© Entomologica Fennica. 27 .VIII.l991

The karyotype, morphology and ecology of aequalis Kieffer (Diptera, )

Mauri Hit-venoja & Paraskeva Michailova

Hirvenoja, M. & Michailova, P. 1991: The karyotype, morphology and ecol­ ogy of Glyptotendipes aequalis Kieffer (Diptera, Chironomidae) .- Entomol. Fennica 2:87-96.

The karyotype, morphology of adult, pupae and larvae, and the ecology of Glyptotendipes aequalis Kieffer (1922) are described from material from Finland, and a neotype from the (probably) original German material is designated. 2n = 8. 1st, lind and Illrd chromosomes are metacentric, IVth acrocentric. The IVth chromosome has a nucleolus and a Balbiani ring. G. aequalis is karyologically closely related to G. caulicola , differing by a simple inversion step. Mauri Hirvenoja, Sotilaskorventie 13,01730 Vantaa, Finland Paraskeva Michailova, Bulgarian Academy of Sciences, Bout. Rouski 1, 1000 Sofia, Bulgaria

1. Introduction locality is missing (there being no data label), this is very probably the specimen mentioned by The male of Glyptotendipes aequalis Kieffer Lenz, because only one side is provided with an (1922:79) was originally described from material anal comb. collected in Westfalen, Gennany by A. Thiene­ The origin of the specimen is, however, not mann. Up to now, however, no original adult completely certain and thus we cannot know if it specimens have been found in any coll ection. is the cast skin of the lost holotype. Thus, (ICZN, Lenz ( 1954- 1962: 176) gives a short descrip­ Opinion 1147, 1980) it cannot be considered as a tion of the pupa of G. aequalis as a species of the lectotype, but the probability that this specimen caulicola group in the subgenus Phytochironomus belongs to the original material is very great. Kieffer: Because it may thus provide the best existing evi­ "Analkamm mit nur 2-3 Zahnen, die bei der e in zigen dence of the characters of the original G. aequalis, vorh andenen Exuvie sogar auf der e inen Seite fehlen. Auf this pupal skin (cf. below p. 93) has here been der Analflosse oral-median e in Paar he lle, starre, mittel­ designated as the neotype of the species. grol3e Borsten. " According to "Limnofauna Europaea" (Fitt­ According to the same paper, G. aequalis kau & Reiss 1978) G. aequalis is still known from was at that time known only from Westfalen southern Germany only. By contrast, Glypto­ (Mi.in ster) , Germany. Thus the cast pupal skin tendipes foliicola Kieffer according to the same which Lenz has seen may be a part of the original paper is widely distributed and has perhaps in material. Dr. F. Reiss (Zoologische Staatssamm­ some cases been confused with G. aequalis. The lung, Munich, Germany) in 1988 found and pre­ correct identification of G.foliicola seems so far pared one pupal skin from Thienemann 's coll ec­ to be impossible and this has been considered as tion. Although information about the sampling a nomen dubium (Hirvenoja 1990). 88 Hirvenoja & Michai/ova: Glyptotendipes aequalis • ENTOMOL. FENNICA Vol. 2

A species detem1ined as G. aequalis by the aid 2.2. Male (N=6, Fig. !A) of the neotype and the literature is found in two different parts of Finland. An attempt to give A quite dark species with more or less pale more definitive descriptions of adults and imma­ scutellum and more or less pale parts on legs as ture stages follows in the present paper. follows: bases offemora, middle of mid and hind In addition the present work deals with the tibia; the tarsi darken towards the last joints in first description of the karyotype under the name the mid and hind legs. G. aequalis. We demonstrate the photomap of Antenna! ratio 3.0-4.2. Frontal tubercles about the polytene clu·omosomes, as well as present 10-15 j..lm. Palp segments (2-5): 90-115, 205- information about the chromosome polymor­ 275, 205-230 and 240-275 j..lm. Clypeus with phism in one Finnish population. The karyotype < 35 and postocular region with < 24 setae. relationships with other species of the genus Number of acrostichals < 18; dorsocentrals (16- Glyptotendipes Kieffer are also investigated. 34) in 2-3 rows, sometimes also mostly irregul ar standing scutellars (25-29) in two rows. Tracheal scar of pupal respiratory organ about 3/5 length of postpronotum. Wings 3.0-3.2 j..lm. Costa not

