Paratanytarsus oconnori, Moubayed-Breil & Ashe & Langton, 2012

Paratanytarsus oconnori View in CoL sp. nov.

(Figures 3, 6, 18, 22, 31, 34, 43-49)

Type material

Holotype. FRANCE: Hérault , Roque-Haute Domain, temporary deep pools, altitude 25-30 m, 21.v.1995 (1 pharate male), leg. J. Moubayed-Breil.

Paratypes: same locality as holotype, 23.iii.1997 (1 male adult and 1 female pupal exuviae), leg. J. Moubayed-Breil (in the collection of the senior author) ; same locality as holotype, basaltic lake at the cliff, altitude 15 m, 3.viii. 2001 (1 pharate male), leg. J. Moubayed-Breil (in the collection of P. H. Langton) . Holotype deposited in the collections of the National Museum of Ireland, Kildare Street , Dublin 2, Ireland .

Etymology

This species is named after our colleague Dr James P. O’Connor who recently retired from the National Museum of Ireland. He continues his research as Emeritus Entomologist in the museum and has to date to his credit 452 publications.

Diagnosis

In general, male adult characters of P. oconnori sp. nov. are closely similar to those of both P. natvigi Goetghebuer and P. dissimilis Johannsen. Distinguishing features for the male are: hairs present on anterior half of inner margin of the eyes; palp without sensilla clavata; 3 rd and 4 th palpomeres bearing a patch of minute macrotrichia (apically on segment 3, basally and apically on segment 4); thorax with minute scutal tubercle mediodorsally; sternapodeme with a swollen rectangular base, bearing two characteristic nozzle-like horns; anal point wider at base and progressively narrowed distally with rounded apex, in lateral view crests include 2 smooth peaks medially, proximal peak a little higher than the distal one; superior volsella subcircular to subrectangular with rounded posterior margin; digitus wider at base, abruptly tapering medially and ending with a rounded apex; median volsella short, slightly flattened distally, bent downwards, bearing only simple setae, posterior margin with 7-8 setae distinctly bent downwards; inferior volsella with basal half swollen, distal half less swollen, 5 straight and slender setae are present ventrally. Pupal exuviae characters key to those of P. bituberculatus but can be easily separated by the following features: thoracic suture with dense granulations only on posterior half, anterior half faintly granulose; scutal hump densely granulose and weakly domed; pearl row on the wing sheath short, not reaching the nose, median patch on tergite VI circular, bearing dark short to medium sized spines, 2 longer spines projecting posteriorly; genital sac of male covered with granulations as in P. bituberculatus and P. corsicanus , genital sac of female bearing granulations apically as in P. corsicanus .

Description

Adult male (n=3) ( Figures 43-49 View Figures 43-49 )

Large species. Total length 4.60-4.65 mm. Wing length (paratype) 2.30 mm. Dark brown to blackish in general. Head brown to dark brown with both eyes and pedicel blackish. Antenna (partial in holotype and paratype) with dark brown proximal segments. Palpomeres with dark brown segments. Thorax dark brown to blackish with black mesonotal stripes; mesosternum, scutellum and prosternum dark brown. Wing with both membrane and veins medium brown to dark brown. Legs dark brown to blackish. Abdomen and anal segment dark brown to blackish including both sternapodeme gonocoxite and gonostylus; only anal point and IX tergite are brown.

Head. AR not measurable (distal ¼ of last flagellomere missing). Eyes with bare ommatidia; hairs present on anterior half of inner margin of eyes ( Figure 43 View Figures 43-49 ). Clypeus with 23-24 setae located in 5 rows: 3, 3, 5-6, 6, 6. Palp with 5 segments; 3rd and 4th palpomeres ( Figure 44 View Figures 43-49 ) bearing patch of minute microtrichia: apically on segment 3, basally and apically on segment 4; last segment bearing minute sensilla chaetica apically, length (in µm) of palpomeres: 65, 88, 195, 205, 265. Frontal tubercles domed. Coronal triangle well marked bearing 4 small stout coronals. Temporal setae 11 including 2 inner and 9 outer verticals.

