MARTIN SPIES
Munich, Germany
A CONTRIBUTION TO THE KNOWLEDGE OF HOLARCTIC PARACHIRONOMUS LENZ (DIPTERA: CHIRONOMIDAE), WITH TWO NEW SPECIES AND A PROVISIONAL KEY TO NEARCTIC ADULT MALES
Spies, M. 2000. A contribution to the knowledge of Holarctic Parachironomus Lenz (Diptera: Chironomidae), with two new species and a provisional key to Nearctic adult males. – Tijd- schrift voor Entomologie: 143: 125-143, figs. 1-13, tables 1-5. [ISSN 0040-7496]. Published 5 July 2000. The Palaearctic Parachironomus monochromus (van der Wulp, 1874) and the Holarctic P. tenuicaudatus (Malloch, 1915) are redescribed in the male adult, pupal and larval stages. North American records of monochromus are shown to be misidentifications, and two new Nearctic species are described: P. hazelriggi, sp. n. (adult male, pupa, larva), and P. gillespieae, sp. n. (adult male). P. pediformis Lenz, 1951 is a new junior synonym of P. tenuicaudatus. A provisional key is given for Nearctic adult males of the genus, including several unnamed species. P. digitalis (Edwards, 1929), guarani Spies, Fittkau & Reiss, 1994, and parilis (Walker, 1856) are newly recorded from the Nearctic region. Correspondence: Martin Spies, Schrämelstr. 151, D-81247 München, Germany. E-mail: [email protected] Key words. – Diptera; Chironomidae; Parachironomus; Holarctic; taxonomy; description; key.
During studies on nuisance Chironomidae in the autapomorphies or simple character sets have been greater Los Angeles urban area (USA, California), identified distinguishing them from all their con- adult males of the genus Parachironomus Lenz were geners. Thus, the selection of species for the present encountered, which according to the North Ameri- work perhaps does not form a monophyletic group- can reference literature represented the originally Eu- ing, but is rather a result of the author’s interpretation ropean species monochromus (van der Wulp, 1874). of the available information. More associations of still Simultaneously collected pupal exuviae, however, did undiscovered or poorly known Parachironomus imma- not match the descriptions of monochromus, but ap- ture stages, and the integration of data on all stages are peared to be P. tenuicaudatus (Malloch, 1915). Cer- required for an authoritative Holarctic revision. tain transatlantic differences in monochromus male hypopygial morphology – first noticed by Goetghe- MATERIAL AND METHODS buer (1951) – were also apparent. A wide array of material and references has been The species analyzed here have in the course of 140 studied to resolve these inconsistencies and identify years been mentioned in many publications. For prac- the southern Californian species, extending the origi- tical reasons it is not possible to evaluate all that mate- nal problem. The North American Parachironomus rial. Analogously, the historic listings under each fauna clearly is not as well described as assumed, and species heading below do not contain all known refer- a full review of Holarctic scope is required. As a first ences, but instead only those essential to finding nec- step, the present study attempts to resolve a limited essary information. For example, since most authors set of outstanding problems, and identifies others. treating North American material followed Townes’ The species treated here are presently inseparable as (1945) concepts, such contributions are only cited in larvae (3 of 4 known), and differ little to moderately in special cases. Unverified records are indicated by a ‘?’. pupal and adult male morphologies (P. monochromus Relevant details are discussed in ‘Notes’ sections fol- more noticeably in both stages). However, no shared lowing the historic listings.
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Wherever possible, standard morphological termi- INHS Illinois Natural History Survey (Cham- nology is followed (combined from Sæther 1980, paign) Cranston & Reiss 1983, Coffman 1986, Pinder & ISNB Institut Royal des Sciences Naturelles de Reiss 1986, and Oliver & Dillon 1989). Some special Belgique (Brussels, Belgium) adult characters are evaluated according to Spies et al. JES James E. Sublette (Tucson, Arizona) (1994: 62). Pupal setae showing at least partial lamel- JHE John H. Epler (Crawfordville, Florida) lar widening are called ‘semi-taeniate’ or ‘taeniate’ af- MJB Michael J. Bolton (Columbus, Ohio) ter a suggestion by Langton (1994). The whorl of OCVCD Orange County Vector Control District clear spines in the ventral, postero-lateral corners of (Garden Grove, California) some abdominal segments on the pupa is called ‘vor- PHL Peter H. Langton (Coleraine, Northern tex’ (plural ‘vortices’) after Langton (1999). Also see Ireland) the latter reference for definitions of the pupal WFBM W. F. Barr Entomological Museum, Uni- paratergites and parasternites. versity of Idaho (Moscow) Two new terms are here introduced to facilitate de- USNM United States National Museum of Natural scription of group- or species-diagnostic character History (Washington, D.C.) states. ‘Caudal’ setae of the adult male anal tergite are ZMAN Zoölogisch Museum, Universiteit van Am- those grouped around the origin of the anal point sterdam (Netherlands) and/or lining a proximal stretch of the latter. ‘Portal se- ZSM Zoologische Staatssammlung München tae’ identifies a bilaterally paired set often clearly distin- (Germany) guishable anterior to the ventralmost caudal setae, on a ledge or prominence(s) posteroventral to (‘at the en- SYSTEMATIC PART trance of’) the inner genital area (see figs. 1-5). Outside of the present study, examples for portal setae and their Parachironomus monochromus (van der Wulp) taxonomic value (see Spies et al. 1994) are Parachirono- (figs. 1, 3) mus manaos Spies, Fittkau & Reiss, 1994, P. puberulus (Edwards, 1931), and P. vistosus Paggi, 1979. Chironomus unicolor van der Wulp, 1859a: 5, 1859b: 5, For the adult male stage, table 5 allows direct com- 1859c: 32, 1859d: 162 (primary homonym of C. unicolor parisons of non-hypopygial characters. The text de- Walker, 1848; adult male). Chironomus monochromus van der Wulp, 1874: 129 (re- scriptions give features not listed in the table, but also placement name for C. unicolor van der Wulp, 1859 nec repeat those warranting special mention. Walker, 1848). Meristic data are generally presented in the format: Chironomus (Cryptochironomus) claviforceps Edwards, 1929: value range (number of values). If the value distribu- 389 (adult male; see below under ‘Notes’, item 1). tion is significantly skewed, n is preceded by the me- Tendipes (Parachironomus) monochromus: Kruseman, 1933: 192 (synonymization of claviforceps Edwards; adult dian value: x-y (M; n). male). Data on bilaterally paired and discrete structures Tendipes (Cryptochironomus) monochromus: Goetghebuer, (e.g. number of thoracic dorsocentral setae) are given 1937-1954: 46 (adult male). for one body side only. Structures interpretable as ? ‘Parachironomus monochromus v. d. Wulp. var. Goetgh.’: fused pairs without a distinct gap are evaluated in full Lenz, 1938: 712 (larva, pupa; see below under ‘Notes’, (e.g. caudal setae around anal point origin). item 2). Parachironomus monochromus: Lehmann, 1970: 146 (adult male); ? Rodova, 1978: 99 (adult female); Albu, 1980: Abbreviations of life stages: ex = exuviae, L = larva, 131 (adult male; see below under ‘Notes’, item 4); Lang- P = pupa, ph = pharate. ton, 1991: 274 (pupa); Kobayashi & Suzuki, 1999: 82 Abbreviations of names and institutions (USA un- (adult male; see below under ‘Notes’, item 4). less otherwise stated): AEI American Entomological Institute (Gaines- Notes on nomenclature and identification ville, Florida) 1. Goetghebuer (1921) was the first subsequent au- ANSP Academy of Natural Sciences of Philadel- thor to apply the name monochromus van der Wulp to phia (Pennsylvania) a morphological concept. He used it for specimens BAC Broughton A. Caldwell (Lawrenceville, clearly belonging to P. tenuicaudatus (Malloch), and Georgia) for that reason later (1928: 81, 155) called baciliger BMNH The Natural History Museum (London, Kieffer a junior synonym. Edwards (1929) followed UK) Goetghebuer’s opinion, and separately described his FAMU Florida A&M University (Tallahassee) new claviforceps. Kruseman (1933) verified that van GLACVCD Greater Los Angeles County Vector der Wulp’s type of monochromus was lost, and reinter- Control District (Santa Fe Springs, Califor- preted the species because Goetghebuer’s specimens nia) had the legs unicolorous, not partially darkened as
