 Corynoaera • * + 47 * Cryptoohironoima • * * • • 47 i Cryptotendipea • u u 47 • • • 47 • • 47 DemicryptochironomuB * • 37, 47 * Dicrotendipea • • • * * 47 • * 32, 47 Endochironomua • • * * • 47, 88 Fleuria 47 Gillotia + • 47, 50 Glyptotendipee • • • • 3, 47 i GoeIdichironomua + • 47. 53 Sraoeua • • 47 THE ZOOGEOGRAPHICAL DISTRIBUTION" OF CHIRONOMIDAE

References Barniachia + + + + + 47 Harrisiua + 26 Henrordia + 28 Hyporhygma • + 47,53 Imparipecten + 26 Kiefferulua • + + + + 11, 47 Kiefferulua + + + + + 11, 47 Wirthiello. * 15 Klooeia + 94 Krenopaectra + + 47 Kribioooemua + 28 Kribiodoaie + 28 Kribiomyia + 28 Kribiothauma + 28 Kribioxenua + 28 £

Paryiterffum • 26 Pauciapinigera + 25 P. ASHE, DA. MURRAY, F. REISS

phaenopaeetra t • • • 3, 38, Po lypedi tun* • 47 Pentapedilum • • » 22. 26, Polypedilwn • 47 Tripodura » • u • 37, 42, U * • • 14, 47 PaeudoohironormB • * • 47 Fsiloahironomua » Rheotanytaraua 47 Riethia * • 26 • • 0 47, 50 • 22. 66 47 BaicaloBergentia * 47 Sergentia • • • 47 Skueella • • 26, 28 Skutzia + 52 Stelechomyia * 47 Stensjellina * * • • * 47 Stempel t-inella * u 75 StenochironomUB * • 47 Petalopholeua • • * 2 Stenochironorma * • • » 47 Stictoahironorma * * • • • 26, 47 Sublettea • • • 47, 50 Tanytaraua • • • • 47 Thienemannicla • 47 Tenjnaia 23 Tribeloa + • 47 Trichotendipea • 29 Virgatanytaraua * 47 Xenoehironorme • * • * + + 23 XeatoahironomuB * • M Yama • " « • • + 47 • 47 (19) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 45

Table 7 : Known Chironomidae of the Anlarctic region and iheir distribution.

Species Distribution

Telmatogeton amphibius (Eaton) Kerguelen Is. Crozet Is. Microzetia mirabilis Seguy Crozet Is. Parochlus steinenii (Gercke) South Sandwich Is., South Georgia Is. Parochlus n.sp. Edwards & Usher 1985 South Georgia Is. Belgica albipes (Seguy) Crozet Is. Belgica antarctica- Jacobs offshore islands and mainland Antarctica Eretmoptera murphyi Schaeffer South Georgia Is., South Orkney Is. Limnophyes minimus (Meigen) Kerguelen Is. Limnophyes sp. Brundin 1970 South Georgia Is.

extention of the Patagonian fauna. More work is requi­ 3.2. Chilenomyiinae red in south Patagonia (Tierra del Fuego) to investi­ gate the possible occurrence of Antarctic elements This is the most rçcently recognised subfamily as which may contradict recognition of a true Antarctic well as one of the most primitive. The Chilenomyii­ region. P. steinenii from South Georgia is not conspe- nae is the smallest subfamily with only a single spe­ cific with the Patagonian « steinenii » of Brundin cies, i.e. Chilenomyia paradoxa Brundin. The subfa­ (1966) and the latter represents an undescribed mily is exclusively Neotropical and has only been species. found to date in Southern Chile (Brundin 1983a). Seve­ ral genera in other subfamilies e.g. Podonominae and Aphroteniinae, exhibit a circumantarctic distribution 3. Subfamily ecology and zoogeography with sister species and/or genera occurring in the sou­ thern Neotropics, Australia-New Zealand and southern 3.1. Telmatogetoninae Africa (Brundin 1966). It would not therefore be very The Telmatogetoninae is the sixth largest subfamily surprising if the Chilenomyiinae were discovered in in terms of described species and now contains only New Zealand or S.E. Australia. two valid genera, Telmatogeton and , both of which have a worldwide distribution. Little is known regarding the ecology of this subfa­ The subfamily is almost exclusively marine, the only mily since the larvae and pupae have yet to be disco­ freshwater representatives known belong to the genus vered. The adults were found flying near small brooks Telmatogeton where five species inhabit rapid moun­ in dense Notofagus forests (Brundin 1983a). The imma­ tain streams in the Hawaiian Islands. The immature ture stages, as in the majority of species in the family, stages of the marine species are found in the inter- will probably prove to be aquatic. tidal zone on rocky coasts where the larvae feed on 3.3. Tanypodinae algae and algal detritus. Larvae of the marine species construct tubes in various algae including Enteromor- This subfamily is the third largest in terms of the pha, Monostroma, Porphyra and Viva (Tokunaga 1935). number of genera and species. Currently there are 44 Recently, Robles (1984) has shown that the heavily chi- recognisable genera with subgenera in Ablabesmyia, tinized anterior end and the peculiar developement of Clinotanypus, Procladius and Tanypus. Only eight the terminal segments in the pupa protect it from evic­ genera, Ablabesmyia, Clinotanypus, Conchapelopia, tion from its tube by aggressive larvae of the same, Larsia, Macropelopia, Nilotanypus, Procladius and or allied, species. Tanypus, are known from all zoogeographical regions 46 P. ASHE, DA. MURRAY, F. REISS (20)