Thorax. Scutum bearing a minute scutal tubercle mediodorsally. Thoracic setae: antepronotals absent; 18 biserial acrostichals; 29-31 dorsocentrals often biserial; 0-1 prealars; 2 humerals; 8 uniserial scutellars.

Wing. All veins bearing setae except M; membrane covered with macrotrichia progressively becoming dense in distal ¼.

Legs. Pulvilli moderately developed, a little longer than claw; tibial comb of P 3 semicircular, maximum height 117 µm, maximum width 235 µm, composed of 19-20 short spines with rounded apex, of equal size (about 37-39 µm). Legs (P 1 and P 2 incomplete), measurements, in µm, as follows.

Hypopygium ( Figures 45-49 View Figures 43-49 ). Tergite IX subtriangular, ending with a quite wide V-like apical margin, bearing 8 setae situated mediodorsally on 2 weakly projecting tubercles, 10 setae present distally. Anal tergite with weakly developed tubercles medially, tergite bands very thick at base, becoming progressively thin from base to median part of anal tergite, parallel side medially and terminated with an outwards arc close to the base of the anal point crests. Anal point in dorsal view ( Figure 45 View Figures 43-49 ) 46 µm long, maximum width 35 µm at base, wider at base and narrowed distally with rounded apex; in lateral view ( Figure 46 View Figures 43-49 ) crests include 2 smooth peaks medially, proximal peak a little higher than the distal; 11-12 setae present basally including 6 laterally (3 on each side) and 4-5 ventrally. Sternapodeme with a swollen rectangular base, bearing two characteristic nozzle-like horns. Gonocoxite moderately swollen at ventral inner base, which bears dense microtrichia; inner basal margin bearing 4 stout setae, distal one is much stouter and longer than the preceding. Superior volsella ( Figures 47-48 View Figures 43-49 ) subcircular to subrectangular with rounded posterior margin; bearing 9 setae including 4 lateroventrally and 5 medially; digitus 48 µm long, maximum width 16 µm, wider at base, abruptly tapering medially and ending with a rounded apex. Median volsella ( Figures 45, 49 View Figures 43-49 ) 55-57 µm long, short and slightly flattened distally, bearing short to medium sized simple setae; posterior margin with 7-8 comb-like setae which are bent downwards. Inferior volsella ( Figure 45, 49 View Figures 43-49 ) 113 µm long, maximum width 41 µm, nearly divided longitudinally into two swollen lobes, basal half wider than distal half; in dorsal view, 12-13 curved stout setae near the apex; in ventral view, presence of 5 slender setae which are bent inwards. Gonostylus bearing two longitudinal rows of fine setae on inner margin (about 8-9 setae on each row).

Pupal exuviae (n = 3) (Figures 18, 22, 31, 34)

In general pupal exuviae of P. oconnori sp. nov. key close to those of other species in the bituberculatus- complex and to P. corsicanus . Main distinguishing characters are: thoracic suture (Figure 18) covered with dense granulations on posterior half, anterior half with faint granulations; scutal hump densely granulose and weakly domed; pearl row on the wing sheath short (Figure 22), not reaching the nose, extending to end of apical curvature of sheath; postero-median patch of spines on tergite III bearing about 15 long spines on each side, longest spines are inserted in a short rounded row, D5 taeniate; median patch on tergite VI (Figure 31) circular, bearing dark short to medium sized spines, 2 longer spines project posteriad; genital sac of male covered with granulations as in P. bituberculatus and P.corsicanus , genital sac of female (Figure 34) bearing granulations apically as in P. corsicanus .

Female adult and larva: Unknown.