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van der Wulp had described them. Thus, claviforceps Pex+(; Bavaria, pond nr. Wolfsegg E of Landshut, 14.v.1983, F. Reiss (ZSM). 2(; Bavaria, Inn river at Perach, Edwards, instead of baciliger Kieffer, became a junior 9.vi.1977, leg. Utschick (ZSM). – NETHERLANDS: 3(; synonym of monochromus. Goetghebuer (1937-1954) Voorst, ‘Yselarm’, 18.viii.1930, M. C. Jansen & G. Kruse- accepted Kruseman’s solution which has been the ba- man (ZMAN, author’s coll.). – UNITED KINGDOM: 1(; Eng- sis of the Palaearctic concept for monochromus van der land, Hertfordshire, Letchworth, vi.1918, F. W. Edwards Wulp ever since. (syntype claviforceps Edwards; at BMNH). 1(; England, The author has slide-mounted three males from the Cambridgeshire, March, fishpond in Maple Grove, 4.v.1973, G. W. Simpson (PHL). 1 LexPex; as previous exc. Kruseman collection at ZMAN, as well as the male 5.vi.1973 (PHL). 1 Pex; as previous exc. at Station Rd., amongst the two syntypes of claviforceps at BMNH (the 17.v.1973, W. M. H. Peace (PHL). other syntype is a female), and finds these to be con- specific. It would be preferable if the no longer type- Adult male (also see table 5) based monochromus had been avoided in favour of Colour. – Green without distinct thoracic mark- claviforceps. However, an action to this effect today ings. Fore fe and most of ti greenish to yellowish, apex would overturn long-standing and frequent practice, of ti brown; fore ta1 mostly whitish, apex brown; ta2- and is therefore not in the interest of taxonomic sta- ta5 brown; mid and hind legs darkened only apically. bility. A neotype designation for monochromus is not Head. – Temporal setae in laterally single, medial- considered as necessary as required by the current ly double to triple rows. Frontal tubercles present in Code of nomenclature (International Commission on only 1 of 8 specimens examined. Zoological Nomenclature 1999: Article 75). Thorax. – Dorsocentral setae in at most briefly 2. Lenz (1938) briefly described larval and pupal double rows. Scutellar setae in 2 rows, the posterior characters under the name ‘monochromus v. d. Wulp. generally more and larger. var. Goetgh.’, from material reportedly reared in Aus- Wing. – M1+2 bare. tria (Danube oxbow nr. Vienna, leg. J. Vornatscher). Legs (table 1). – Mid and hind tibiae each with the Some immature specimens found at ZSM are possibly combs fused and the spurs subequal to slightly un- from that material, but they are incompletely labeled even, about as long as their respective comb height. and in poor condition. The larvae are presently insep- Hypopygium (figs. 1, 3). – Laterosternite IX with arable from those of other Parachironomus species. 1-3 setae. Anal tergite bands of T- to Y-type, some- The pupae key (in Langton 1991) not to monochro- times separate at the juncture, longitudinal part occa- mus, but to the unassociated provisional taxon ‘Pe 2’. sionally split posteriorly to embrace field of setae. 19- The adults, which had been determined by Goetghe- 37 caudal setae, including 7-14 stronger and longer buer (Lenz 1938: 713), have not been recovered. ones in a dorsal cluster. Anal point with basal, mi- Thus, the Vornatscher material can not be placed to crotrichiose part not distinctly delimited from the ter- species at this time. No information is available on minally subconical anal tergite; distal, bare part in the reason for Lenz’ (1938) term ‘var. Goetgh.’, but dorsal aspect narrow, sometimes slightly widened api- in a later publication (1954-1962: 203) Lenz listed cally, strongly angled to ventral (fig. 3). Portal setae 2, the same material and his own earlier treatment under widely separate. Superior volsella straight to slightly the modified name ‘monochromus v. d. Wulp nec curving, 85-110 µm long, distally widened to medi- Goetgh.’ (see above ‘Note 1.’). On present evidence it an, without conspicuous apicolateral projection; me- is considered unwarranted to formally recognize dian setal pit always smaller than distal pit and often Lenz’ variety as a separate taxon on subspecific or positioned a little proximal. Inferior volsella rounded- higher level. ly rectangular, or with low projection to posterior; 3. For ‘Tendipes monochromus (Wulp)’ of Townes (1945) and subsequent North American authors, see below under Parachironomus hazelriggi sp. n., ‘Notes’. 4. The meristic data given by Albu (1980) and Table 1. P. monochromus, adult male: lengths of leg seg- ments in µm (largest / smallest (= syntype claviforceps) com- Kobayashi & Suzuki (1999) are for the most part in plete specimen). good agreement with those found in the present study. Albu’s (1980: 131) palpomere length values P1 P2 P3 are much higher when related to body (wing) size. For both the Romanian and Japanese specimens fe 890 / 660 940 / 720 1010 / 780 slightly lower wing lengths are reported. ti 760 / 580 860 / 640 1130 / 860 ta1 1150 / 950 470 / 370 740 / 560
ta2 580 / 460 250 / 200 370 / 300 Material examined. – GERMANY:1(; Baden-Württem- ta 440 / 360 180 / 130 290 / 240 berg, Schussen river at rail bridge nr. Moos/Langenargen, 3 ta4 300 / 220 110 / 80 170 / 150 27.v.1963, F. Reiss (ZSM). 1(; as previous exc. Stockacher ta5 130 / 100 80 / 60 90 / 80 Aach nr. Ludwigshafen (Bodensee), 4.vi.1963 (ZSM). 1(, 1
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ps
ps
1 2
Figs. 1, 2. Parachironomus, adult male hypopygia (dorsal). – 1, P. monochromus; 2, P. tenuicaudatus. Scale: 100 µm; ps = po- sitions of portal setae. microtrichia not elongate. Gonostylus mostly slender, some with few, small surface denticles. Prealar tuber- slightly curving in dorsal and lateral aspects, with cle with narrow, low anterior, and broad, higher pos- weak near-basal expansion bearing 3-5 lateral and terior sublobe, overall height < length. ventral setae isolated by an adjacent bare section, and Abdomen. – Dorsal armament: Segment II bare with distinct widening to dorsal beginning beyond except for hook row, the latter covering 25-30% of midlength and peaking in distal ⅓; 4-6 distalmedian segment width, consisting of about 30-35 hooks ap- setae. parently without dorsal accessory points; III with sha- green limited to the usual posteromedian patches; VII Adult female with anterolateral shagreen only. Ventral armament: I Not studied. Specimens not seen for the present with extensive shagreen from anterolateral to postero- study have been described by Rodova (1978). median; II: parasternite covered with conspicuous shagreen, sternite with posterior transverse band in- Pupa cluding some elongate points (up to 50 µm), band Similar to P. hazelriggi sp. n., except as follows. with lateral projections to anterior, not connected to Length. – Exuviae 4.4-5.2 mm (Langton 1991). anterior transverse band or triangular field of weak Head. – Frontal seta 15-25 µm long, cephalic tu- shagreen (Langton 1991: fig. 112e, parasternite sha- bercle longer overall, its distal cone may appear sube- green not shown); III at most with little, anterolater- qual in flattened exuviae mounts. al shagreen; IV with fully developed vortex; V, VI Thorax. – Thoracic horn of numerous branches, with small point fields between posterolateral adhe-