(List). The information currently available indicates 3.4. Buchonomyiinae that there are several other genera which could be expected to have a worldwide distribution : these This subfamily contains a single genus, Bucho- include Apsectrotanypus, Coelotanypus, Fitt- nomyia, and two described species, B. thienemanni kauimyia, , Paramerina, Psectrotanypus Fittkau from the western Palaearctic and B. burma- and Thienemannimyia. On present evidence the nica Brundin & Saether from the Oriental region. Nearctic region is the richest in genera with 36 B. thienemanni occurs in western and central recorded, followed by the Palaearctic with 29, the Europe in a band stretching from Ireland through Afrotropical with 19, the Oriental with 17, the Aus­ southern Britain, France and Germany apparently tralasian with 16 and the Neotropical with 15 as far as Iran (pupal exuvia from Iran probably genera. The Neotropical and Oriental regions, which belong to B. thienemanni) (Murray & Ashe 1981). theoretically should have rich generic faunas, are This species, therefore, could be expected to occur among the poorest which reflects the lack of work in Italy, Yugoslavia, Hungary, Greece, Bulgaria, Tur­ done on the chironomid fauna in these areas when key and Iraq. B. bumxanica is only known from Kam- compared to the Palaearctic and Nearctic. Although baiti in Northern Burma close to the border with the numbers of described lanypodine genera recor­ China. In Iran (Dowling pers. comm.) and Burma ded from the Neotropical and Oriental regions is (Brundin + Saether \978) Buchonomyia has been quite low some undescribed genera have been dis­ found at altitudes over 2,000 m so that the genus covered in these regions. In addition, the discovery could be expected to occur at similar altitudes in of genera already known from other regions is to Pakistan, India and China (Yunnan Province). Tht be expected. genus has not yet been discovered in the Easten Palaearctic or the Nearctic region. The descriptio . The Tanypodinae are generally adapted to warmer of the distinctive pupa (Murray & Ashe 1981), 1st in - waters (middle to higher range of temperature) and tar larva (Ashe 1985) and 4th instar lanva (Ashe, n mostly prefer standing water. Representatives of prep.) will ensure recognition of the immature s';i- some genera (e.g. Tanypus)ha\c been reported from ges. The subfamily Buchonomyiinae in the northt m hot springs at temperatures up to 44.5° C (Thiene- hemisphere seems to have a more southern dis 'ri- mann 1954 : 567, sub Protenthes). When they occur but ion with all reported occurrences betw :en in rivers they are usually more common in the slow 25° N-53° N. flowing lower course or in marginal lentic areas in upland or mountain regions where the current is The larvae and pupae are only known for B. : Hie- weaker. Some genera, such as Rheopelopia and Nilo- nemanni and are found in rivers in areas of mode­ ianypus, are secondarily adapted to running waters rate currents. It is likely that B. burmanica also and may be found in moderately strong currents. occurs in lotie habitats. The subfamily seems o be Some genera which possess haemoglobin are able adapted to the middle range of water temperature to survive in warm stagnant water bodies with typi­ and in Western Europe B. thienemanni emerges cally low oxygen concentrations. Representatives of mainly during the summer months. Recently a sin­ the subfamily are commoner in the warmer clima­ gle 4th instar larva has been discovered which was tic regions, especially in tropical and subtropical obtained from a probable trichopteran case, indica­ areas. The diversity decreases towards colder ting that there may be an association betweer regions or with increasing altitude. The larvae are Buchonomyia and a trichopteran species. generally regarded as being carnivorous but some species are known to feed on diatoms and detritus 3.5. Podonominae (Oliver 1971). The true situation in many species is There are 14 genera currently recognised in thu probably a combination of both types of feeding Podonominae (List). None of these genera are world with reliance on diatoms and detritus when prey wide in distribution but every zoogeographical items are scarce. The larvae of the majority of spe­ region is represented by at least two genera. Pa roc h cies are free-living and none are known to produce lus, which occurs in four regions, i.e. Palearctic. larval or pupal cases. However, one species, Abla Nearctic, Neotropical and Australasian regions, U besmyia (A.) janta Roback is know to be symbiotic the most widely distributed genus. The genus con on Mollusca (Unionidae : Quadrula) (Roback 1982). tains 47 described species but only one, P. kief fen (21) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 47 is found in the northern hemisphere where it has much wider distribution. It is possible that mem­ a Holarctic distribution. The remaining species bers of the subfamily still survive in the Palaearc­ occur only in the southern hemisphere. Boreochlus tic in an area climatically similar to those areas in is the next most widely occuring genus and it has the southern hemisphere where the only known been found in the Holarctic and the Oriental regions. living species occur. Four recent genera, Anaphro- The Palaearctic and Nearctic share five genera in tenia, Aphrotenia, Aphroteniella and Paraphrotenia common i.e. Boreochlus, Lasiodiamesa, Paraboreoch- are recognised. In the Neotropical and Australasian lus, Parochlus and Trichotanypus. Five genera are regions both Aphroteniella and Paraphrotenia occur found in both the Neotropical and Australasian whereas the recently described Anaphrotenia has regions : Parochlus, Podochlus, Podonomopsis, Podo- only been recorded from the latter region. The genus nomus and Rheochlus. The Australasian region con­ Aphrotenia is apparently endemic to Southern tains two further genera, Zelandochlus which is Africa. endemic to New Zealand and Archaeochlus which has been found in Southern Africa and Western Aus­ Aphroteniine larvae are found in mountain tralia. The Oriental region contains two genera, streams in the southern temperate zone (Southern Boreochlus (with two species from Northern Burma) Chile, S.E. Australia and Cape Province in South and Neopodonomus from Bhutan. The genus Micro­ Africa) except for the recently described genus and zetia is only known from the isolated Crozet Islands species, Anaphrotenia lacustris Brundin which is between the African continent and Antarctica. found in the littoral zone of a subtropical lake in Queensland, Australia. Larvae and pupae of the The Podonominae, in general, can be regarded as rheophilic species occur in the algal layer on stones an essentially cold-tolerant predominantly rheophi- exposed to the current in water temperatures ran­ lic group requiring high oxygen concentrations. Lar­ ging from 8.8-20.4° C and seem to be adapted to the vae may be found in waters with a temperature middle range of temperature. They do not show the range 0-24° C (Brundin 1966 : 96) and are adapted tolerance to very low water temperatures exhibited to the lower to middle range of temperature. Most by the Podonominae and are not found in streams species occur in cool rivers and streams but some coming directly from glaciers. The only known len- are found in warm, more or less, temporary streams tic species, A. lacustris, occurs in the sandy littoral (Archaeochlus and Afrochlus), among mosses in cool zone of a warm (25° C), humic, low lying, subtropi­ springs and creeks (Parochlus kiefferi (Garrett), cal lake on Frazer Island, Queensland. The larvae Boreochlus and Paraboreochlus), meltwater tarns at of most species appear to be algal and organic detri­ high latitude (Parochlus steinenii (Gercke) and Tri­ tus feeders except Paraphrotenia cf excellens Brun­ chotanypus) or in standing bog waters (Lasiodia­ din which is carnivorous and feeds on other ar­ mesa) (Brundin 1966). Larvae of some species are thropods such as chironomid larvae and crustaceans found in the most extreme of all freshwater habi­ (Brundin 1966, 1983c). tats i.e. in glacier fed rivers and streams close to the glacier in very powerful currents, in very tow tem­ 3.7. Diamesinae peratures (0-5° C) and in waters containing large amounts of suspended abrasive material (coarse After the Tanypodinae the subfamily Diamesinae sand and gravel) (Brundin 1966: 97). As far as is is the next largest in both numbers of genera and known the larvae feed on unicellular algae, mainly species. None of the genera are worldwide in dis­ diatoms, and algal detritus (Brundin 1966) ; no car­ tribution but Diamesa is the most widely distribu­ nivorous forms have been reported. ted. Diamesa is a good example of a cold adapted Holarctic genus which is typical of cool mountain 3.6. Aphroteniinae streams and lakes, a habitat which is only found in tropical latitudes on very high mountains. In the The subfamily Aphroteniinae is confined in its dis­ Neotropical region Diamesa has been found from tribution to the southern hemisphere. However, the the Mexican Highlands (Serra-Tosio 1977), the Boli­ presence of the fossil amber genus Electrotenia in vian Andes and as far south as Patagonia (Brundin Asiatic Russia (Kalugina 1980) within the Palaearc­ 1966 : 366) to about latitude 55° S. In the Afrotropi­ tic indicates that the subfamily historically had a cal region the most southerly known occurrences 48 P. ASHE, DA. MURRAY, F. REISS (22) are Mt. Kenya and Mt. Ruwenzori on the equator in Afrotropical and Australasian regions it will pro­ the East African Highlands (Willassen & Cranston bably only be discovered by intensive collecting in 1986) and the genus has not been found in Southern the preferred habitat. All four genera i.e. Compte- Africa where suitable habitats exist. Diamesa is not romesa, Monodiamesa, Odontomesa and Prodia­ known from the Australasian region but the nearest mesa, occur in the Nearctic, three in the Palaearc­ occurrence is Mount Kinabalu in Borneo (Willassen, tic (except Compteromesa), two (Monodiamesa and pers. comm. to P. Ashe) at about latitude 6° N and Prodiamesa) from the Neotropical region and two it seems unlikely that the genus could occur in the (Monodiamesa and Odontomesa) from the Oriental New Guinea Highlands or further south. region.