Remarks

Very similar to both P. natvigi Goetghebuer and P. dissimilis , the male adult of P. oconnori sp. nov. is mainly distinguished from other related species on the basis of the legs and hypopygium: tibial comb of P 3 bearing more than 30 smooth spines in P. dissimilis , less than 20 smooth spines in P. oconnori ; anal point in dorsal view uniformly tapering apically in P. oconnori ( Figure 45 View Figures 43-49 ), parallel sided to drop-like in P. dissimilis ( Giłka 2011, Figures 123-124; Palmén 1960, Figure 2); in lateral view presence of 2 smooth peaks in P. oconnori (Figures. 6 and 46), presence of only 1 peak in P. sp. A (near dissimilis ) (Figure 7); superior volsella subcircular to subrectangular, bearing a rounded posterior margin; median volsella short, weakly flattened distally, bearing a different combination of simple setae on apical part; inferior volsella wider basally, constricted medially and moderately swollen distally, stout and curved setae present on dorsal apex, 5 slender setae present ventrally, which are bent inwards. Pupal exuviae of P. oconnori resembles those of P. bituberculatus ; it is separated from other related species by the following characters: hump on thoracic suture bearing dense granulation (Figure 18); granulations on thoracic suture extended anteriorly and posteriorly; median patch of spines on tergite VI dark and bearing 2 longer apical spines (Figure 31); only genital sac of female bearing granulations apically (Figure 34).

Ecology and geographical distribution

The species P. oconnori is only known from the type locality at Roque-Haute Domain, in southern France, where it was found in pools, lentic springs, a basaltic lake and slow flowing streams. Aquatic plants which occur in these habitats include Potamogeton, Scirpus, Ranunculus, Juncus, Alisma, Isoetes and Marsilea .

Paratanytarsus bituberculatus (Edwards 1929) (Figures 16, 20, 29, 33a, 33b, 42, 50-55)

Synonym: Paratanytarsus atrolineatus (Goetghebuer 1937) Imago : Reiss 1968; Langton & Pinder 2007; Giłka (2011).

Pupa: Thienemann 1951; Reiss & Säwedal 1981 (? pupa); Langton 1991; Langton & Visser (2003).

Female adult and larva: Undescribed.

Material examined

FRANCE: Aude, Aude River , epipotamal zone, altitude 50-60 m, 23.iv.1995 (1 male pupal exuviae), leg. J. Moubayed-Breil ; Normandy, Valmont River , 5.xi.1997 (1 male pupal exuviae), leg. J. Moubayed-Breil ; SPAIN: Andalucía, Sierra Almijara, Barranco de Luna , 21.iii.1999 (1 pharate male), leg. J.J. Casas (in collection of P.H. Langton) ; IRELAND: Co. Meath, Navan, Randalstown, N 855704, leg. D.A. Murray (specimen in National Museum of Ireland ) , 14.v.1996 (1 male adult); ITALY: Milan, Botanic Garden , 12.viii.1980 (2 pharate males and 1 male pupal exuviae), leg. B. Rossaro.

Diagnostic characters

Only the main distinguishing characters of both adult males and pupal exuviae are given.

Male adult. AR = 1.25-1.35. Palpomere 3 with only one apical patch of microtrichia, palpomere 4 with two patches of microtrichia, basally and distally ( Figure 50 View Figures 50-55 ); frontal tubercles 22-25 µm long ( Figure 51 View Figures 50-55 ), well projecting. Anal tergite ( Figure 52 View Figures 50-55 ) with 2 marked tubercles bearing 2 setae apically (Italian specimen has only one seta on each tubercle); anal point in lateral view with one median well projecting peak bearing a domed proximal margin ( Figure 53 View Figures 50-55 ); superior volsella semicircular with apical margin nearly straight and bearing a narrowed long digitus; median volsella ( Figures 42 View Figures 35-42 , 55 View Figures 50-55 ) short with numerous spatulate setae distally; inferior volsella ( Figures 54-55 View Figures 50-55 ) strongly widened and swollen at base, constricted distally, bearing a triangular lobe medially, setiferous ventral lobe well defined and bearing 4 stout setae ( Figure 55 View Figures 50-55 ).

Pupal exuviae. General shape pattern of abdomen as in Figure 23. Granulations on thoracic suture absent (Figure 16), only scutal hump is faintly granulose; thoracic horn absent; pearl row on wing sheaths short (Figure 20). Pair of spine patches on posterior half of tergite III composed of about 13-15 spines inserted nearly in a L-shaped line, D5 bristle-like or taeniate (taeniate in Italian specimens); point patch on tergite VI (Figure 29) subcircular, not darkened, composed of faint points. Anal lobe (Figures 33a, 33b) with rounded apical margin, genital sac of male with dense granulations including apical part, genital sac of female without granulations.