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sion marks and seta L4; VIII with 2-4 anal comb East Palaearctic, the species has been recorded from spines (up to 20 µm long). Lateral setae: V-VII: 4 the Russian Primorye Territory (Makarchenko et al. (taeniate); VIII: 5 (taeniate, the posterior two usually 1999), and Japan (Kondo & Hamashima 1985). next to each other). Anal lobe with 1 postero-dorsal On present evidence, P. monochromus does not oc- taenia and about 35-50 in fringe, the latter in single cur outside of the Palaearctic region. All new world row, setae not more densely set toward fringe ends. records instead belong to one of the two species new- ly described below, or may represent additional, un- Larva (from one Lex individually associated with recognized ones. Listings from Mexico after Spies & Pex+() Reiss (1996) must be changed to P. hazelriggi, sp. n. Similar to P. hazelriggi sp. n., except as follows. P. monochromus van der Wulp has been found in Head. – Epipharynx see P. tenuicaudatus below. creeks and small rivers as well as in standing waters Antenna: AR = 2.0, length ratio segments 1 / 2 (scle- ranging from small, temporary habitats to lakes, and rotised sections only) = 4.8; antennal blade see including inland saline waters (Fittkau & Reiss tenuicaudatus. Postmentum length 185 µm. 1978). The larvae are said to live in filamentous algae Body. – Procercus with 7 anal setae (about 625 µm (Langton 1991). long) and 1 lateral seta (50 µm). Supraanal setae length 350 µm. Parachironomus tenuicaudatus (Malloch) (figs. 2, 4, 6, 9) Differential diagnoses Adult males of P. monochromus (van der Wulp) are Chironomus tenuicaudatus Malloch, 1915: 475 (adult male). distinguished by the following combination of hy- Frison, 1927: 169 (lectotype designation). Chironomus monochromus: Goetghebuer, 1921: 161 (adult popygial features: anal tergite with distinct cluster of male; see below under ‘Notes’, item 1). enlarged posterodorsal setae; anal point basal section Cryptochironomus baciliger Kieffer, 1922: 357 (adults; see intergrading with anal tergite, distal part strongly an- below under ‘Notes’, item 2). gled to ventral; portal setae 2, far apart; superior Chironomus (Cryptochironomus) monochromus and C. (C.) volsella without apicolateral projection; inferior baciliger: Goetghebuer, 1928: 75, 81, 155 (adults; see be- volsella with lobe at least to median; gonostylus most- low under ‘Notes’, items 1 and 2). Chironomus (Cryptochironomus) ‘monochromus (v.d.W.), ly slender, slightly curving, with distal widening to Goet. bacilliger, Kieff.’ (the latter an incorrect subsequent dorsal peaking around ⅔ of gonostylus length. Out- spelling): Edwards, 1929: 389 (adult male; see below un- side of the hypopygium, no morphological character der ‘Notes’, item 1). could be found separating P. monochromus from all Tendipes (Parachironomus) baciliger: Kruseman, 1933: 193 other specimens studied (table 5): the frontal tuber- (synonymization of monochromus sensu Goetghebuer nec cles were not always absent in monochromus, and a van der Wulp; adult male). bare M of the wing also occurred on some P. hazel- Paraharnischia bacilliger: Lenz, 1941: 37 (pupa). 1+2 Harnischia (Harnischia) tenuicaudata: Townes, 1945: 160 riggi, sp. n. (synonymization of baciliger Kieffer; adults; see below un- The pupa of monochromus can be separated from der ‘Notes’, item 3). known Palaearctic congeners as keyed in Langton Parachironomus bacilliger: Lenz, 1951: 105 (unjustified (1991). Among the species in the present work it emendation of species name); ? Sæther, 1977: 172 (adult stands out most strikingly by possessing vortices, anal female). Parachironomus bacilliger var. pediformis Lenz, 1951: 105 comb spines, longer points posteriorly on S II, and (adult male, larva, pupa; see below under ‘Notes’, item 4). fewer anal lobe fringe taeniae. Tendipes (Cryptochironomus) tenuicaudatus: Darby, 1962: The larva is presently practically inseparable from 50, etc. (adult male, larva, pupa). several other Parachironomus known to the author, Parachironomus tenuicaudatus: Lehmann, 1970: 152 (adult including tenuicaudatus and hazelriggi sp. n. male); Langton, 1991: 272 (pupa); Epler, 1995: 7.79, etc. (larva). Distribution and ecology The species has been reported from many western Notes on synonymy and identification European countries (Ashe & Cranston 1990), reach- 1. See chapter on P. monochromus, ‘Notes’, item 1. ing south to the Rhone valley (Laville & Serra-Tosio 2. Goetghebuer (1928) in an appendix treated ‘chi- 1991), southern Switzerland (Lodz-Crozet 1998), ronomids from Germany collected by Thienemann’, and subalpine Italy (Boorman et al. 1995). Notably, reportedly consisting of specimens ‘after which Kief- P. monochromus so far has not been found in Norway fer established his new species descriptions’ (op. cit.: and Denmark (Lindegaard 1997), nor in the 147). This listing includes P. baciliger (Kieffer). In Mediterranean. Known eastern European localities the ISNB Goetghebuer collection, in the section main- are the Volga north of Moscow (Shilova 1976), and tained as ‘types de Kieffer’, there is one male speci- the Danube delta in Romania (Albu 1980). In the Far men, mounted on a pin also holding a strip of mica
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with the hypopygium. The species represented clearly gium matching the details Lenz figured (1951: figs. 9 is P. tenuicaudatus (Malloch) as here understood. and 3, respectively). This establishes pediformis Lenz However, since there is no type label nor any infor- as a junior synonym of P. tenuicaudatus (Malloch). mation on source or sample data, the specimen can- not be recognized as a type for baciliger. Material examined. – CANADA: Ontario: 1(; St. In the Thienemann collection at ZSM, the present Lawrence Islands, Grenadier Isl., 19.vi.1975 (ZSM). – USA: Illinois (all at INHS): lectotype male adult; Mason Co., Ha- author has discovered exuviae labeled ‘bacilliger n. sp.’, vana, 28.iv.1914, C. A. Hart & J. R. Malloch; on slide, in and ‘Woltersteich’, the type locality of the species. Euparal. 1( (paralectotype); as previous except 27.iv.1914. These specimens are considered quite likely to have 1( (paralectotype); Champaign Co., St. Joseph, 3.v.1914. come from the same batch as Kieffer’s adult types. 1( (paralectotype); as previous exc. Urbana, 20.v.1914. Their observable morphology is consistent with that Minnesota: 1(; Ramsey Co., St. Paul, Highwood Park, interpretation. 10.v.1935, H. & M. Townes (AEI). New Jersey: 1(; The documentation available on the above speci- Burlington Co., Moorestown, 10.vi.1939, H. K. Townes (AEI). New Mexico: 1(; Torrance Co., ix.1925, C. H. Mar- mens is not sufficient to prove type status for them tin (AEI). New York: 1(; Schenectady Co., Niskayuna, beyond all doubt (see also Spies 1998). Nevertheless, 23.viii.1934, H. K. Townes (AEI). Ohio: 2 Lex+Pex+(; they give added support to the conclusion drawn Geauga Co., Punderson Lake, 20.ix.1992, M. J. Bolton from the morphological comparisons below: to main- (MJB). 