2. Morphology of adults and immature extended over R4+5; setae on R 1 and on the distal stages part of R4+5; squama with 16- 18 setae. About 15-mm-long sensilla chaetica near 2.1. Material and methods distal end of basitar sus of mid leg 6- 11 in number. Fore leg: leg ratio 1.23- 1.35; beard ratio 2.5-3.2. A great number of adults were reared from lar­ Mid leg: leg ratio 0.48-0.50; beard ratio 2.8-3.0. vae from a garden pond in Vantaa near Helsinki Hind leg: leg ratio 0.57-0.62; beard ratio 3.8- (May 1988). Measurements were made on non­ 4.5. Legs in j..lm : macerated material mounted in Euparal. Mor­ Fore leg Mid leg Hind leg phologically identical specimens had been reared Femur 1200- 1400 1315- 1570 1430- 1630 earlier from a forest clay pond on the Hirvenoja Tibia 1115- 1285 1230- 1455 1515- 1715 farm, Riihimaki (May to August 1970), also in Tarsus 1430- 1640 585-745 855-1055 southern Finland. Some of the latter were also 1 Tarsus2 785-855 415-455 540-630 included in the measurements in addition to the Tarsus 630-700 300-370 385-485 neotype (Fig. 1G) mentioned above. 3 Tarsus4 1500-515 12 15-255 255-285 The morphological terminology as used here Tarsus 230-240 155- 185 145- 185 is in regard to some details more classical than, 5 for instance, in "Chironomids of the Holarctic Hypopygium (Fig. 1A) resembles that of the region" (Wiederholm ed. 1983, 1986, 1989). The common type in the genus Glyptotendipes. nomenclature of the male genitalia used here is Processus analis dark, apically rounded and illustrated, for instance, in Reiss & Fittkau ( 197 1: broadest near the apex. Appendage 1 dark, quite fig. 1) and in Pinder (1978:13). The classical straight, proximally haired, apically with a strong nomenclature of the genitalia has been used be­ hook. Appendage 2 relatively broad compared to cause (according to the opinion of the author its length. About 10 setae on the pale spot of MH, cf. Hirvenoja 1973) most appendages of the tergite IX, not differing from the nearest setae in gonocoxite are of phallic origin, arising during surroundings of the pale area. the evolution of the chironomid and hav­ ing no true homological connections with Hy­ menoptera. In the nomenclature of the larval head 2.3. Female (N=6, Fig. 1B-C) Gouin's (1959, 1968) school of thought is ac­ cepted; defining the ventral wall of the head Colouration as well as type of chaetotaxy as in capsule of the larva as a submentum in recent the male, but the number of setae a little higher. papers represents an oversimplified morphologi­ Antennae with 5 flagellomeres or occasionally cal concept. with 6 if first is more or less clearly divided. ENTOMOL. FENNICA Vol. 2 • Hirvenoja & Michailova: Glyptotendipes aequalis 89

lfXJI'm E

lfXJI'm I IEF6

~ F A AJr4

Fig. 1. G/yptotendipes aequalis Kieffer.- Adults. A: Male hypopygium; B: Genitalia of female; C: Genitalia and shape of cercus seen more laterally.- Pupa. D: Variation of anal combs; E: Basal ring of the respiratory organ; F: Variation of cephalic tubercles in the slides; G: Variation of the racket-shaped plates of tergite (I I) Ill-VI (first row left drawn from the neotype).-Original. 90 Hirvenoja & Michailova: G/yptotendipes aequalis • ENTOMOL. FENNICA Vol. 2