Remarks and geographical distribution

Reiss & Säwedal (1981: 94) in a key indicate that the pupal exuviae of P. bituberculatus has a thoracic horn but in Langtom (1991: 327, Plate 132b) the thoracic horn is regarded as absent in this species and the latter viewpoint is accepted here.

The pupal exuviae of P. bituberculatus resemble closely those of P. corsicanus , P. curvispinus and P. oconnori . Main separating characters for P. bituberculatus are: thoracic suture (Figure 16) lacking granulations on both anterior and posterior half; thoracic horn absent; scutal hump faintly granulose and not domed (Figure 16); pearl row on the wing sheath (Figure 20), not reaching the nose; median patch on tergite VI (Figure 29) circular, bearing faint to weakly darkened points; genital sac of male covered with granulations (Figure 33a), genital sac of female (Figure 33b) lacking granulations apically. However, some variation in the male imagines and the pupal exuviae P. bituberculatus from Italy include: tubercles on anal tergite of male bearing only a single seta; thoracic suture with faint granulations on anterior half; D5 on tergite III taeniate.

DISCUSSION

Associated material composed of pupal exuviae and pharate males belonging to the bituberculatus -complex (with pupae that key to P. bituberculatus in Langton 1991 and Langton & Visser 2003) described here demonstrates that there is a risk of misidentifying pupal exuviae similar to P. bituberculatus as P. bituberculatus . Consequently, we believe that populations of P. bituberculatus from Europe and neighboring regions include about five new species, which key to P. bituberculatus as pupal exuviae. It appears that true P. bituberculatus is more a northern element than a widespread species throughout the Palaearctic Region. Therefore, the presence or absence of P. bituberculatus from some subregions in Europe and the Mediterranean Region needs to be reviewed and confirmed on the basis of pharate males or adult males, in particular from southern Europe (including Portugal, Spain, France, Italy, Greece) and the Near East. Details on both the taxonomy and biogeography of the bituberculatus -complex are given in a separate paper in preparation.

In the bituberculatus -complex, based on the morphology of both male adult and pupal exuviae, there appear to be several species groups, one centering on P. bituberculatus with several close but distinct sister species and another which includes only P. corsicanus sp. nov. Further investigations throughout Europe, North Africa and the Near East are necessary to determine if imaginal and pupal features support each other and enable the creation of species groups within the entire genus.

Paratanytarsus . bituberculatus has been previously accepted as having a widespread distribution throughout the Palaearctic Region including southern Europe, the Near East and North Africa. Our investigations show that several species morphologically similar to P. bituberculatus , occur in the Mediterranean Basin region. Two of these, P. curvispinus and P. oconnori , which we consider members of the bituberculatuscomplex, are described in this paper. In the light of these new discoveries the geographical distribution of P. bituberculatus needs to be reviewed by focusing especially on examining pharate males and adult males. Taxonomic information given in this paper provides additional characters for keying any further related species which fall into the bituberculatus -complex. Additional future work needs to be done on both the taxonomic composition and geographical distribution of all members of the bituberculatus -complex.

The biological quality of both water and sediment in wetland areas and lower reaches of coastal rivers is seriously damaged by the impact of organic and chemical pollution in many countries around the Mediterranean Basin. Furthermore, despite extensive investigations during the last three decades in southern France and Lebanon, only a few pharate males and pupal exuviae belonging to the bituberculatus -complex have been obtained.

Coastal aquatic ecosystems including coastal rivers in southern and western France (Mediterranean and Atlantic coastal areas) have been heavily affected by toxic chemical pollutants such as HAP’s and PCB’s. The latter products combined with the anti-mosquito solution BTI and the past use of pesticides such as Fipronil and Lindane (which remain in the sediments and take a long time to breakdown) can lower the chironomid diversity in coastal habitats by 30 to 50 % (Moubayed-Breil, personal observation).

Key to related known adult males and pupal exuviae

Adult male

As in most Tanytarsini View in CoL , variation in the superior and median volsella, and the shape of the anal point, provide good distinguishing characters which are often used as main key features for the identification of species. Details of the median volsella in particular, for accurate observation, are more clearly visible when the hypopygium is temporally mounted ventral side uppermost or if the median volsella is dissected from the rest of the hypopygium.