1 Lex+Pex+(, as previous exc. Fern Lake Bog, 26- tain baciliger Kieffer as a junior synonym under P. 30.v.1993 (MJB). Pennsylvania: 1(; Montgomery Co., tenuicaudatus (Malloch). A type designation for ba- Philadelphia, Mingo Creek, 26.viii.1953, S. S. Roback (ANSP). South Dakota: 1 PM; Charles Co., Lake Francis ciliger is not deemed as necessary as required within Case, 14.viii.1967, P. L. Hudson (JES). 1(; as previous exc. the nomenclature Code (International Commission 17-24.viii.1967 (ANSP). – GERMANY: Baden-Württemberg: 1 on Zoological Nomenclature 1999). Pex+(; Bodensee at Süssenmühle, 29.viii.1962, F. Reiss 3. Townes (1945) lists a number of previous au- (ZSM). Bavaria (all ZSM): 1 Pex (det. PHL); Würm river at thors’ records whose material he apparently exam- Mühltal, 23.viii.1984, G. Wyrwa. 2(; Chiemsee, viii.1988, ined. Most notable among those are the specimens H. W. Riss. 1(; as previous exc. Starnberger See, v.1992. Schleswig-Holstein (all ZSM): 2 Pex + fragments; ‘Wolterste- described by Johannsen (1905: 228 sub ‘Chironomus ich’ (type locality of baciliger Kieffer), 2.vii.1918. 1 L, 1 Lex, modestus Say Var. b’; 1938: 43). The male genitalia 1 Pex, 2 Pex+ph ( abdominal ends; Grosser Plöner See, figured (Johannsen 1905: pl. 32, fig. 9) likely repre- vii.1950, ex coll. F. Lenz (syntypes pediformis Lenz). 1(; sent tenuicaudatus (Malloch), and contradicting pu- Grosser Plöner See, 1953, I. Müller-Liebenau. 1(; as previ- pal characters – abdominal tergite II with shagreen, ous exc. Dieksee. – UNITED KINGDOM: 1 Pex+(; England, segment VIII with 4 lateral setae (Johannsen 1938: Cumbria, Esthwaite Water, 19.viii.1985, P. H. Langton (PHL). – SOURCE UNCERTAIN: 1(; ?Germany (ISNB: Goet- 44) – may just be observation errors. However, Epler ghebuer coll., ‘Types de Kieffer’ section, No. 18.073). (1995) showed that at least Johannsen’s larvae must have been misassociated, as Johannsen (1938: 44) Adult male (also see table 5) had thought possible. For the present study, no rele- Colour. – Mostly green with usually distinct, yel- vant material could be found in the Johannsen collec- lowish to reddish thorax markings: vittae, median tion (E. R. Hoebeke, Cornell Univ., pers. comm.). anepisternum II, most of preepisternum, and postno- As suspected previously (Darby 1962, Epler 1995), tum (except anteriorly). Fore leg brownish at least
Roback’s (1957) notes on supposed tenuicaudatus lar- from tip of ta1, darkest specimens with fore leg entire- vae were also based on misassociations. The present ly brown; mid and hind legs darkened over variable author has examined a variety of ANSP specimens la- extents of tarsi. belled Harnischia tenuicaudata (Malloch) by Roback. Head. – Temporal setae in mostly double, laterally Only the adult males were correctly determined, in- single, medially occasionally triple rows. Frontal tu- cluding one from the samples reported on in 1957. bercles at least rudimentary (4 µm л circles free of mi- Four accompanying larval slides, however, proved to crotrichia), usually semiglobular or slightly elongate. hold Kiefferulus. Three larvae seen from another study Thorax. – Dorsocentral setae in single to double (label codes ‘SRP #5’, etc.) represent two different rows. Scutellar setae in 2 rows, the posterior generally species, probably of Glyptotendipes. Four pupae from more and larger. a further Roback survey (Pennsylvania, 1959) turned Wing. – M1+2 with number of setae widely variable. out to be Cryptotendipes. Legs (table 2). – Mid and hind tibiae each with the 4. Lenz’s (1951) ‘bacilliger’ var. pediformis (type lo- combs fused and the spurs subequal to slightly un- cality: Germany, Schleswig-Holstein, Grosser Plöner even, about as long as their respective comb height. See; collected vii.1950) was listed as a species under Hypopygium (figs. 2, 4). – Laterosternite IX with ‘Nomina dubia in Parachironomus’ by Ashe & Crans- 0-3, usually 2 setae. Anal tergite bands separate, me- ton (1990). Syntypic material of all life stages is still dian longitudinal part at most weakly developed. 8- preserved at ZSM, including a Pex and a male hypopy- 15 caudal setae, never strong or in dorsal positions,
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Table 2. P. tenuicaudatus, adult male: lengths of leg seg- except for hook row, the latter covering 25-35% of ments in µm (largest / smallest complete specimen). segment width, consisting of 26-37 (n=5) hooks of
P1 P2 P3 which at least the median ones bear one dorsal acces- sory point each; III with shagreen not reaching much fe 890 / 660 940 / 720 1010 / 780 anterior of setae D5; VII similar to VI, but medially fe 990 / 650 980 / 650 1100 / 740 variably thinning or interrupted, and posterior points ti 810 / 530 910 / 600 1190 / 800 and bases not much enlarged. Ventral armament: I ta 1240 / 860 460 / 320 810 / 540 1 with anteriorly widespread shagreen narrowing to ta2 630 / 440 270 / 180 450 / 290 ta3 490 / 360 200 / 140 340 / 230 mid-segment, strongest anterolaterally (extending to ta4 370 / 270 140 / 90 200 / 130 parasternite); II with posterior transverse band usual- ta5 160 / 120 80 / 60 100 / 70 ly, but not always including some scattered, slightly elongate points (up to 25 µm), band at most poorly connected to more anterior, triangular field of weak shagreen, parasternite covered to anterior of PSB; III mostly lateral and ventral (some far distal) on anal usually with anteromedian sternite and anterior point. Anal point in dorsal-aspect slide mounts usual- and/or central parasternite shagreen, both may be ly surpassed by superior volsellae, 95-125 µm long, strongly reduced. Anal lobe with 1 postero-dorsal tae- ventrally and laterally microtrichiose and setose to nia and 65-100 (n=9) in fringe, the latter in stag- about ⅔ of length, bare along most of dorsal surface gered, single row, fringe setae anteriorly more densely and in entire distal ⅓; anal point distally strongly set than medially. curving to ventral (fig. 4). Portal setae 3-6 (3; 34), usually loosely spaced in a shallow arc. Superior Larva (based mostly on two Nearctic Lex volsella straight or with slight bends near base and individually associated with Pex+() apex, 120-160 µm long, distally moderately widened Similar to P. hazelriggi sp. n., except as follows. (mostly dorso-ventrally), with a lateral, subrectangu- Head. – Labral chaetae 1 or 2 (material of Lenz lar to subacute projection sometimes longer than dis- 1951). Epipharynx with pecten of 7 teeth strongly de- tal volsella width; setal pits variable, subequal and ad- creasing in size toward lateral, and with about 6 jacent (but not parallel), or median pit smaller and a chaetulae of different sizes. Mandibular seta interna little proximal, 2 of 13 specimens unilaterally with 3 with a third, apically simple branch. Antenna (fig. 9): pits and setae. Inferior volsella roundedly rectangular AR = 1.8, 1.9, length ratio segments 1 / 2 (sclerotised or with low prominence to posterior, and with the sections only) = 4.7, 4.8, segments 3 / 4 = 0.8, 1.0; distal microtrichia usually distinctly elongate; mi- blade strong-walled almost to apex, reaching near tip crotrichia field often extending onto base of superior of segment 3, accessory blade strong-walled only prox- volsella and to adjacent area of gonocoxite (fig. 4). imally, reaching near tip of segment 2. Ventromental