Frontal tubercles 6-10 11m . Palp segments 2-5: are situated in 2- 3 rows, especially on the anal 70-80, 170-210, 185-220 and 220-345 11m . part of the lobes; dorsal pair of setae on the anal Wings 3. 1-3.8 11m. Sensilla chaetica on middle segment often present but found in variable and/or more on the apical part of the basitarsus places. of mid leg (39-55) and on middle of basitarsus of hind leg (39-54 in number). Fore leg: leg ratio 1. 37-1.41; beard ratio 2.0-2.3. Mid leg: leg ratio 2.5. Larva (N=9, Fig. 2) 0.47-0.48; beard ratio 2.5-3.1. Hind leg: leg ra­ tio 0.56-0.57; beard ratio 3.8-4.0. Legs in 11m: Length < 9 mm. Anterior parapods with simple, nearly invisible plumose or distictly serrate claws. Fore leg Mid leg Hind leg Behind the parapods, near the claws an unpaired, Femur 11 70-1285 1300-1430 1315-1430 triangular group of very small claw like chaetae, Tibia 1085-11 70 1270- 1355 1485- 1570 about 40-50 in number (in some species of Tarsus 1530- 1600 600- 655 840- 885 1 Glyptotendipes this group has been divided into Tarsus 770-830 385-415 5 15-555 2 two groups!). One pair about 150-200 11m long Tarsus 630- 655 315- 315 4 15-415 3 and 50-70 11m thick ventral tubules present on Tarsus 500-515 200- 230 240-255 4 segment IX; the anal tubules about 200-250 (the Tarsus 240-245 155-155 145- 170 5 upper pair) up to 300 11m long. Head brownish, Genitalia as in Fig. 1; spermathecae round or about 600- 630 11m long, 460-470 11m broad. somewhat oval, 140--18511m long. Gonocoxite IX Antenna! sockets and boundaries of frontal with 3 setae; segment X wi th 7-1 1 setae on each apotome a little darkened, forming a V-shaped side. figure, seen in the unprepared specimens but may partly disappear in slides. Borders of foramen occipitale dark, a quite large triangulum brown 2.4. Pupa (N= l2, Fig. IE-G) beginning at the tentorial pits; ventral part of the postgenal bridge a little darkened between the Length of the cast skins 8-10 11m . Frontal foramen occipitale and hind margin of the apotome with about 85-11 5 long cephalic tuber­ postmentum; also a narrow unsclerotized, pale cles, the preapical seta present. Thorax sulcus of this length ventromediall y in the area dorsomediall y granulated, more orall y smooth of fusion of both sides. Dorsal sclerites of the but darkened with reti cul ate pale lines. Small , head (Fig. 2K) agree with fig. 10.30.E in Pinder furrowed, sometimes not very clearl y visible & Reiss (1 983:381). anterolateral tubercles on the first abdominal The setae SI about 40 11m long, proximall y segment present; small PB and 76-88 booklets with one side feathered but apicall y with strong, present on segment II; ordinary pedes spurii A somewhat variable serration on both sides; setae on segment IV, the whole conjunctiva on seg­ SII simple. Great lateral chaetae feathered, few ment V with chagrin , segment VI with few small simple spinulae also present; between these spines on the hind corner. Structures of the ab­ groups a group of very small, hardly visible dominal segments much as in fig. 10.26 by Pinder chaetae (in some other species of G/yptotendipes & Reiss (1986). Racket- shaped plates on tergites clearl y to be found). Pecten epipharyngis with III-VI, in every fourth skin at the same place a 19-23 teeth , chaetul ae lateralis strongly serrated, pigmented spot found on tergite II. A pl ate (up to chaetulae basales (2) rudimentary. 350 11m ) covers about 1/4 - 1/3 of the total length Premandibles with 2 apical teeth , inner tooth of the tergites; there is great indi vidual variabil­ more or less sharpl y pointed, a little smaller than ity in the size of the plates, but the first (on outer. In mandibles with pale dorsal tooth, other segment III) is a li ttle smaller than the others, 4 teeth darkened. Apex of max illary sclerite which are all quite similar (Fig. I D). Anal comb clearl y dentate with 5- 7 rounded denticles; outer consists of some more or less loose curved spines. side of the galeolacinia (region of pal pi fer; Imms The anal segment is bordered with (one side) 64- 1957:24) with lengthened chaetae. Prementum 90 often uniserial filaments but sometimes these conical; lateral chaetulae quite narrow, not ENTOMOL. FENNICA Vol. 2 • Hirvenoja & Michailova: Glyptotendipes aequalis 91