In the following keys, for the identification of adult males and pupal exuviae, only those species, known from Europe and the Near East, belonging to the bituberculatus- complex ( P. bituberculatus , P. curvispinus and P. oconnori ) and P. corsicanus are fully keyed.

Adult males

1. Anal tergite without tubercles medially; Anal tergite bands terminating abruptly, well before base of anal point ( Figure 9 View Figures 9 ); Anal point (in dorsal view) broad basally, constricted medially, expanded apically to drop-like apex ( Figure 9 View Figures 9 ), (in lateral view) triangular (Figure 4); Superior volsella suboval (Figures 10-11); Digitus long and finger-like, equally broad, with rounded apex (Figures 10-11); Median volsella short, triangular bearing only long bristlelike setae (Figure 12); Inferior volsella wider at base, straight and uniformly tapering ( Figures 9 View Figures 9 , 13), bearing 2 widely spaced setae ventrally ( Figure 9 View Figures 9 ), setiferous ventral lobe absent ....................................….. P. corsicanus View in CoL sp. nov.

- Anal tergite with or without tubercles medially; Anal tergite bands terminating near base of anal point ( Figures 38 View Figures 35-42 , 45 View Figures 43-49 , 52 View Figures 50-55 ); Anal point (in dorsal view) evenly tapered to rounded apex ( Figures 38 View Figures 35-42 , 45 View Figures 43-49 ), (in lateral view) not triangular (Figures 5-6, 39, 46, 53); Superior volsella subcircular, subtriangular or subrectangular; Combination of digitus, median volsella and inferior volsella not as described above ...................................................................... 2

2. Anal tergite without tubercles medially ............................ other Paratanytarsus species [Not keyed]

- Anal tergite with tubercles medially bituberculatuscomplex ............................................................................... ...... 3

3. Median volsella with simple setae ( Figures 45, 49 View Figures 43-49 ); Digitus very broad basally, abruptly constricted medially, and evenly continuing to rounded apex ( Figures 47-48 View Figures 43-49 ); Anal tergite with weakly developed tubercles medially ( Figure 45 View Figures 43-49 ) ............................................................ P. oconnori sp. nov.

– Median volsella with spatulate lamellae ( Figures 41-42 View Figures 35-42 ); Digitus not as above, with pointed apex ( Figure 40 View Figures 35-42 ); Anal tergite with very distinct tubercles medially ( Figures 38 View Figures 35-42 , 52 View Figures 50-55 ) .............................................................................................. 4

4. Inferior volsella straight, with a triangular lobe projecting beyond the inner margin ( Figure 54 View Figures 50-55 , Reiss & Säwedal 1981, Figure 9 View Figures 9 ), ventrally bearing 4 setae on a distinct subapical lobe ( Figure 55 View Figures 50-55 ); Tubercles on anal tergite smaller, usually with 2 setae (rarely 1 seta) ( Figure 52 View Figures 50-55 ); Median volsella, short and broad, not tapering, with simple setae and 10 to 11 spatulate setae distally, simple setae reaching to about one-half length of inferior volsella ( Figures 42 View Figures 35-42 , 55 View Figures 50-55 ); Palpomeres 3 (apically) and 4 (basally and apically) with patches of microtrichia ( Figure 50 View Figures 50-55 ) ........................................... P. bituberculatus (Edwards 1929)

– Inferior volsella bent inwards apically, with a distinctive tuft of dark setae on inner margin ( Figure 38 View Figures 35-42 ), ventrally bearing 3 setae but without a distinctive subapical lobe ( Figure 38 View Figures 35-42 ); Tubercles on anal tergite larger, with 4 or 5 setae ( Figure 38 View Figures 35-42 ); Median volsella triangular, tapering towards apex, with both simple setae and about 4 to 6 spatulate setae distally ( Figures 38, 41 View Figures 35-42 ), simple setae almost reaching tip of inferior volsella; Palpomeres 3 and 4 lacking patches of microtrichia ( Figure 37 View Figures 35-42 ) .............................................................. P. curvispinus sp. nov.