The Diamesinae are a cold adapted group which Larvae of the « Prodiamesa group » (Brundin occur in rivers and lakes and, like the Podonominae, 1966 : 367) [= Prodiamesinae] are detrivores and live are adapted to the low to middle range of tempera­ preferably in cool, slow-flowing streams, in the pro­ ture (0-22° C) and require high oxygen concentra­ funda! zone of oligotrophic lakes and in the littoral tions. Rheophilic species are more common in zone of high mountain lakes. Members of the sub­ upland or mountainous areas being particularly family prefer the low to middle range of water tem­ common wherever the current is strongest although perature and no species have been reported from some genera such as Lohodiamesa and Potthastia warm tropical waters. Prodiamesa olivacea (Meigen) prefer slower current. The rheophilic forms include is reported to be predatory (Beck 1977). Boreoheptagyia whose larvae cling to rocks at the splash line in cool mountain or glacier-fed streams 3.9. Orthoclàdiinae (Oliver 1983). Diamesa is the most commonly encountered lotie genus but it is also known from Currently the Orthoclàdiinae is the largest sub­ springs, shallow still waters (Oliver 1983), meltwa- family with 118 recognisable genera and 31 subge­ ter trickles under snow at high altitude (Kohshima nera. Fourteen genera : , , Cory- 1984) and even exposed wave-washed lake shores noneura, , Eukiefferiella, Limnophyes, (Aagaard 1978). Other genera, e.g. Potthastia and , , , Parame- Pseudodiamesa, also occur in both standing and flo­ triocnemus, , Rheocricoiopus, wing water. Protanypus is the only genus known to Smittia and Thienemannieila are known from all be restricted to standing waters and it is usually regions. In addition, there are nine genera (i.e. Bryo- found in oligotrophic lakes (Oliver 1983) although phaenocladius, , , Brundin (1966 : 366) states that three undescribed Parakiefferiella, , , species from Northern Burma may be rheophilic. , Symbiocladius and Tvetenia) which, based on their present distribution, are likely to The larvae of most species are algal feeders have a worldwide distribution. The Nearctic region (mainly diatoms), Pseudodiamesa is regarded as at present has the richest generic fauna with 79 omnivorous and at least two genera, Lohodiamesa genera followed by the Palaeartic with 74, the Aus­ and Protanypus, are carnivorous (Brundin 1966: tralasian with 36 and the Neotropical, Oriental and 364-366). Afrotropical regions with 34, 33 and 31 genera respectively. 3.8. Prodiamesinae The Orthoclàdiinae can be regarded as the most This subfamily is the seventh most speciose with broadly adapted group ecologically. The subfamily four genera currently recognised. No genera of Pro­ includes numerous species in various types of fresh­ diamesinae are known to have a worldwide distri­ water habitats as well as many terrestrial forms and bution but Monodiamesa, recorded from four some marine genera. In general they are cold adap­ regions, is the most widely distributed. Both Mono­ ted, become more common towards higher latitu­ diamesa and Prodiamesa are found in the Palaearc­ des or with increasing altitude and are poorly repre­ tic and Nearctic and south to Patagonia in the Neo- sented in tropical standing waters. The majority of tropics (Brundin 1966 : 366). In the regions where species prefer the low to middle temperature range the subfamily is found it is usually represented by (0-25° C) but some genera (e.g. Cricotopus) are com­ only a few species. If the subfamily exists in the mon in warm tropical waters and even in hot (23) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 49

springs. Some freshwater genera may be confined 1979 ; Saether & Halvorsen 1981) and Eurycnemus to running water (Eurycnemus, Rheosmitlia) or stan­ was found in trichopteran cases (Hydropsychidae : ding water (Mesocricotopus, Tokunagayusurika) Hydropsyche) although the type of association is whereas others may occur in rivers, lakes, ponds unknown (Murray & Ashe 1981). Cardiocladius lar­ and puddles (, Hydrobaenus). vae are reported to be ectoparasitic and predatory Representatives of some genera are found in very on trichopteran pupae (Parker & Voshell 1979) and restricted habitats such as seeps (), on Diptera (Simulidae) (Steffan 1968). Nanocladius plant held water or phytotelmata (Metriocnemus in subg. Plecopteracoluthus is phoretic on Plecoptera pitcher plants, rot holes, etc.) and temporary pools (Perlidae : Acroneuria and Paragnetina) and both (Lapposmittia). Other genera occur not only in fresh­ subgenera of Nanocladius have been found in pho­ water but also in brackish water (Propsilocerus) or retic association on ( : have representatives that occur in aquatic, semi- Chauliodes, Corydalus and Nigronia) (Gotceitas & terrestrial and terrestrial habitats (Bryophaenocla- Mackay 1980) although Nanocladius s.str. are more dius, Limnophyes) (Cranston et al. 1983). The terres­ typically free-living. A species, placed originally in trial forms include Belgica and Eretmoptera which Dactylocladius (Tonnoir 1922), whose generic affi­ are found in peaty soils among mosses and higher nities are uncertain, is phoretic on Diptera (Blepha- plants (Usher et Edwards 1984, Cranston 1985), riceridae : Neocurupira). A probable Eukiefferiella living in cow dung and Gymnome- species is phoretic on aquatic Hemiptera (Naucori- triocnemus and Parasmittia in the soil of meadows dae : Cryphocricos) (Roback 1977). In lamellibran- and woodlands (Cranston et ai 1983). Marine genera chiate molluscs two orthoclad genera, Baeoctenus include Clunio, which occurs in the littoral to sub- and an undescribed genus, are found among the gills littoral zone (to about 20 m), as well as , of Anodonta species and larvae of Baeoctenus are Tethymyia and (Neumann 1976). known to feed on the gills of the molluscs (Gordon Most aquatic orthoclad genera produce larval/pupal et al. 1978). The only known association between a cases constructed from one or more of a variety of chironomid and a vertebrate was described by Fitt­ materials including detritus, sand grains, salivary kau (1974) where species of the genus Ichthyocladius secretions, diatoms, plant debris. Such cases may are found in cases attached to opercular spines in be fixed to solid objects or covered by sediments. of the families Austroblepidae and Lorica- Larvae of some genera, and Heterotany- riidae although the exact type of relationship in tarsus, produce transportable cases similar to those unknown. constructed by some Chironominae. Within major habitats, e.g. lakes or rivers, some taxa may be seve­ It is obvious that the Orthoclàdiinae display a rely restricted in where they occur : Symposiocla- wide variety of food preferences which include detri­ dius and Xytotopus mine in saturated dead wood tus, diatoms, other algae, living tissues of higher (Cranston et al. 1983), Cricotopus subg. Nostococla- plants, saturated dead wood, dung, etc. and some dius live within colonies of the blue-green alga Nos- are predatory on various invertebrates. Some genera toc (Ashe & Murray 1980), or they may be associa­ feed on only one type of food whereas others are less ted with other . specific and may feed on a wide range of available food types. The immature stages of several orthoclad genera and species are associated (i.e. parasitic, symbiotic, 3.10. Chironominae phoretic, commensal or inquiline) with other aqua­ tic invertebrates and even aquatic vertebrates. Epoi- The Chironominae is the second largest subfamily cocladius is phoretic on ephemeropteran nymphs (List) in terms of genera (101) but the largest if the (Ephemera and Hexagenia) (Steffan 1968) whereas number of described species is taken into account Symbiocladius is ectoparasitic on ephemeropteran (Table 3). Subgenera are recognised in , nymphs (Atalophlebioides, Ecdyonurus, Epeorus, , , Sergentia and Stenochiro-