I JOpm ·OEFGHIJ

Fig . 2. Glyptotendipes aequalis Kieffer.-Larva. A: Head in ventral vi ew; 8: Front border of the maxillary sclerite; C: Eye spots; D: Seta Sl; E: Pecten epipharyngis; F: Apex of premandible; G: Front border of paralabial plate; H: Claws of parapods; 1: Maxille antiaxially; J: Prementum; K: Head in dorsal view; L: Last abdominal segments. ­ Original. markedl y differentiated, abundantly present and 3. Karyological studies those standing most laterally seem to bend in­ wards in direct ventral view. Middle tooth of the 3.1. Material and methods chin or hypochilum (Gouin 1959) with small lateral denticles; from the six other teeth the Nine larvae from the garden pond in Vantaa were outermost 3rd clearly smallest. Paralabials totally fixed in 3 : 1 ethanol : aceti c acid. The analysis was striated, front border smooth but figured (depends petformed on squashed preparations of salivary somewhat on preparation technique) as in Fig. gland polytene clu·omosomes made by the standard 2G. Antennae about 180 J•.Un long. RO in first third aceto-orcein technique. The cluumosome ann s of of basal segment. Proportions of lengths of the G. aequalis have been called A toG on the basis of antenn a! segments in fourth instar (6 larvae): their homology with some sections of G. glaucus (Meigen) clu·omosomes. The latter species has been 100 : 30 : 24 : 20 : 6 100 : 35 : 28 : 23 : 7 suggested as a standard (Michailova 1989). G. 100 : 32: 29: 24: 6 100 : 37 : 27 : 23 : 7 aequalis polytene chromosomes have distinctive 100 : 3 2 : 21 : 18 : 6 100 : 38 : 27 : 19 : 5 banding pattems, mak ing them advantageous for (One smaller, obviously third instar larva: karyological analysis. Evety chromosome is divided 100:45:24:24: 10.) conditionall y into secti ons. 92 Hirvenoja & Michailova: Glyptotendipes aequalis • ENTOMOL. FENNICA Vol. 2

8

F

c BR

Fig . 3. 1st, lind, ll lrd and IVth chromosomes of Glyptotendipes aequa/is Kieffer. Original.

3.2. Karyotype. cunianus Edwards and G. gripekoveni Kieffer (Michailova 1979). The double bands in section 2 n = 8. 1st (AB), lind (CD), and Illrd (EF) 12/13- 13/14 are typical to this chromosome. chromosomes are metacentric, IVth (G) Section 9/10- 12 is similar to region 7/8 - 9/10 acrocentric. The centromere region is the darkest in G. cauJicola (Michailova 1979:29). Near the band in every chromosome. C. aequalis has no telomere 13/4- 14/15) there is a pale section cytologicall y obvious sex-differential segment or which looks li ke a constriction. Such a section chromosomes. ex ists in G. caulicola, G. glaucus, G. gripekoveni 1st (AB) chromosome (Fig. 3). This chromo­ and G. paripes, but has a different locali zation in some is divided into 15 sections. The centromere this chromosome. region is in section 5/6. Arm A: all investi gated lind (CD) chromosome (Fig. 3). Thi s chro­ indi viduals exhibit heterozygous inversion in this mosome contains 18 sections. The centromere arm. Arm B: in section 5/6- 6/7 is a constric­ region is localized in secti on 9/10. Arm C: at the tion. Such a constriction has been observed in telomere there is a group of dark bands. This the Ist chromosome of G. caulicola Kieffer, G. region is very often active. This arm is very glaucus Kieffer, G. paripes Edwards, G. man- similar in gross morphology to the arm C in G. ENTOMOL. FENNICA Vol. 2 • Hirvenoja & Michailova: Glyptotendipes aequalis 93

Illrd (EF) chromosome (Fig. 3). This is di­ vided into 8 sections. The centromere region is localized in secti on 3/4. Arm E: secti on 1/1 - 1/2 is similar to section 1 - 2/3 of G. gripekoveni. Section 1/1 -2/3 is similar to section 1 -2/3 of G. caulicola. The banding pattern in section 3/4 is characteristic in this arm of the chromosome. Arm F: Sequences of bands in sections 5 - 6!7 of G. aequalis differs by homozygous inversion from sequence of bands 4/5- 5/6 of G. gripekoveni. IVth (G) chromosome (Fig. 3). Thi s is the shortest of the complement, and is divided into 6 sections. There is a Balbiani ring and a nucleolus. The last one is in the telomere region. Very often in connection with the nucleolus there is a "Dark G knob" (Fig. 4c). The band pattern in section 6/7 is similar to section 4/5 of G. caulico/a. Section 1/ 2- 2/3 is a functionally active heterozygous puff G or different th ickness of a dark band (Fig. 4b).