Pupal exuviae

1. Thoracic horn present; Pearl row on wing sheath present or absent, when present, pearl row is usually short, stopping well before nose; Hump on suture of thorax present or

-

2.

–

3.

-

4.

-

absent; when hump is present, granulations are dense ........................… other Paratanytarsus species [Not keyed] Thoracic horn absent; Pearl row present, either very long or short (Figures 19-22); Hump on thoracic suture present, usually distinct (Figures 15, 17-18), if indistinct, hump is defined by a cluster of granulations (Figure 16) .................................................................................................… 2

Pearl row very long, almost reaching the nose (Figure 19); Median patch on tergite VI consists of short spines (Figure 27); Thoracic hump large, smoothly rounded, domed, with minute faint granulations, only anterior part of suture with dense granulations (Figure 15); D5 on tergite III taeniate (Figure 24); Apical margin of anal lobe straight (Figure 32) ................................................................ P. corsicanus sp. nov. Pearl row short, stops well before the nose (Figures 20-22); Median patch on tergite VI with dark points or bearing faint short spines (Figures 29-31); Thoracic hump and suture granulations not as described above (Figures 16-18); D5 on tergite III taeniate or bristle-like; Apical margin of anal lobe rounded (Figure 33) (bituberculatuscomplex)......................................................................................3

Thoracicsuturewithoutanteriorandposteriorgranulations, hump weakly domed, indistinct, indicated by a cluster of faint granulations (Figure 16)

........................................... P. bituberculatus (Edwards 1929) Thoracic suture with anterior and posterior granulations, hump distinct, pointed or strongly granulose (Figures 17, 18) .............................................................................................. 4

Hump on thoracic suture bent backwards and pointed apically (Figure 17); Median patch on tergite VI composed of dark points (Figure 30); D5 on tergite III bristle-like .............................................................. P. curvispinus sp. nov. Hump on thoracic suture almost indistinct, strongly granulose (Figure 18); Median patch on tergite VI composed of dark spines (Figure 31); D5 on tergite III taeniate .................................................... P. oconnori sp. nov.

ACKNOWLEDGMENTS

The authors are grateful to Dr J. Giudicelli, Dr B. Roché (†), Dr A. Orsini and Dr C. Mori for their hospitality in Corté City who provided facilities to the senior author in Corsica at the meeting of the “French Association of Limnology” in July 1997. We are also indebted to Prof. B. Rossaro for providing specimens of P. bituberculatus from Italy.

REFERENCES

Delettre YR. 2001. An annotated checklist of Chironomidae (Diptera) trapped in Brittany (France) since 1975. Annales de Limnologie 37: 143-149.

Ekrem T. 2001. A review of Afrotropical Tanytarsus van der Wulp (Diptera: Chironomidae). Tijdschrift voor Entomologie 144: 5-40.

Ekrem T. 2002. A review of selected south- and east Asian Tanytarsus v. d. Wulp (Diptera: Chironomidae). Hydrobiologia 474: 1-39.

Furon R. 1950. Les grandes lignes de la Paléogéographie de la Méditerranée (Tertiaire et Quaternaire). Vie et Milieu 1: 131- 162.

Furon R. 1972. Eléments de Paléoclimatologie. Vuibert, Paris, 210 pp, 24 figs.

Giłka W. 2009. Order Diptera, family Chironomidae Tribe Tanytarsini. Arthropod Fauna of the UAE 2: 667-682.

Giłka W. 2011. Ochotkowate - Chironomidae, plemię: Tanytarsini, postaci dorosłe, samce. Klucze do oznaczania owadów Polski. [Non-biting midges - Chironomidae, tribe Tanytarsini, adult males. Keys for the Identification of Polish Insects]. Nr 177 serii kluczy. Część XXVIII, Muchówki - Diptera, zeszyt 14 b. Polskie Towarzystwo Entomologiczne. Biologica Silesiae, Wrocław, 95 str. [in Polish]

Giłka W, Paasivirta L. 2008. On the systematics of the tribe Tanytarsini (Diptera: Chironomidae) – three new species from Finland. Entomologica Fennica 19: 41-48.

Giudicelli J. 1975. Analyse de la l’endémisme dans la faune des eaux courantes de la Corse. Ecologia Mediterranea 1: 133-147.