Gonostylus slender, gently curving or bending to me- plate with 7-9 blunt peaks at anterior margin, 13 lon- dian, with near-basal expansion bearing 4-13 (7; 30) gitudinal striae creases, and 2-4 posterior, transverse lateral and ventral setae isolated by an adjacent bare striae. Seta submenti at least 50 µm long. Postmentum section, and with distal widening to dorsal peaking at length 200, 210 µm. about ⅔ to ¾ of length; 5-8 distalmedian setae. Body. – Procercus hardly higher than wide, with 1 or 2 weak lateral setae. Supraanal setae 350, 400 µm Adult female long, basally swollen. Not studied. For unverified published descriptions see the taxonomic references listed above. Intercontinental differences The Palaearctic adult male specimens examined Pupa show setation patterns slightly differing from their Ne- Similar to P. hazelriggi sp. n., except as follows. arctic counterparts (table 5): while temporal, scutellar, Length. – Exuviae 3.9-6.6 mm (Langton 1991 and and especially wing vein setae counts are relatively high author’s data, combined n = 8). (compare respective wing size ranges), those of the Head. – Frontal seta 25-35 µm long, cephalic tu- mesonotal and squamal setae are not, and numbers of bercle longer overall, its distal cone may appear sube- clypeals are even relatively low. In addition, the distal qual in flattened exuviae mounts. macrotrichia on the inferior volsella tend to be not Thorax. – Thoracic horn of numerous branches, quite as long in the Palaearctic. with surface denticles of at most moderate size (fig. 6). The Nearctic pupal exuviae studied are smaller Prealar tubercle with narrow, low anterior, and broad, than the known Palaearctic size range (3.9-4.6 versus higher posterior sublobe, overall height < ½ length. 5.0-6.6 mm, respectively). Abdomen. – Dorsal armament: Segment II bare Future studies may resolve these differences as arte-
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3
4
5
Figs. 3-5. Parachironomus, adult male hypopygia (lateral; upper gonocoxite/gonostylus removed). – 3, P. monochromus; 4, P. tenuicaudatus; 5, P. hazelriggi sp. n. ps = positions of portal setae.
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facts resulting from the relatively low number of Eu- Alcocer et al. 1993), with the latter two records again ropean specimens analyzed. In any case, they are con- based on immatures only. sidered insufficient to warrant taxonomic separation Lenz (1951) observed the larvae to feed on (mostly of the two continental forms. deceased) pupae of other chironomids. Darby (1962) found them to live as rather solitary omnivores build- Differential diagnoses ing loose tubes on submerged plants. Adults emerged Adult males of P. tenuicaudatus (Malloch) are dis- throughout the summer season, but the life cycle was tinguished by the following combination of hypopy- interpreted to include 1-2 generations per year. gial features: anal tergite without posterodorsal setae; Swarms were observed over ground-level markers, in anal point narrow and long, but shorter than superior aggregation with several other species. volsella, basal setiferous part extended to ⅔ of anal point length, distal bare section curving to ventral; Parachironomus hazelriggi sp. n. portal setae loosely spaced; superior volsella with api- (figs. 5, 7, 8, 10, 11) cal lamellar projection; inferior volsella with lobe at least to median, distal microtrichia elongate; gonosty- Harnischia (Harnischia) monochromus: Townes, 1945: 160 lus slender, curving, with distal widening to dorsal (adults, in part; see below under ‘Notes’, item 1). ? Sub- ⅔ ¾ lette, 1957: 392 (pupa, NOT larva; see below under peaking at to of gonostylus length. Outside of ‘Notes’, item 2). the hypopygium, no morphological character could ? Harnischia (Harnischia) sp.: Sublette, 1957: 392 (larva; see be found identifying adult male P. tenuicaudatus below under ‘Notes’, item 2). against all similar species studied (table 5). ? Parachironomus ‘monochronomus (Wulp)’: Sublette, 1960: The pupa of P. tenuicaudatus shows only minor 223 (incorrect subsequent spelling; adult male). differences from that of hazelriggi sp. n., as given in Parachironomus monochromus: Beck & Beck, 1969: 285 (adults, larva, pupa; see below under ‘Notes’, item 3); the description above. For separation from other Epler, 1995: 7.79, etc. (larva). species, see the diagnosis under hazelriggi sp. n.. Parachironomus tenuicaudatus: Beck & Beck, 1969: 286 The larva is presently practically inseparable from (adult male, pupa, at least in parts; see below under several other Parachironomus known to the author, ‘Notes’, item 3). including monochromus and hazelriggi sp. n. Notes on synonymy and identification Distribution and ecology 1. Although Townes (1945) admitted he had not In the Nearctic region, P. tenuicaudatus (Malloch) compared Palaearctic material for his identification, is very widely distributed, occurring from British Co- his species concept for ‘monochromus (Wulp)’ has lumbia to Québec in Canada (Oliver et al. 1990), and been uncritically followed in numerous publications all across the USA to southern California (Sublette on North American Chironomidae. For obvious rea- 1960) and Florida (Caldwell et al. 1997). Further dis- sons, only a small amount of the related material tribution into Central America is thus not unlikely. could be examined in the present study. The selection However, a record from central Mexico (Alcocer et al. from Townes’ (1945) specimens received did not 1993) is considered uncertain as it appears to be based consist of P. hazelriggi sp. n. exclusively. At least part on immature specimens only. In the Palaearctic, the of the material from New Jersey, Medford Lakes (at species has been found from Ireland to Denmark and AEI) is of a different species falling within the polytyp- Norway (Ashe & Cranston 1990, Lindegaard 1997), ic category ‘Unresolved’ in the key below. P. gillespiae and south to northeastern France (Laville & Serra- sp. n. also runs to ‘monochromus (Wulp)’ in Townes Tosio 1991), prealpine Switzerland (Lodz-Crozet (1945). Therefore, all historic specimens and data 1998) and Germany. A record from China (after with the latter identification should be carefully reex- Wang & Zheng 1993) based on larvae is unreliable amined for their proper specific placement. since the species is not identifiable in that stage. 2. Sublette (1957) briefly characterized two In Europe, P. tenuicaudatus has been said to inhab- Parachironomus species (sub Harnischia after Townes it macrophyte stands in the litoral of lakes (Lehmann 1945) from a reservoir. The two larval descriptions 1970, Fittkau & Reiss 1978). The original samples of were later said to have been inadvertently switched baciliger Kieffer came from ‘chalk encrustations on (Beck & Beck 1969). One of them was linked to a Potamogeton lucens’ (Lenz 1941). In North America pharate male pupa determined as ‘monochromus’, but the species has also been found in rivers (Caldwell et the association was no better than circumstantial. All al. 1997) and springs, but in the latter only where descriptions given contain discrepancies to the mor- there were larger pools with partly submerged macro- phologies of P. hazelriggi sp. n.. No specimens could phytes around the edges (Ferrington et al. 1995). P. be examined, and with unassociated larvae of the rel- tenuicaudatus habitat has been reported to include evant species still inseparable the identification will oligohaline waters (Kruseman 1933, Colburn 1988, remain unsolved.