4. Discussion.

4.2. Morphology

c The adults were originally determined as G. b aequalis with the aid of the key in "Lindner" (Goetghebuer 1937-1954). The comparison be­ tween the neotype (pupal skin) and the cast skins from Finland shows that some specimens of the latter material are quite similar. Most of the skins Fig . 4. Glyptotendipes aequalis Kieffer. - a: Hetero­ of Finnish origin , however, have stronger racket­ zygous inversions in lind chromosome. b: IVth chro­ mosome, different function al activity. c: Telomere shaped dorsal plates on tergites III-VI, but for heterochromatin "Dark knob" in IVth chromosome. the present there do not exist suffic ient Original. taxonomical differences to make it possible to reject the determination of the populations from Finland as G. aequa/is. caulico/a. Section 3/4-6/7 of G. aequalis differs Kieffer (1922:79) describes the male foreleg from section 1/1 - 3/4 of G. caulicola by an of G. aequalis as follows: " .. . , extremite du homozygous inversion. Arm D has a constriction metatarse et 2e article brievement barbus, poils in section ll/ 12, and distinct bands in section 12/ trois fo is aussi longs que Ia grosseur des arti­ 13 and 13/14. The band sequence in section 14/ cles, ... " 15- 16 is inverted in compari son with the band The latter agrees well with the specimens pattern in section 6/7- 8 of G. caulicola. Section described above as G. aequo/is. The genitalia of 14/1 5- 15/16 is similar to section 5/6- 6!7 of G. many species of G/yptotendipes are very similar. gripekoveni and section 8-9 of G. glaucus The structure of the male foreleg, the colouration (Michailova 1979). Heterozygous inversions were of the adult, and the characters of the pupa con­ observed in both arms in telomere regions in sidered together make the correct identification 3.22% (Fig. 4a). of the species possible. 94 Hirvenoja & Michailova: Glyptotendipes aequalis • ENTOMOL. FENNICA Vol. 2