Habroleptoides, Heptagenia, Meridialarisr Rhithro- nomus. The Chironominae are the dominant group gena and Thraulodes) (Wiens et al. 1975 ; Hynes in the tropical and subtropical lowlands and with 1976 ; Soldan 1978). Dratnalia is phoretic on Tri- more work on the taxonomy in these regions the choptera (Limnephilidae : Potamophylax) (Dratnal number of genera and species can be expected to 50 P. ASHE, DA. MURRAY, F. REISS (24)

increase to make this the dominant subfamily. Four­ 1982) and the genus has five described teen genera have a worldwide distribution : Chiro- marine species from the Pacific area (Hashimoto nomus, Cladopelma, , Cryptochiro- 1976). Some genera and species are very restricted nomus, , , Parachirono- in their occurrence within major habitats (lake, •mus, Polypedilum, , , Ste- river, etc) : some mine in and feed on saturated dead nochironomus, Tanytarsus, and wood (Harrisius, Stelechomyia, Zavreliella. There are over 20 Chironominae genera s.str.) or in living tissues, leaves and stems, of known from five of the six zoogeographical regions macrophytes ( (some species), and it is likely that most of these will prove to have Hyporhygma, Stenochironomus sub. gen. Petalopho- a worldwide distribution. The Nearctic is the richest leus)(Pinder & Reiss 1983, Borkent 1984)ormaybe in genera followed by the Palaearctic, Afrotropical, associated with other invertebrates. Neotropical, Australasian and Oriental regions. Most of the reported cases of Chironominae living The Chironominae are adapted to a wide variety in association with other aquatic invertebrates of freshwater habitats and also occur in brackish involve molluscs. species are and marine waters. At least one terrestrial species, known to be parasitic in Gastropoda with Parachi­ belonging to the genus Tanytarsus, is known to exist ronomus varus Gcetghebuer ectoparasitic in Physa (Pinder & Reiss 1983). As a general rule the number fontinalis Linnaeus and P. varus var. limnaei Guibé of species of Chironominae increases towards the reported as endoparasitic in Limnaea limosa (Guibé equatorial regions, with decreasing altitude and 1942), Lamellibranch molluscs are also known to be with reducing current speed in flowing waters. They parasitized by chironomid larvae. (?) are the dominant group in lowland standing waters paripes Edwards is reported in Anodonta cygnea Lin­ in tropical and subtropical areas of the world. The naeus (Beedham 1966) and Xenochironomus canter- majority of species prefer the middle to high range buryensis (Freeman) from New Zealand is found in of temperature (15-35° C) but some species exist in Hyridella menziensis (Gray) (Forsyth & McCallum hot springs or in streams fed by hot springs at tem­ 1978). Demeijerea species mine in and feed on fresh­ peratures up to 50° C. Many species possess haemo­ water sponges and Bryozoa (Neff & Benfield 1970) globin, a factor which may partly account for their and some species of Xenochironomus mine in spon­ dominance in lowland tropical standing waters ges (Pinder & Reiss 1983). Polypedilum jallax (Joh.) where oxygen concentrations are likely to be low. is reported to be parasitic and predatory on trichop­ Although the ecology of the Chironominae is still teran pupae of the genus Potamophylax (Otto & poorly known because the tropical forms are little Svensson 1981) and Collartomyia also appears to investigated they do not seem to be as broadly adap­ behave similarly (Borkent 1984 : 22 sub Collartiella). ted ecologically as the Orthoclàdiinae - especially Dicrotendipes peringueyunus Kieffer in tropical with regard to marine and terrestrial forms. Most Africa is phoretic on freshwater crabs (Potamonau- genera are represented in standing waters and when tes) and was never found free-living (Disney 1975). they occur in flowing waters it is usually in the The Chironominae, being one of the largest sub­ lower reaches of large, slow flowing rivers or in mar­ families, displays a wide range of food preferences ginal lentic areas in fast flowing rivers and streams, and as indicated above includes those that feed on Some genera are only known from running waters aquatic plants (diatoms, other algae, higher plants, (Rheotanytarsus, Subiettea) or standing waters (Cory- etc.), saturated dead wood, dead leaves, enriched nocera, , Nilodorum) whereas sediments, etc. Predatory forms occur sporadically other genera occur in a wide range of aquatic habi­ throughout the Chironominae but members of the tats including rivers, lakes, thermal springs and Hamischia - complex are generally regarded as brackish water (Cladotanytarsus, Stempellina). Some being carnivorous. genera appear to be restricted to certain types of water bodies : in dystrophic lakes and pools, Lithotanytarsus in limestone rich streams and Baeo- 4. Fossil record and the age of tendipes in polyhaline waters (Pinder & Reiss 1983). Chironomidae The only exclusively marine genus, Pontomyia, is found near coasts to a depth of 30 m (Bretschko A neglected but very important element for the zoogeography, phylogeny and ageing of chironomid (25) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 51 taxa is the fossil component, primarily fossils in and have a history which goes back far into the amber. The amber chironomid fossils have not been Jurassic (Brundin 1966 : 439). Since the adult stage comprehensively dealt with even though this family is not dependant on any specialised food source it of insects is one of the commonest components. A follows that the evolution of chironomids was not study of the amber fossils would be particularly use­ dependant on the presence of vertebrates or higher ful in elucidating past distributions of genera espe­ plants as are some other families of Diptera. The cially in areas where such genera may now be phylogenetic relationships between the chironomid absent. Additionally such a study would help in esta­ subfamilies have been well documented. Recently blishing a minimum geological age for many genera, the phylogenetic position of the Buchonomyiinae tribes and subfamilies. Fossil ambers are known has been resolved with its placement as the sister from several periods of the Cenozoic and Mesozoic group of the Podonominae + Aphroteniinae (Mur­ eras. From the collection of Baltic amber in the Zoo­ ray & Ashe 1985). However, the phylogenetic posi­ logical Museum, Copenhagen (Larsson 1965) it is evi­ tion of the Telmatogetoninae is still uncertain. The dent that the family Chironomidae is by far the lar­ Telmatogetoninae were originally considered as the gest component represented by 671 specimens apomorph sister group of the Diamesinae by Brun­ which is 27.4 % of the Diptera or 13.2 % of all arth­ din (196*6 : 373) a view which was initially suppor­ ropod inclusions. Around the turn of the century ted by Saether (1976) but later rejected and placed Meunier, in various papers, described numerous chi- as the plesiomorphic sister group of the other chi­ ronomids from Baltic and Zanzibarian amber but ronomid subfamilies (Saether 1977). The ecology of all the descriptions are inadequate and the material the Telmatogetoninae (predominantly marine) does needs to be redescribed and compared with existant not agree with the presumed primitive habitat of the species. Only three amber chironomid fossils have ancestral chironomid in the « upper course of small, been adequately described : Libanochlites neocomi- cool mountain streams and springs » suggested by cus Brundin (Podonominae : Boreochlini) from the Brundin and with its present phylogenetic position Lower Cretaceous of Lebanon (120-140 million years as the sister group of all the other chironomid sub­ old) (Brundin 1976) ; Electrotenia bnmdtni Kalugina families. It seems that the Chilenomyiinae should (Aphroteniinae : Electroteniini) from the Upper Cre­ more logically occupy this position with the Telma­ taceous of Siberia (Kalugina 1980) and Cretodiamesa togetoninae placed as a sister group of the Diame­ taimyrica Kalugina (Diamesinae : Cretodiamesini) sinae. There is little evidence to suggest that Gs IX also from the Upper Cretaceous of Siberia (Kalugina of Telmatogetoninae (Saether 1977 : 4) is identical 1976). The subfamilies Tanypodinae, Orthoclàdiinae, with structures found in Sciaridae and « possibl) Chironominae (Boesel 1937) and Buchonomyiinae in some » Nymphomyiidae or that the presence ol (probably B. thienemanni) (Brundin 1966 : 80) are a gonostylus in the adult female is a plesiomorphic known from Baltic amber (Upper Eocene, about 40 feature throughout the Diptera since such structu­ million years old) but the only subfamily reported res are not reported elsewhere in the whole of the from Canadian amber (Upper Cretaceous, about 75 Diptera. Two plesiomorphic characters ; gonostylus million years old) is the Orthoclàdiinae (Boesel 1937). IX (Gs IX) present and gonapophysis VIII (Gp VIII) Only the subfamilies Chilenomyiinae, Telmatogeto­ very large and elongate, are used by Saether (1977 : ninae and Prodiamesinae have not been reported 31, Trends 1, 2) to support the phylogenetic place­ from amber. ment of the Telmatogetoninae since most of the apo- morphies are autapornorphies. The eggs of Telma­ « There is strong reason to suppose that the basic togetoninae, i.e. Telmatogeton (and presumably also and first steps in the phylogenetic evo­ Thalassomya), are always laid singly, without a gela­ lution of the family took place in the upper course tinous medium, with the micropyle uppermost of small, cool mountain streams and their springs, (Terry 1913, sub Charadromyia). The gonostyli in Tel­ and that the present widespread occurrence of the matogetoninae, with the cerci, probably help in chironomid in other types of habitats is due orientation and the correct positioning of the indi­ to secondary adaptative radiation » (Brundin 1966 : vidual eggs during attachment to the substrate. 455). The larvae of most rheophilic chironomids feed Much more control is required for this kind of ovo- on diatoms and this diet has permitted chironomids position than is found in those chironomids to evolve and radiate independantly of higher plants 52 P. ASHE, DA MURRAY ET F. REISS (26) which lay an egg-mass. It would seem therefore that when the continuous land connection from Austra­ most of the unique features of the female genitalia lia through Antarctica to South America was finally of Telmatogetoninae are highly apomorphic and broken. Those genera (and subgenera) which occur have developed to suit the needs of a very harsh envi­ in both the Palaearctic and Nearctic regions date ronment. The female genitalia of Telmatogetoninae back at least to the time just prior to the final sepa­ have developed some remarkable features but prac­ ration of Europe and Eastern Russia from North tically all these features appear to be apomorphies America. (loss of the gelatinous egg-mass) or autapomorphies (loss of spermathecae, Gp VIII very large and elon­ 5. Taxonomic notes and new faunal gate, Gs IX present with elongated cerci to facili­ records tate ovoposition). It is not within the scope of this work to deal with the phylogeny of the Telmatoge­ Apsectrotanypus toninae in any greater detail since it requires a much more detailed analysis and because of this no for­ According to Fittkau & Roback (1983) species from mal change is proposed for the phylogenetic posi­ the Neotropical and Australasian regions identified tion of the Telmatogetoninae. as Macropelopia may belong to Apsectrotanypus. Larvae similar to Apsectrotanypus are known from According to Brundin (1976 : 153) the S. African/W. the Neotropics. Australian vicariance pattern displayed by Archaeochlus is a consequence of the Upper Juras­ Conchapelopia sic separation between S. Africa and E. Antarctica Australasian region : Adult males have been col­ and (Brundin, loc. cit. : 148) that the existance of the lected from the McLeod River about 18 km west of genus in the Upper Jurassic supports the idea that Mt. Carbine in Queensland (F. Reiss). the Podonominae with its two tribes, Boreochlini and Podonomini, are at least 175 million years old. Djalmabatista Brundin (1976 : 159) in summary states : « there is This genus has been found in a river in the tran­ good reason to conclude that representatives of all sition zone between the Palaearctic and Oriental Chironomidae subfamilies and many tribes existed regions (Fittkau & Roback 1983) and it is therefore in the Jurassic ». Prior to the breakup of the sou­ regarded as occurring in both these regions. thern continents India occupied a position between Southern Africa and Western Australia and Labrundinia Archaeochlus must have existed in Southern India In Fittkau & Roback (1983) Labrundinia is erro­ at that time. Climatic changes that occurred as India neously stated to have a worldwide distribution, drifted northwards were probably not significantly there are no published records from the Afrotropi­ different to that which exists today and it is possi­ cal or Australasian regions. ble that Archaeochlus still survives in Southern India or Sri Lanka. Nilotanypus Neotropical region : Two new species have been An estimate of the minimum age of some chiro­ discovered in the Amazon basin region (F. Reiss). nomid genera can be made in a similar way to Brun- din's estimate regarding Archaeochlus. The shared Australasian region : Pupal exuviae have been presence of a particular genus or group of genera found in recent collections from creeks in New on different land masses which are now separated South Wales and Queensland (F. Reiss). from one another, but which were formerly in close Oriental region : Pupal exuviae have been found contact, indicates that the minimum age of those in a stream in northern Sulawesi, Indonesia (P. Ashe) genera, can in some cases, be dated to at least that and from Yunnan Province, China (F. Reiss). time prior to the break-up of the original land-mass. The shared presence of certain genera and a subge­ Procladius nus in S.E. Australia/New Zealand and the Chaudhuri & Debnath (1983) describe several spe­ Andean/Patagonian region (Table 6) indicates that cies of Procladius from the Oriental region (India) all these taxa are at least 45 million years old dating and assign them to the subgenera Calotanypus, Psi- from the separation of Australia from Antarctica lotanypus and Procladius s.str. Roback (1982c) (27) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 53 recently redescribed Procladius and its subgenera adult male collected near Panguana, Peru (F. and it is apparent that the subgenera identified in Reiss). Chaudhuri & Debnath (op. cit.) are incorrect but from their illustrations of the male genitalia it is pos­ sible to recognise the correct subgenera. The spe­ Oriental region : Pupal exuviae have been collec­ cies included in Calotanypus belong in Djalmaba- ted in surface drift in a tributary of the River Mung tista, the Psilotanypus species appear to belong to Lun, Yunnan Province, China (F. Reiss). Procladius s.str. and those included in Procladius s.str. belong to Holotanypus. One of the Indian spe­ Eurycnemus cies, Procladius noctivagus (Kieffer), which also This genus is only known with certainty from the occurs in the Afrotropical region, belongs in the sub­ Palaeartic region and reports to date of the genus genus Holotanypus and this subgenus now occurs from the Nearctic, Neotropical and Australasian in both the Oriental and Afrotropical regions. regions are erroneous.