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3. For Beck & Beck’s (1969) concept of Florida Peekskill, 6.vii.1941, H. K. Townes (AEI). Ohio: 1(; ‘monochromus (Wulp)’, only one insect, a complete Ottawa Co., N Bass Island, Fox’s Marsh, 6.vii.1995, rearing spread over 2 slides, could be obtained in the L. Brohl (MJB). – MEXICO: Guanajuato: 1(; reservoir present study. The labels show source data only as W of San Miguel de Allende, 20.i.1970, F. Reiss sample codes, which are identified below according to (ZSM). Mexico City D.F.: 1(; pond betw. Mexico FAMU records. The corresponding locality, however, City and Toluca, 19.i.1970, F. Reiss (ZSM). is not among those given in Beck & Beck (1969). Nevertheless, from their description and figures as Etymology well as the specimen seen, the present author feels jus- Named after Jack Hazelrigg, manager of the tified in assuming that all ‘monochromus’ sensu Beck Greater Los Angeles County Vector Control District & Beck are in fact hazelriggi, sp. n. (Santa Fe Springs, California, USA), without whose Beck & Beck’s (1969) ‘tenuicaudatus’ was based on patient, yet active support the work presented here one Pex+( association and another single male adult. could not have been performed. The former was recognized as a misidentified ‘mono- chromus’ by Epler (1995), and is here transferred to P. Adult male (also see table 5) hazelriggi sp. n.. The second male specimen was not Colour. – Variable seasonally and regionally: most available for the present study. often thorax yellowish green with distinctly darker vittae, median anepisternum II, preepisternum, and Type material. – Holotype. – Adult male; USA: Cal- postnotum (except anteriorly); fore leg mostly ifornia: Orange County, Anaheim, Santa Ana River brownish, the mid and hind legs only apically; ab- retarding basin N of Ball Rd. (‘Burris Pit’), 9.xi.1992, domen green, hypopygium brownish with superior leg. M. Spies; on slide, in Euparal (at ZSM). volsellae and bases of gonostyli lighter. Darker speci- Paratypes. – CANADA: Manitoba: 1(; Delta Marsh, mens generally greyish to brownish-olive with tho- Bone Pile Pothole, 16.v.1980, D. Wrubleski (ZSM). racic markings little contrasting, lighter specimens USA: California (all leg. M. Spies): Los Angeles Co. greenish with darker markings indistinct. (at GLACVCD): 1 Pex; Pico Rivera, San Gabriel River Head. – Temporal setae in laterally single, medial- at Washington Blv., 13.iv.1993. 1(; Long Beach, ly double to triple rows. Frontal tubercles at least Los Angeles River at Pacific Coast Hwy, 24.iii.1993. rudimentary (4 µm л circles free of microtrichia), Orange Co., Anaheim: 3 Pex (on 1 slide), 1(; Krae- usually semiglobular or slightly elongate. mer retarding basin, 27.viii.1992 (OCVCD). 2 Pex, 1 Thorax. – Dorsocentral setae in at most briefly Pex + part of ph(; same as holotype (ZSM). 1(; San- double or triple rows. Scutellar setae in 2 rows, the ta Ana River S channel E of Fwy 91, 17.v.1993 (on posterior generally more and larger. slide with Pex of Parachironomus sp.; at OCVCD). Wing. – M1+2 with less than 10 distal setae, on some Florida: 1 Lex+Pex+(; ?Leon Co., ?Lake Munson, wings bare. ?6.v.1964, E. C. & W. M. Beck (2 slides; FAMU; see Legs (table 3). – Mid and hind tibiae each with the above ‘Notes’, item 3). 1 Pex+(; Polk Co., Lake St. combs fused and the spurs subequal to slightly un- Claire, 21.i.1966, E. C. & W. M. Beck (FAMU). New even, about as long as their respective comb height; 2 Mexico: 1 Pex+(; Eddy Co., Pecos River nr. state of 17 specimens with unilaterally 3 spurs. line, 21.xi.1974, M. Beard (JES). 1 Pex+(; San Hypopygium (figs. 5, 11). – Laterosternite IX with Miguel Co., Conchas Lake, 9.ix.1964, J. E. & M. F. 0-4, usually 2 setae. Anal tergite bands separate, medi- Sublette (JES). New York: 1(; Rockland Co., Haver- an longitudinal part present, rudimentary, or absent. straw, 24.viii.1936, H. K. Townes (AEI). 1(; 8-22 (14; 15) caudal setae, often including a few in Westchester Co., Hudson River, Peekskill Bay at dorsal position, these not always stronger. Anal point far surpassing tips of superior volsellae, 105-145 µm long, its basal section distinctly widened in dorsal as- Table 3. P. hazelriggi, sp. n.: lengths of leg segments in µm pect, the distal, bare part (>50% of anal point length) (largest / smallest complete specimen). narrow throughout in dorsal view, in lateral aspect gradually narrowing from wider base (fig. 5). Portal P1 P2 P3 setae 3-6 (5; 28), usually all in a tightly placed arc or row, occasionally one seta at greater distance. Superior fe 920 / 600 960 / 620 1080 / 690 volsella straight, 80-115 µm long, distally little to ti 820 / 500 890 / 580 1160 / 770 moderately widened, with a lateral, subrectangular to ta1 1090 / 830 410 / 290 730 / 510 ta2 540 / 440 250 / 170 430 / 290 subacute, lamellar projection usually shorter than dis- ta3 440 / 370 190 / 130 330 / 220 tal volsella width; setal pits variable, subequal and ad- ta4 330 / 260 140 / 80 190 / 130 jacent (but not parallel), or median pit smaller and a ta5 140 / 90 90 / 60 110 / 80 little proximal; median pit missing on 1 of 34 volsel-
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6 7 8
Figs. 6-8. Parachironomus, pupae. – 6. Part of thoracic horn, P. tenuicaudatus; 7, Part of thoracic horn, hazelriggi sp. n.; Scale: 50 µm; 8, Abdominal segment II (ventral), hazelriggi sp. n.; not to scale. lae. Inferior volsella with median contour in dorsal present (antero-)lateral of the precorneals. view quickly receding to proximal, without distinct Abdomen. – Dorsal armament agreeing with usual prominence to median or posterior; microtrichia not Parachironomus pattern (Langton 1991): Segment I elongate. Gonostylus slender, slightly curving in dor- bare; II bare except for hook row, the latter covering sal aspect, with near-basal expansion usually bearing 30-40% of segment width, consisting of 27-41 (n=6) 4-9 lateral and ventral setae isolated by an adjacent hooks of which at least the median ones bear one dor- bare section, in 1 specimen (Pecos River, NM) with sal accessory point each; III with or without variably 13, 14 setae merging into more distal ones; gonostylus extensive fine shagreen anterior to the usual postero- with rapid widening to dorsal at about midlength, median patches; IV, V with median longitudinal sha- slowly narrowing thereafter; 5-8 distalmedian setae. green, anteriorly spreading to lateral, points coarser posteriorly; VI similar to V, but posteriormost points Adult female enlarged and with elongate bases; VII similar to VI, but Not studied. A few individual associations with Pex posterior points and bases not much enlarged; VIII are available from the authors California rearings. with anterolateral shagreen patches nearly meeting me- dially; IX bare; conjunctives III/IV to V/VI with fine, Pupa needle-like