Pinder & Reiss (1983, 1986) divide the known 1989, Miseiko et al. 197 1, Miseiko & Minsarino­ pupae and larvae of the genus Glyptotendipes into va 1974; Belyanina 1982). In all species 2n = 8; three groups: A, B and C, a division which in 1st, lind and IIIrd chromosomes are metacentric, some detail s differs from the older grouping of IVth acrocentric. Comparing the banding pattern Lenz ( 1954-1962). The new key characters bring, of the sali vary gland chromosomes of G. aequalis however, new problems: with other species of the genus Glyptotendipes The pupa of G. aequalis has an anterolateral indicates that G. aequalis is closely related to G. tubercle present on the sides of the first abdomi­ caulicola, G. glaucus, G. paripes and G. gripe­ nal segment as in species of the species group A, koveni. The band patterns of all chromosomes of but it is very small and moreover the features G. aequalis differ from those of G. sa linus and G. otherwise match group B better. The dark "mark" barbipes. This supports the idea (Michail ova on the tergite II in some individuals from Finl and 1979) that G. glaucus, G. gripekoveni, G. paripes, gives, however, an intermedi ate impression be­ G. caulicola and G. aequalis have evolved from tween the groups A and B. one ancestor in one direction while G. barbipes Key characters of group A also include lateral Staeger and G. salinus Michai lova have evolved denticles in the median tooth of the hypochilum in another directi on. of the larva of G. aequalis, although in many re­ G. gripekoveni, G. caulicola, G. glaucus and spects the larva agrees with those of group B. G. aequalis showed remarkable stability in some Furthermore in smaller details, for in stance the chromosome secti ons, vi z. 1st chromosome, sec­ front border of the paralabials and the apex of the tion 13/14 - 14 - 15 and 5/6 - 6/7; lind chromo­ maxillary sclerite, it agrees more with G. barbipes some, section 14/ 15- 15/ 16; IIIrd chromosome, sensu Pankratova ( 1983) than with G. caulicola section 1 - 1/2. These pattern s could be consid­ figured in the same paper. The presence of the ered as "basic patterns" in the sense of Wl.ilker ventral tubu li should li kewise not be included in ( 1980). Perhaps these common patterns ex isted the characteristics of Phytochironomus sensu in a hypothetical stem species. Lenz ( 1954-1962), to wh ich for in stance G. G. caulicola and G. aequalis are very similar aequalis and G. caulicola were referred by thi s and closely related, certain sequences of their author. chromosomes being identical. They differ only The karyological compari son (see above) by a simple inversion step (lind chromosome, shows G. aequalis and G. caulicola to be closely sections 3/4-6/7, 14/15 - 16), thu s indicating related species. Morphologicall y the re lations derivation from a common ancestor. They are between these species are not so clear, but there not far removed from the ancestral species and is no true di sagreement: if the features di scussed have a slow rate of divergency. above, di stingui shing G. aequalis from C. cauli­ cola , are not interpreted as synapomorphies, they can be di scarded in th e definition of species group 5. Ecological observations B. It seems therefore better to change the key characters for groups A and B in Pinder & Reiss Kieffer (1922:79), as well as Lenz ( 1954-1962: (1983, 1986), rather than to create a new group 176), mentions that the original specimens were for G. aequalis . (probably semi -) mining Alisma plantago L. Ac­ cording to Thienemann ( 1954:95, sub Phyto­ ) thi s species was reared in June 19 14 4.2. Karyotype from the stem of Alisma in Westfa len. The specimens described here from Yantaa, 2 Comparison of the ka ryotypic features of G. Finland, were found in an artific ial (about 15m ) glaucus with those of the other Glyptotendipes water reservoir in the garden while emptying the studied demonstrates simil arities with respect to fo rmer on 15 May 1988. The larvae had over­ th e diploid chromosome set (7 species of this wintered among the leaves of Populus tremula genus have been cytotaxonomicall y anal ysed; see L. and Betula spp. and in mud in a small amount Martin & Porter 1973, Michail ova 1979, 1987, of completely frozen water on the concrete bot- ENTOMOL. FENNICA Vol. 2 • Hirvenoja & Michai!ova: Glyptotendipes aequalis 95 tom. The water was somewhat polluted by the overwintered chironomid species in this pond. decomposing leaves of the trees and especiall y As is seen from the values for conductivity, by the excrement of the ducks that had been conditions in the forest pond (Riihimaki) were living on it during the previous summer. The not as eutrophicated as in the garden pond in water was strongly coloured by algae, and the Vantaa, as is clear also from the species combi­ populations of the Cladocera species were dense. nation. The latter might give a slightly lower The electrical conductivity of the water (on 15 saprobity index than that of the pond in Vantaa. May 1988) was 13.7 mS/m (y25), alkalinity 0 .50 The water most probabl y here too completely and the colour about 80 mg Pt/ 1. freezes in winter. There has been, however, more Using the saprobity tables of S l