Rheopelopia Krenosmittia Oriental region : Pupal exuviae has been collected Oriental region : A male adult has been collected in drift samples from a stream in northern Sulawesi, in Guandong Province, China (F. Reiss). Indonesia (P. Ashe). Archaeochlus Orthocladius (Eudactylocladius) Downes (1974) reports the presence of an undes- Afrotropical region : A pupa belonging to this sub­ cribed species of Archaeochlus from Western Aus­ genus has been found high on Mount Kenya (Crans­ tralia. The species posesses well developed mandi­ ton pers. comm. to P. Ashe). bles in the adult. Pa racricotopus Podonomopsis Oriental region : Pupal exuviae have been collected This genus was incorrectly included as a nomen in a stream in northern Sulawesi, Indonesia (P. Ashe). nudum only in the generic catalogue (Ashe 1983) and the fact that the genus had been validly described Pa ralimnophyes by Brundin (1966 : 272) was unfortunately omitted. Nearctic region : This genus has been recently dis­ covered in the northern Nearctic (Cranston & Oli­ Arctodiamesa ver, pers. comm. to P. Ashe). The uncertain presence of Arctodiamesa in the Parasmittia Nearctic (Saether et al. 1984) has been confirmed from specimens collected in Alaska (Cranston & Oli­ Nearctic region : One species of the genus has ver, pers. comm. to P. Ashe). been found in Canada (Cranston & Oliver, pers. comm. to P. Ashe). Monodiamesa Pa ratrissocladius Oriental region : Two species have been found at Neotropical region : Adults of this genus have Taoyuan, Hunan Province, China (F. Reiss). been discovered in Southern Chile (Halvorsen, pers. Apometriocnemus comm. to P. Ashe). Palaearctic region : This genus has been discove­ Rheocricotopus red in material collected in Ireland (Heneghan, pers. The subgenera to which the Neotropical and Aus­ comm.). tralasian members of the genus belong is at present unknown. Diplosmittia Neotropical region : This genus which was origi­ Semiocladius nally described from the West Indies within the Neo­ Oriental region : This genus has been found tropical region (Saether 1981) has recently been dis­ recently in the Oriental region (Cranston, pers. covered on the mainland of South America as an comm. to P. Ashe). * 54 P. ASHE, D.A. MURRAY, F. REISS (28)