spinules, fields at least as wide as preceding Length. – Exuviae: male 3.8-5.2 mm (4.9; 9), fe- tergite flap. Ventral armament: I with usually wide- male 5.4 (1). spread shagreen strongest anterolaterally (extending to Colour. – Exuviae pale yellowish to mostly trans- parasternite), weak and often variously interrupted me- parent. dially; II (fig. 8): parasternite covered with conspicuous Head. – Frontal seta 30-50 µm long, cephalic tu- shagreen, sternite with posterior transverse band in- bercle longer overall, its distal cone may appear short- cluding some slightly elongate points (up to 20 µm), er in flattened exuviae mounts. band with lateral projections to anterior, and more or Thorax. – Mesonotum with extensive, low granula- less connected medially to more anterior, triangular tion. Thoracic horn of numerous branches, some with field of weak shagreen; III similar to II, but without conspicuous surface denticles (fig. 7). Antepronotals: posterior transverse band, parasternite shagreen weaker 1 dorsal (usually semi-taeniate), 1 lateral (about twice and limited to anterior ⅓ to ⅔; IV bare, vortex absent; as long and taeniate); precorneals: 1 anterior (at least V at most with few, weak pointlets anterior to seta V1 ; semi-taeniate), 1 posterior (longer and taeniate); dor- VI-VIII with anterolateral shagreen patches, VIII with- socentrals evenly spread or the posterior 2 or 3 ap- out anal comb spines; IX bare. Dorsal setae I: 2 (an- proximated, Dc4 longest and usually at least basally terolaterally and centrally, up to 150 µm long); II: 3 semi-taeniate, Dc2 > Dc3, both weak, Dc1 shorter (D1 longest, D2 weak, D5 on posterior tergite flap); III- than Dc2 but stiff. Prealar tubercle with single peak VII: 5 (D2-4 weak, D5 short, stiff, on tergite flap); VIII: near middle, anterior sublobe not distinct, overall 1 (posterior, long, semi-taeniate). Lateral setae I: 1 height < length. A small, sclerotised tubercle usually (near mid-length of segment); II (fig. 8): 3 (1 dorsally
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as figured in Epler (1995: 7.78) next to lower half of key couplet 7: with 15 teeth, median tooth about 2x width of first lateral, penultimate lateral tooth project- ing as far as its median neighbor, outer mentum mar- gin not dissected. Ventromental plate slightly narrow- er than mentum (measured perpendicular to longitudinal axis of head), with 7-10 blunt peaks at anterior margin, 10-11 longitudinal striae creases, and 3-4 posterior, transverse striae. Seta submenti near posteromedian corner of ventromental plate, 40 µm long, thin and flexible. Postmentum length 180 µm. Body. – Anterior parapods with long and short claws bearing at least a few inner teeth. Posterior para- 9 10 pod claws simple (without the inner teeth figured by Beck & Beck 1969). Procercus with 7 anal setae (about 600 µm long) and 1 lateral seta. Supraanal se- tae length 300 µm.
Figs. 9, 10. Parachironomus, larval antennae. – 9, P. Differential diagnoses tenuicaudatus; 10, P. hazelriggi sp. n.. Scale: 50 µm. Adult males of P. hazelriggi, sp. n., are distinguished by the following combination of hypopygial features: anal tergite without dense cluster of enlarged pos- and 1 ventrally near mid-segment, 1 under PSB, the terodorsal setae; anal point longer than superior volsel- two ventral ones stronger); III, IV: 3 (2 near middle, 1 la, basal section distinctly widened, distal bare part a posterior); V-VII: 4 (all long, individually variable tapering blade; portal setae base marks closely adja- from semi-taeniate to taeniate); VIII: 5 (as preceding, cent; superior volsella with apical lamellar projection; the posterior two usually next to each other). Ventral inferior volsella reduced, without lobe to median or setae I: 2 (both near mid-segment, up to 100 µm long); posterior; gonostylus slender, slightly curving, with II (fig. 8), III: 3 (1 anterior, 2 posterior); IV-VII: 4 (1 rapid widening to dorsal near midlength. Outside of anterior, 3 posterior, V3 weak); VIII: 1 (central, long, the hypopygium, no morphological character could be semi-taeniate). O-setae present on T II-VII and (fur- found identifying P. hazelriggi sp. n. against all similar ther laterally) on S II-VIII. PSB present anteriorly on I, species studied (table 5). Compared to P. gillespieae sp. posteriorly on II. Anal lobe with 1 postero-dorsal tae- n., hazelriggi sp. n. carries far fewer setae on M1+2 of the nia and about 90-150 in fringe, the latter in staggered, wing, but more sensilla chaetica on the mid ta1. single row, fringe setae anteriorly and sometimes poste- The pupae of hazelriggi sp. n. and tenuicaudatus riorly more densely set than medially. can be distinguished from other Holarctic Parachi- ronomus by the following character combination: vor- Larva (from one Lex individually associated with tex and anal comb spines absent; abdominal segment Pex+() II with 2 lateral setae near mid-length and 1 under Head. – Labral S I, II simple, strong, their sockets PSB; sternite II with anterior and posterior shagreen, with long projections to oral, S II longer; S III fine, including slightly elongate points posteriorly; tho- S IVA 2-segmented, S IVB a short peg; 1 long chaeta racic horn with surface denticles. P. hazelriggi sp. n. lateral of S II; seta praemandibularis present. Pre- differs from tenuicaudatus only in details, most no- mandible not as dark as mentum margin and tably by the longer thoracic horn surface denticles, mandibular teeth, with 2 teeth and a more proximal, and the simple prealar mound. However, reliable de- median lobe. Pecten epipharyngis apparently as usual termination of unassociated material remains impos- in the genus. Mandible with apical plus 2 inner teeth, sible as long as pupae of P. gillespieae sp. n. and sever- all brown; seta subdentalis a curving, basally broad al other Nearctic species are still unidentified. blade, reaching to tip of proximal inner tooth; pecten The larva of hazelriggi sp. n. is presently practically mandibularis of two long blades; seta interna with two inseparable from several other Parachironomus known major branches visible, both apically serrate. Antenna to the author, including monochromus and tenuicau- (fig. 10) 5-segmented; AR = 1.7, length ratio segments datus. The slight differences described above could be 1 / 2 (sclerotised sections only) = 4.1, segments 3 / 4 = due to the very low numbers of reliably associated 0.9; antennal seta subapically on segment 1, ring or- specimens available, or to different individual sizes gan at about ⅓ of segment length from base. Mentum (e.g. antennal segment proportions, see figs. 9, 10).
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ps
ps
11 12
Figs. 11, 12. Parachironomus, adult male hypopygia (dorsal). – 11, P. hazelriggi sp. n.; 12, P. gillespieae sp. n. Scale: 100 µm; ps = positions of portal setae.