Fittkau, E. J. & Reiss, F. 1978: Chironomidae. - In: lilies, VI . VI .) 197 1: The karyotype structure in natural popu­ J. (ed.), Limnofauna Europaea. 2 ed.: 404--420. lations of (Diptera, Chiro­ Goetghebuer, M. 1937-1954: Tendipediclae (Chirono­ nomidae). (In Russian with Engli sh summary) - midae). b) Subfamilie Tendipedinae (). Tsitologiya 13: 150 1- 1505. A. Die Imagines. - In: Lindner, E. (eel .): Die Fl iegen Mi seiko, G. N. & Minsarinova, B. Kh . (Mi·ICeFIKO , I . H . & cl er palarktischen Region 13c: 1-138. M HH cap HH OIJa, 6. X) 1974: The karyological structure Gouin, F. 1959: Morphology of the larval head of some of natural populations of Glyptotendipes glaucus and chironomidae (Diptera, ). - Smithsonian G. paripes (Chironomiclae). (In Russian with Engli sh Mi sc. Coli. 137: 175- 20 I. summary)- Tsitologiya 16: 893-897. - 1968: Morphologie, Histologie und Entwicklungs­ !CZN, Opinion 1147, 1980: Status for the purposes of type geschichte der Insekten und der Myriapoclen. - Fort­ fi xations of the remains of Chironomid larvae (Insecta, schr. Zoo!. 19: 194-282. Diptera) provided by Thienemann to Kieffer for the Hirvenoja, M. 1973: Revision cler Gattung Cricotopus van description of new species based on the adults reared der Wulp und ihrer Verwandten (Diptera, Chirono­ from those larvae.- Bull. Zoo!. Nomencl. 37( I): 11 - miclae). - Ann. Zoo!. Fennici 10:1 -363. 26. 1990: What is Gl yptotenclipes foliicola Kieffer (Diptera, Pankratova, V. Ya. (naH KpaTOilB , B. 51. ) 1983: [Larvae and Chironomidae)?- Nachr.Bl. Bayer. Entomol. 39:29- pupae of midges of the subfamil y Chironominae of the 30. USSR fa una (Diptera, Chtronomiclae =T endipeclidae)]. Imms, A. D. 1957: A general textbook of entomology, (In Russ ian)- Oprecl. Faune SSSR 134:1 - 295. including the anatomy, ph ysiology, development and Pinder, L. C. V. & Re iss, F. 1983: The larvae of classificati on of . 9. eel. - London. 886 pp. Chironominae (Diptera: Chironomiclae) of the Holarctic Kieffer, J. J. 1922: Chironomides nouveaux ou peu connus region. - Keys and diagnoses. - Entomol. Scand., de Ia region palearctique.- Ann. Soc. Sci. Bruxe lles Suppl. 19:293-435. 42( 1):7 1- 128. - 1986: The pupae of Chironomin ae (Diptera: Chi ro­ Laaksonen, R. 1970: Water quali ty in the water systems. A nomidae) of the Holarctic region. - Keys and diag­ study based on observat ions carried out by the water noses. - Entomol. Scand., Suppl. 28:299-456. pollution control authority 1962-1968. (In Finnish with Reiss, F. & Fittkau, E. J. 197 1: Taxonomie und Okologie English abstract) - Maa ja vesiteknillisia tutkimuksia europaisch verbreiteter -Arten (Chirono­ 17: 1- 132. miclae, Diptera). - Arch. Hyclrobiol. , Suppl. 40:75- Lenz, F. 1954-1962: Tenclipeclidae (Chironomidae). b) 200. Subfamilie Tenclipeclinae (Chironomi nae). B. Die Slaclecek, V. 1973: System of water quality from the bio­ Metamorphose cler Tendipeclinae. - In: Lindner, E. logical point of vi ew.- Ergebn . Limnol. 7 :1 -208. (eel.) : Die F li egen der palarktischen Region 13c: 139- Thienemann, A. 1954: Chironomus. Leben, Verbreitung 260. und wirtschaft li che Bedeutung der Chironomiden. ­ Martin , J. & Porter, D. 1973 : The salivary gland chromo­ Binnengewasser 20:1 - 834. somes of Glyptotendipes barbipes Staeg. (Diptera , Thienemann, A. & Zavrel, J. 19 16: Di e Metamorphose cl er Chironomidae) description of inversions and com­ Tanypinen. - Arch. Hydrobiol., Suppl. 2: 566-654. parison of Nearcti c and Palaearctic karyotypes. - Wiecl erholm, T. (ed.) 1983: Chironomiclae of the hoi arctic Stud. Naatur. Sci. 6:1 - 25. region . Keys and di agnoses. Part I. - Larvae. - Michai lova, P. 1979: Comparati ve karyological analysis Entomol. Scancl., Suppl. 19:1 -457. of the species of the genus Glyptotendipes Kieff. - 1986: Chironomiclae of the holarcti c region. Keys and (Diptera, Chironomiclae). - Caryologia 32:23-44. diagnoses. Part 2. - Pupae.- Entomol. Scand., Suppl. - 1987 : Comparative karyological studies of three species 28:1 - 482. of the genus Glyptotenclipes Kieff. (Diptera , Chiro­ - 1989: Chironomidae of the holarctic region. Keys and nomiclae) from Hungary and Bulgari a and Glypto­ diagnoses. Part 3 - Adult males. - Entomol. Scand. , tendipes salinus sp.n. from Bulgaria. - Foli a Bioi. Suppl. 34:1 - 532. 35 :43- 56. Wli lker, W. 1980: Basic patterns in chromosome evolution - 1989: The po lytene chromosomes and their significance in the genu s Chi ronomus (Diptera). - Zeitschr. Zoo!. for systematics and phylogeny of fam. Chironomidae, Syst. Evolutionforsch. 18: 11 2-11 3. Diptera. - Acta Zoo!. Fennica 186: 1- 107. Mi seiko, G. N., Minsarinova, B. Kh . & Kiknaclze, I. I. (M1·1ceF1K o, I . H ., M~o~HcapHH O I Ja, 6 . X . & I