Stictocladius type-material of C. brasiliensis and N. heterochirus Sticîocladius should not be regarded as a subge­ is completed. Until this problem is resolved the con­ nus of Diplocladius but warrants full generic status cept of Nilodorum should be based on N. brevibucca because it has a larva of « Genus nr. Lopescladius » and on the diagnoses of the larva, pupa and adult type as described by Coffman & Roback (1984) of the genus as given in the key work on the « Chi­ (Cranston & Edward, pers. comm. to P. Ashe). ronomidae of the Holarctic region » (Wiederholm 1983 et seq). Chemovskiia Oriental region : Adults of this genus have been found at Taoyuan, Hunan Province, China (F. Reiss). Oriental region : Adult males, pupae and pupal exuvia have been collected from the Rangeet River, Cladopelma Darjeeling, India ; pupal exuvia which probably belong to this genus have been collected in a tribu­ Neotropical region : Cryptocladopelma boydi tary of the River Mung Lun, Yunnan Province, China (Lichtenberg) from Columbia now belong in (F. Reiss). Cladopelma. Afrotropical region : See note under Oriental region : There are unpublished records . from Sri Lanka, India and China (Yunnan Province) (F. Reiss). Cryptotendipes Australasian region : A single adult male has been discovered near the Mc Leod River, 18 km west of Oriental region : Pupal exuviae collected in a tri­ Mount Carbine, Queensland (F. Reiss). butary of the River Mung Lun, Yunnan Province, China (F. Reiss). Afrotropical region : According to Cranston (pers. Neotropical region : Adult males belonging to this comm. to P. Ashe) the single African species inclu­ genus have been found in Venezuela (F. Reiss). ded in Paracladopelma in Freeman & Cranston Polypedilum (Tripodura) (1980) is really a Cryptotendipes species. Oriental region : Several species belonging to the Goe Idichi ronomus subgenus Tripodura have been found in the Orien­ G. holoprasinus (Goeldi) has been found in Hawaii tal part of China (R. Reiss). and is apparently a recent aircraft introduction Pontomyia (Reiss & Sublette 1985). Nearctic region : Larvae have been collected in a Nilodorum bottom grab at Sapper Point, near Key Largo, Flo­ The concept of Nilodorum as used by all recent rida (Caldwell, pers. comm. to P. Ashe). authors and in this paper is based on Nilodorum bre- Robackia vibucca Kieffer which was described from Africa and which was until recently regarded as the type- Afrotropical region : Adult males have been col­ species of the genus. When compiling a catalogue lected around Samburu Lodge, Kenya (leg S. Koch, of chironomid genera (Ashe 1983) it was discovered det. Saether). that the genus was described in an earlier paper and Oriental region : Pupal exuviae and a mature pupa N. heterochirus Kieffer automatically became the were collected from the Darjeeling River at type-species by monotypy. N. heterochirus was des­ 250 m.a.s.l. The genus is also known from pupal exu­ cribed from the Island of Trinidad (Neotropical viae collected from a tributary of the River Mung region) and was regarded by Edwards (1931 : 320) Lun, Yunnan Province and at Taoyuan, Hainan Pro­ as a synonym of Chironomus brasiliensis Wiede­ vince, China (F. Reiss). mann, 1828. It seems likely that a new name will be required for Nilodorum sensu auctt. with N. brevi- Saetheria bucca as the type-species but this cannot be done Neotropical region : Adult males and pupal skins until a re-examination of the have been found in the Amazonian region (F. Reiss). (29) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 55