Goetghebuer M. 1921. Chironomides de Belgique et spécialement de la zone des Flandres. Mémoires du Musée Royal d’Histoire Naturelle de Belgique 8 (Fascicule 4, Mémoire 31): 1-208 + [2] p.

Johannsen OA. 1905. Aquatic nematocerous Diptera II. Chironomidae. Pp 76 -330, 333-352, pls 16-37. In Needham, J. G., Morton, K. J. and Johannsen, O. A. (eds) May flies and midges of New York. Bulletin of the New York State Museum 86: 1-352.

Kieffer JJ. 1909. Diagnoses de nouveaux chironomides d’Allemagne. Bulletin de la Société d’Histoire Naturelle de Metz 26: 37-56.

Langton PH. 1991. A key to pupal exuviae of West Palaearctic Chironomidae. Privately published. Huntingdon, England, 386 pp.

Langton PH, Pinder LCV. 2007. Keys to the adult male Chironomidae of Britain and Ireland. Volume 1 (Pp: 1-239) and volume 2 (Pp: 1-168). Freshwater Biological Association, Scientific Publication No. 64.

Langton PH, Visser H. 2003. Chironomidae exuviae. A key to pupal exuviae of the west Palaearctic Region. Amsterdam: Biodiversity Center of ETI. CD-Rom.

Langton PH, Cranston PS, Armitage P. 1988. The parthenogenetic midge of water supply systems, Paratanytarsus grimmii (Schneider) (Diptera: Chironomidae). Bulletin of Entomological Research 78: 317-328.

Laville H, Serra-Tosio B. 1996. Additions et corrections à l’inventaire des Chironomidés (Diptera) de France depuis 1990. Annales de Limnologie 32: 115-121.

Laville H, Langton PH. 2002. The lotic Chironomidae (Diptera) of Corsica (France). Annales de Limnologie 38 (1): 53-64.

Moubayed-Breil J. 2007. Non-biting midges from Continental France: new records, faunal and biogeographical outline (Diptera, Chironomidae). Ephemera 9 (1): 17-32.

Palmén E. 1960. Paratanytarsus -Arten (Dipt., Chironomidae) aus dem β-mesohalinen und oligohalinen Brackwasser der Finnischen Meerbusens. Annales Entomologici Fennici 26: 280-291.

Reiss F. 1968. Ökologische und systematische Untersuchungen an Chironomiden (Diptera) des Bodensees. Ein Beitrag zur lakustrischen Chironomidenfauna des nördlichen Alpenvorlandes. Archiv für Hydrobiologie 64 (2/ 3): 176-323.

Reiss F, Säwedal L. 1981. Keys to males and pupae of the Palaearctic (excl. Japan) Paratanytarsus Thienemann & Bause, 1913, n. comb., with descriptions of three new species (Diptera: Chironomidae). Entomologica Scandinavica Supplement 15: 73-104.

Saether OA. 1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae). Entomologica Scandinavica Supplement 14: 1-51.

Saether OA, Spies M. 2011. Chironomidae. In: Fauna Europaea Service (ed.). Fauna Europaea, internet database at http://www. faunaeur.org [visited April 2011].

Säwedal L, Langton PH. 1977. Redescription of Paratanytarsus tenellulus (Goetghebuer, 1921) (Diptera: Chironomidae). Entomologica Scandinavica 8: 167-171.

Shilova AI. 1976. Khironomidy Rybinskogo Vodokhranilishcha. Izvestiya Academii Nauk SSSR 1976, 249 pp.

SchneiderA. 1885. ChironomusGrimmii undseineParthenogenesis. Zoologische Beiträge 1: 301-302.

Stur E, Ekrem T. 2006. A revision of West Palaearctic species of the Micropsectra atrofasciata species group (Diptera: Chironomidae). Zoological Journal of the Linnean Society 146: 165-225.

Thienemann A. 1951. Tanytarsus-Studien II. Die Subsectio Paratanytarsus. Auf Grund der nachgelassenen Papiere Friedrich Wilh. Carl Krüger’s. Archiv für Hydrobiologie Supplement 18: 595-632.