StempeîUneîîa subfamilies to prevailing environmental conditions Afrotropical region : One species has been found (oxygen concentration, water temperature, climate, recently at Lake Tana, Ethiopia (F. Reiss). altitude, current speed in flowing water, etc.). The Telmatogetoninae are an exception to the above rule in that they are predominantly marine and a diffe­ 6. Overview of chironomid distribution rent set of factors apply. Our knowledge of the eco­ Of the 307 described valid genera only 38, repre­ logy of the three smallest subfamilies, Chilenomyii­ senting the subfamilies Telmatogetoninae, Tanypo­ nae, Buchonomyiinae and Aphroteniinae, is very dinae, Orthoclàdiinae and Chironominae, are known incomplete but they appear to mostly avoid clima­ to have a worldwide distribution. The investigation tic or environmental extremes. of different habitat types in various areas of each In chapter 4 we discussed the importance of con­ zoogeographical region is important to determine tinental drift in relation to the distribution of some if particular genera are present or likely to be chironomid genera. Such vicariance events have absent. Genera which occur in very specific and dif­ played the primary role in establishing the distri­ ficult to sample habitats are often missed by nor­ bution of chironomid subfamilies, tribes and some mal collecting methods and this may account for genera since the breakup of Pangaea. However, the some of the anomaly in generic representation bet­ distribution of all chironomid genera cannot be ween the Nearctic and the Palaearctic regions. To explained by vicariance events alone and dispersal date the known world chironomid fauna totals some (e.g. Diamesa, Chapter 3.7) has played an important 3700 valid species (Table 3), representing perhaps role, in recent times, since the middle to recent Ter­ only 30 % - 35 % of the estimated chironomid fauna tiary. A comparison of the chironomid fauna of the of 10,000 - 12,000 species. In terms of the numbers Holartic with that of the Neotropical, Afrotropical of species and our knowledge of the distribution of and Australasian regions respectively reveals that species within regions the Palaearctic is the best the Neotropical region has exchanged some cold known followed by the Nearctic, Afrotropical, Aus­ adapted and many warm adapted groups with the tralasian, Neotropical and Oriental. Most Neotropi­ Holarctic ; the Afrotropical region has exchanged cal and Oriental species are only known from one mainly warm adapted groups and the Australasian or two countries with many only known from the region (as present incomplete knowledge indicates) original type-locality. Our knowledge of the world has exchanged the least (mainly some warm adap­ fauna outside the Holarctic can be brought into ted groups). The ability of each of the three regions perspective if the figures for the number of genera to exchange faunal elements with the Holarctic is (Table 1) and species (Table 3) for the other regions directly related to the physical conditions linking are compared with a small, well investigated, region or separating these regions from the Holarctic. An of the Holarctic e.g. Great Britain. Great Britain, almost continuous mountain chain, which stretches with about 128 genera and over 500 species (Lang- from Alaska through Central America and linking ton 1984), occifpies a land area of only to the Andes mountains (which extend from Colom­ 229,979 sq.km. and has a fauna which at present is bia to Patagonia), provides a pathway by which cold greater than the described fauna of each region adapted forms from the Nearctic and Neotropical except the Palaearctic and Nearctic. regions can migrate Southwards and northwards respectively. A similar exchange of warm adapted For the major subfamilies the following general groups between the Neotropics and the Nearctic is rule applies : Progressing from polar regions to the possible along the lowland coastal strips of Central equator (i.e. high latitudes to low latitudes) or from America and by island hopping along the Lesser and high mountains to lowlands there is an increase in Greater Antilles to Florida. In the Afrotropical diversity and numbers of specimens of the Chiro­ region the lack of a continuous high mountain range nominae and Tanypodinae with a corresponding between the East African Highlands and the Dra- decrease in the Orthoclàdiinae, Podonominae, Dia­ kensberg Range has resulted in the fact that most mesinae and Prodiamesinae ; the reverse is true pro­ (if not all) cold adapted Palaearctic groups have not gressing from the equator towards the polar regions reached Southern Africa although a few extend as or from lowlands to high mountains. This rule is a far south as the East African Highlands. A reflection of the adaptations of the major 56 P. ASHE, D.A. MURRAY. F. REISS (30) considerable number of warm adapted groups have permission to include some unpublished generic records ; been able to migrate from the Afrotropical region in addition we wish to thank the following individuals for northwards and from the southern Palaearctic permission to include information or records relating to particular genera : Dr. B.A. Caldwell, Georgia Dept. Nat. southwards mainly via the Middle East and along Res., Atlanta, Georgia, U.S.A. ; Dr. D.H.D. Edward, Zoology the Nile Valley (the Sahara, in more recent times at Dept., University of Western Australia, Australia ; Dr. E.J. least, has acted as a barrier to migration north or Fittkau, Zoologische Staatssammlung, Munich, Germany ; south). The Australasian region is separated from Dr. G.A. Halvorsen, Museum of Zoology, Bergen, ; the Palaearctic by the Oriental region and in order Dr. D.R. Oliver, Biosystematic Research Centre, Ottawa, Ontario, Canada and Dr. E. Willassen, Museum of Zoology, for any exchange to take place between the Palaearc­ Bergen, Norway. We are grateful to Dr. H. Laville, Tou­ tic and Australasian regions groups must occur in louse, France for providing the French abstract. and have migrated through the Oriental region. Migration between the Palaearctic and Australasian References regions is very difficult because there is no conti­ (Numbers opposite some references are the sources for published nuous land connection and large areas of open sea zoogeographical distribution data given in the detailed list of exist interspersed with numerous islands. Very few genera and subgenera). Aagaard (K). 1978. — The Chironomidae of the exposed zone of cold adapted Palaearctic or Australasian groups are 0vre Heimdalsvatn. Holarct. Eco!., 1 : 261-265. likely to be able to migrate south and north respec­ Ashe (P.). 1983. — A catalogue of chironomid genera and subge­ nera of the world including synonyms (Diptera : Chironomi­ tively because of the absence of high mountains dae). Ent. scand. Suppl., 17: 1-68. necessary for temperate conditions to exist and Ashe (P.). 1985. — A larval diagnosis for the subfamily Bucho­ because of the lack of continuity of the mountain nomyiinae and the genus Buchonomyia with a description of the 1 st instar larva of Buchonomyia thienemanni Fittkau (Dip­ ranges of the East Indies with the Himalayas or with tera, Chironomidae). Spixiana Suppl., 11 : 143-148. the New Guinea Highlands. The main exchange (1) Ashe (P.) & Murray (D.A.). 1980. — Nosiococladius, a new sub­ likely is between warm adapted groups but our genus of Cricotopus (Diptera Chironomidae), pp. 105-111. In Murray D.A., ed. Chironomidae - Ecology. Systematics, Cytology knowledge of the Australasian fauna and that of the and Physiology. Pergamon Press, Oxford, 354 pp. lowlands of the eastern Palaearctic (central China) Beck (W.M.).-1977. — Environmental requirements and pollution and S.E. Asia is very incomplete. tolerance of common freshwater Chironomidae. Envir. Monit. Ser., U.S. Envir. Protect. Ag., Cincinnati, 261 pp. Beedham (G.E.). 1966. — A chironomid (Dipt.) larva associated An analysis of the zoogeographical distribution of with the Lamellibranchiate mollusc, Anodonta cygnea L. Ento­ any group relies on a vast amount of accu­ mologist's mon. Mag.. 101 : 142-143. Boesel (M.W.). 1937. — Order Diptera, Family Chironomidae. In rate and reliable data on the distribution of indivi­ Carpenter F.M., ed. Insects and arachnids from Canadian dual taxa. In spite of the detailed information pre­ amber. Univ. Toronto Stud. Geol. Ser.. 40 : 44-55. sented here our knowledge of the distribution of (2) Borkent (A.). 1984. — The systematics and phylogeny of the Stenochironomus complex (Xestochironomus, Harrisius, and individual genera/subgenera throughout the world Stenochironomus) (Diptera : Chironomidae). Mem. ent. Soc. is still very inadequate and it is not possible to deal Can., No. 128: 1-269. with the zoogeography in any greater detail. For the Bretschko (G.). 1982. — Pontomyia Edwards (Diptera : Chirono­ midae), a member of the coral reef community at Carrie Bow majority of genera it is not yet possible to deal with Cay, Belize pp. 381-385. In Riitxler K. & Macintyre I.G., eds. such topics as « centre of origin », minimum age or The Atlantic barrier reef community at Carrie Bow Cay, Belize, 1 : Structure and communities. Smithson. contrib. mar. Sci., even the total distribution within a zoogeographi­ 12, 539 pp. cal region. We need to know more about the present (3) Brundin (L.). 1956. — Die bodenfaunistischen Seetypen und distribution of individual taxa, their phylogenetic ihre Anwendbarkeit auf die Sùdhalbkugel. Zugleich eine Théo­ rie der produktionsbiologischen Bedeutung der glazialen Ero­ relationships, ecology and fossil record and relate sion. Rep. Inst. Freshwat. Res. Drottningholm, 37 : 186-235. this data to geological and climatological events in (4) Brundin (L.). 1966. — Transantarctic relationships and their different regions over the time scale of the existance significance, evidenced by chironomid midges. With a mono­ graph of the subfamilies Podonominae and Aphroteniinae and of the group. the austral Heptagyiae. K. svenska VetenskAkad. Handl, 11 : 1-472. Brundin (L.). 1970. Diptera Chironomidae of South Georgia. Pacif. Ac know ledge ment s Insects Monogr., 23 : 276. Brundin (L.). 1976. — A Neocomian chironomid and Podonominae- Aphroteniinae (Diptera) in the light of phylogenetics and bio- The authors are particularly grateful to Dr. P.S. Crans­ geography. Zool. Scr., 5 : 139-160. ton, British Museum (Nat. Hist.), London, England for his (5) Brundin (L.). 1981. — Croizat's panbiogeography versus phylo­ constructive critism of the paper, for providing figures on genetic biogeography pp. 94-138. In Nelson G. & Rosen D.E., subfamily representation for the Australasian and Nearc­ eds, Vicariance Biogeography : A Critique. Columbia University tic regions based on unpublished catalogues and for Press, New York, 616 pp. (31) THE ZOOGEOGRAPHICAL DISTRIBUTION OF CHIRONOMIDAE 57

(6) Brundin (L.). 1983a. — Chilenomyia paradoxa gen. n., sp. n. and Drainai (E). 1979. — Eukiefferiellaszczensnyi sp.n.(Diptera,Chi­ Chilenomyiinae, a new subfamily among the Chironomidae ronomidae). Bull Acad pot. Sci. Cl. IISér. Sci. biol, 27 : 183-193. (Diptera). Ent. scand., 14: 33-45. Edwards (F.W.). 1931. — Diptera of Patagonia and South Chile. (7) Brundin (L.). 1983b. — The larvae of Podonominae (Diptera : Part II. Fascicle 5. - Chironomidae, pp. 233-331. Trustees of the Chironomidae) of the Holarctic region - keys and diagnoses. British Museum, London. Ent. scand. Suppl, 19 : 23-31. Edwards (M.)&Usher (M.B.). 1985. — The winged Antarctic (8) Brundin(L). 1983c. — Two newaphrotenian larval types from Parochlus steinenii (Gercke) (Diptera : Chironomidae) in the Chile and Queensland, including Anaphrotenia lacustris n. gen., South Shetland Islands. Biol. J Linn Soc., 26 : 83-93 n. sp. (Diptera : Chironomidae). Ent. scand., 14: 415-433. Fittkau (E.J.). 1974. — Ichthyocladius n. gen., eine neotropische Brundin (L.) & Saether (O.A.). 1978. — Buchonomyia burmanica Gaining der Orthoclàdiinae (Chironomidae, Diptera) deren Lar- sp. n. and Buchonomyiinae, a new subfamily among the Chi­ ven epizoisch auf Welsen (Austroblepidae und ) ronomidae (Diptera). Zool. Scr., 7 : 269-275. leben. Ent. Tidskr. Suppl, 95 : 91-106. Chaudhuri (P.K.)&Debnath(R.K.). 1983 — Studies of Indian Tany- (21) Fittkau (E.J.) & Murray (D.A.). 1985. — Radotanypus a new podine (Diptera : Chironomidae), genus Procladius Skuse. Zool. genus of Tanypodinae from the Nearctic (Diptera, Chironomi­ Jb., Syst., 110: 111-123. dae). Spixiana Suppl, 11 : 209-213. (9) Chaudhuri (P.K.) & Ghosh (M.). 1981. — Anew genus of podo- (22) Fittkau (E.J.) & Reiss (F.). 1978. — Chironomidae, pp. 404-440. nomine midge (Chironomidae) from Bhutan. Syst. Ent., 6 : In lilies J., ed„ 2nd edition. Limnofauna Europaea. Gustav Fis­ 373-376. cher Verlag, Stuttgart, 532 pp. (10) Chaudhuri (P.K.) & Ghosh (M.). 1982. — Record of Chaetocla- (23) Fittkau (E.J.) & Reiss (F.). 1979. — Die zoogeographische Son- dius Kieffer (Diptera : Chironomidae) from India. Folia ent. derstellung der neotropischen Chironomiden (Diptera). hung., 43 : 5-7. Spixiana, 2 : 273-280. (11) Chaudhuri (P.K.) & Ghosh (M.). 1986. — Two Indian species (24) Fittkau (E.J.) & Roback (S.S.). 1983. — The larvae of Tany­ of Kiefferulus Gcetghebuer (Diptera : Chironomidae). Svst. Ent., podinae (Diptera : Chironomidae) of the Holarctic region - keys 11 : 277-292. and diagnoses. Ent. scand. Suppl, 19: 33-110. (12) Chaudhuri (P.K.) & Nandi (S.K.). 1980. — On a new species Forsyth (D.J.) & McCallum (I.D.). 1978. — Xenochironomus can- of genus Allotrissocladius Freeman (Diptera, Chironomidae) terburyensis (Diptera : Chironomidae), a commensal of Hyri- from India. J. Bombay nat. Hist. Soc, 77 : 292-294, delta menzieensis (Lamellibranchia) in Lake Taupo ; features (13) Chaudhuri (P.K.), Debnath (R.K.) & Nandi (S.K.). 1983. — of pre-adult life history. N.Z. Jl Zool, 5 : 795-800. Tanypodine midges of the genus Ablabesmyia Johannsen (Dip­ Freeman (P.). 1955. — A study of the Chironomidae (Diptera) of tera : Chironomidae) from West Bengal with a note on their Africa south of the Sahara. Part I. But!. Br. Mus. nat. Hist., Ent., seasonal incidence and sex ratios. /. nat. Hist., 17 : 901-917. 4: 1-67. ( 14) Cheng (L.)&Hashimoto (H.). 1978. — The marine midge Pon- Freeman (P.). 1956. — A study of the Chironomidae (Diptera) of tomyia (Chironomidae) with a description of females of P. Africa south of the Sahara. Part II. Bull. Br. Mus. nat. Hist., Ent., oceana Tokunaga. Syst. Em., 3: 189-196. 4: 285-366. (15) Coffman (W.P.). 1978. — Chironomidae, 345-376. In Merritt Freeman (P.). 1957. — A study of the Chironomidae (diptera) of R.W. & Cummins K.W., eds. An introduction to the aquatic Africa south of the Sahara. Part III. Bull. Br. Mus. nat. Hist., Insects of North America. Kendall/Hunt Publishing Compagny, Ent., 5 : 321-426. Iowa, 441 pp. Freeman (P.). 1958. — A study of the Chironomidae (Diptera) of (16) Coffman (W.P.) & Roback (S.S.). 1984. — Lopescladius (Cor- Africa south of the Sahara. Part TV. Bull Br. Mus. nat. Hist., diella) hyporheicus, a new subgenus and species (Diptera : Chi­ Ent.. 6: 261-363. ronomidae) Proc. Acad. nat. Sci. Philad., 136 : 130-144. (25) Freeman (P.). 1959. — A study of the New Zealand Chirono­ Cranston (P.S.) & Ashe (P.). in press. — Family Chironomidae. In midae (Diptera, ). Bull Br. Mus. nat. Hist., Ent., 7 : Soôs À. & Papp L., eds. Catalogue of Palaearctic Diptera, 393-437. Volume 2 : Psychodidae - Chironomidae. Elsevier, Amster- (26) Freeman(P.). 1961. — The Chironomidae (Diptera) of Austra­ dam/Akadémiai Kiado, Budapest. lia. Aust. 1. Zool, 9: 611-737. Cranston (P.S.) & Oliver (D.R.). in press. — Problems in Holarc­ Freeman (P.). 1962. — Chironomidae from the Batu Caves, Malaya tic chironomid biogeography. Ent. scand. Suppl.,. (Diptera : Nematocera). Pa*Hf. Insects, 4 : 129131. (17) Cranston (P.S.) & Saether (O.A.). 1986. — Rheosmittia (Dip­ (27) Freeman (P.). 1964. — Notes on Chironomidae (Diptera : tera : Chironomidae) : a generic validation and revision of the Nematocera). Proc. R. ent. Soc. Lond., (B) 33: 147-150. western Palaearctic species. /. nat. Hist., 20: 31-51. (28) Freeman (P.) & Cranston (P.S.). 1980. — FamilyChironomidae, (18) Cranston (P.S.), Oliver (D.R.)& Saether (O.A.). 1983. — The lar­ pp. 175-202. In Crosskey R.W.. ed. Catalogue of the Diptera of vae of Orthoclàdiinae (Diptera : Chironomidae) of the Holarc­ the Afrotropical Region. British Museum (Nat. Hist.), London, tic region - keys and diagnoses. Em. scand. Suppl, 19: 149-291. I, 437 pp. (19) Danks (H.V.). 1980. — of Polar Bear Pass. Glover (B.). 1973. — The Tanytarsini (Diptera : Chironomidae) of Bathurst Island, arctic Canada. Sytlogeus, No. 25 : 1-68. Australia. Aust. J. Zool, Suppl Ser., 23 : 403478. Davies (L.). 1973. — Observation on the distribution of surface- Gordon (M.J.), Swan (B.K.) & Paterson (C.G.). 1978, — Baeoctenus living land arthropodes on the subantarctic Ile de la Posses­ bicolor (Diptera : Chironomidae) parasitic in unionid bivalve sion, lies Crozet. 7. nat. Hist., 7 : 241-253. molluscs, and notes and other chironomid-bivalva associations. Disney (R.H.L.). 1975. — Notes on crab-phoretic Diptera (Chiro­ /. . Res. Bd Can., 35 : 154-157. nomidae and Simulidae) and their hosts in Cameroon. Ento­ Gotceitas (V.) & Mackay (R.J.). 1980. — The phoretic association mologist's mon. Mag., Ill : 131-136. of Nanocladius (Nanocladius) rectinervis (Kieffer) (Diptera : Chi­ Dosdall (L.M) & Mason (PG.). 1981. — A chironomid (Nanocladius ronomidae) on Nigronia scrricomis Say (Megaloptera : Coryda- (Plecopteracoluthus) branchicolus : Diptera) phoretic on a sto- lidae). Can. J. Zool, 58: 2260-2263. nefly (Acroneuria lycorias ; Plecoptera) in Saskatchewan. Can. (29) Guha (D.K.), Das (S.K.), Chaudhuri (P.K.) & Choudhuri (D.K.). Ent., 113: 141-147. 1985. — Chironomid midges of the Andaman Islands (Diptera : (20) Downes (J.A.). 1974. — The feeding habits of adult Chirono­ Chironomidae). Proc. nat. Acad. Sci. India, 55 (B) : 22-38. midae. Ent. Tidskr. Suppl, 95 : 84-90. Guibé (J.). 1942. — Chironomes parasites de Mollusques Gastro­ podes. Chironomus varus limnaei Guibé espèce jointive de Chi- ronomus varus varus Gtgh. Bull biol. Fr. Betg., 76 : 283-297. 58 P. ASHE, D.A. MURRAY. F. REISS (32)

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