Distribution and ecology Parachironomus gillespieae sp. n. Parachironomus hazelriggi, sp. n. so far has been (fig. 12) verified to occur all across the southern Nearctic re- Parachironomus dentatus Gillespie, 1974: 174 (unavailable gion including Mexico, and in the central and eastern name, see ‘Comments’ after the Nearctic key below; adult parts of the continent to reach north to Manitoba, male). Ohio and New York. The species has been found in lakes, rivers, and Type material. – Holotype. – Adult male; USA, Ida- streams (Caldwell et al. 1997; author’s data). ho, Kootenai County, Twin Lakes, 22.vi.1971, leg. J. In southern North America, emergence from low- M. Gillespie; on slide, in Diaphane (at WFBM). Note: land sites apparently takes place throughout the year this is not identical to the intended ‘holotype’ of Gille- (California: author’s records; Florida: Beck & Beck spie (1974), but a different specimen from the same 1969). Especially noteworthy are the two males from sample. – Paratypes (as holotype, except as follows). – high-altitude sites in central Mexico, taken in January. 6(; all data as holotype (3 at WFBM, 2 at ZSM, 1 at Parkin & Stahl (1981) reared a species they deter- USNM). 2(; Kootenai Co., Rose Lake, 16.vi.1971 (1 mined as P. monochromus after Townes (1945) from a each at WFBM, ZSM). 1(; as previous exc. 17.vi.1971 power plant cooling reservoir in Illinois (USA). The (USNM). 1(; Benewah Co., St. Maries River 1.5 mi. E larvae were tolerant of elevated water temperatures St. Maries, 15.vi.1971 (WFBM). (frequently above 30 °C), and were mostly collected from Myriophyllum or from artificial substrates sus- pended in near-shore open water.
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Etymology hind coxa dark brown. Fore leg uniformly light brown, Named after the late Dr Janice Marie Gillespie who mid and hind legs stramineous, somewhat darkening collected the specimens here treated, and recognized only at tips. Abdomen uniformly light brown. them as distinguishable from similar, described species. Head. – Temporal setae in briefly staggered single to partially triple rows. Frontal tubercles present at Adult male (also see table 5) least as 5 µm л semiglobes. Colour (mostly after Gillespie 1974). – Head, Thorax. – Dorsocentral setae in at least partially mouthparts, antennal flagellum and plume light double rows. Scutellar setae in 2 rows, the posterior brown, pedicel dark brown; thorax ground color generally more and larger. stramineous, vittae, median anepisternum II, preepis- Wing. – M1+2 with over 15 setae along at least dis- ternum except posterodorsally, postnotum, mid and tal half, never bare.
Table 5. Parachironomus, adult males: non-hypopygial morphologies.
Character monochromus hazelriggi gillespieae tenuicaudatus tenuicaudatus sp. n. sp. n. (Nearctic) (Palaearctic)
Wing length [mm] .-. (.; ) .-. (.; ) .-. (.; ) .-. (.; ) .-. () Temporal setae - (; ) - (; ) , - (; ) , Frontal tubercle [µm], absent (), л () rudim. to x л to x rudim. to x л to x LxW where applicable Eye extension, ommatidia per lat. diagonal per lat. diagonal per lat. diagonal per lat. diagonal per lat. diagonal pattern (see Spies 1994)
Antennal ratio .-. (.; ) .-. (.; ) .-. (.; ) .-. (.; ) .-. () Clypeus setae - () - (; ) , - (; ) - () Palpomere length [µm] - - - - - Palpomere - - - - - Palpomere - - - - Palpomere - - - - -
Scutal tubercle low v. low to distinct v. low to low v. low to low v. low to low Antepronotal setae - () - () , - () , Acrostichal setae - () - (; ) , - (; ) , Dorsocentral setae - () - (; ) - () - (; ) - () Prealar setae - (; ) - () - () - (; ) - () Supraalar setae () () () () () Scutellar setae - () - () , - () - () Wing VR .-. () .-. (.; ) .-. () .-. () .-. ()
C setae, submarginal often absent prox. of var. extent, rarely to about ¾ dist. to at least ½ dist. to at least ½ dist. row of RM, at most ½ absent prox. of RM RM to arculus RM to arculus RM to arculus dist. to arculus R setae - () - (; ) , - () - ()
R setae - () - (; ) , - () - ()
R+ setae - () - (; ) , - () - ()
M+ setae () ()- (; ) - (; ) - () - () Squamal setae - () - (; ) () - () - ()
Leg ratio LR .-. (.; ) .-. (.; ) .-. (.; ) .-. (.; ) .-. ()
LR .-. () .-. (.; ) .-. () .-. () .-. ()
LR .-. () .-. (.; ) ., . .-. () .-. ()
Mid ta sensilla chaetica - () - () - (; ) - (; ) - ()
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Table 4. P. gillespieae, sp. n.: lengths of leg segments in µm (holotype).
P1 P2 P3 fe 880 900 1020 ti 750 840 1130 ta1 1090 430 730 ta2 580 260 420 ta3 440 180 310 ta4 300 130 160 ta5 120 80 90
Legs (table 4). – Mid and hind tibiae each with the combs fused and the spurs subequal to slightly un- even, about as long as their respective comb height. Hypopygium (fig. 12). – Laterosternite IX with 1-2 Fig. 13. Parachironomus sp. Bolton, adult male superior setae. Anal tergite bands separate, median longitudinal volsella (dorsal). Scale: 50 µm. part weak at most. 6-12 caudal setae, mostly lateral and ventral on basal, microtrichiose part of anal point. Anal point surpassing tips of superior volsellae, 90-
120 µm long, narrow throughout in both dorsal and sp. n., gillespieae sp. n. carries far more setae on M1+2 lateral aspects; distal, bare section at least ½ of anal of the wing, but fewer sensilla chaetica on the mid ta1. point length. Portal setae 2-4 (3; 22), separated from each other by distances at least equal to the diameter of Distribution and ecology their bases. Superior volsella straight or slightly bent, So far known only from northern Idaho. ‘Adults 90-110 µm long, distally little to distinctly widened, were swept from dense shoreline vegetation around with a lateral, at least partially sclerotized spur often small lakes and a sluggish stream’ (Gillespie 1974). longer than distal volsella width; setal pits variable, subequal and adjacent (but not parallel), or median pit Provisional key to adult males of Parachironomus smaller and a little proximal. Inferior volsella at least known from the Nearctic region roundedly rectangular, usually with some prominence to posterior; microtrichia not elongate. Gonostylus (P. elodeae and P. pectinatellae are interpreted from slender, gently curving in both dorsal and lateral as- the literature only. Species marked with an * are com- pects, with near-basal expansion bearing 6-8 lateral mented on below the key. Distributions beyond the and ventral setae isolated by an adjacent bare section, Nearctic are indicated as follows: »C = to Central and with rapid widening to dorsal near midlength, America, »P = to Palaearctic, »S = to South America). slowly narrowing thereafter; 5-6 distalmedian setae. 1. Eye with the dorsomedian extension composed of Other life stages diagonals with up to 5 ommatidia each (as in Unknown. Spies et al. 1994: fig. 1) ...... 2 – Several diagonals within eye extension of 6 om- Differential diagnosis matidia each ...... 20 P. gillespieae sp. n. is distinguished by the following 2. Wing with setae restricted to margins and veins . combination of adult male hypopygial features: anal ...... 3 tergite without enlarged posterodorsal setae; anal – Wing cells also with setae, at least near wing tip . point about as long as superior volsella, narrow ...... 18 throughout, its distal bare section at least ½ of total 3. Gonocoxite distally on dorsal to median surface anal point length; portal setae base marks not touch- with dense cover of long setae. Superior volsella ing; superior volsella with apical, partially sclerotized with extensive median lining of microtrichia ...... spur; inferior volsella with lobe at least to median; ...... danicus Lehmann* »P gonostylus slender, gently curving, with rapid widen- – Not with above combination ...... 4 ing to dorsal near midlength. Outside of the hypopy- 4. Superior volsella without apico-lateral projection gium, no morphological character could be found (if terminal setal pit with elevated rim, then the identifying P. gillespieae sp. n. against all similar latter of even height all around). Gonostylus species studied (table 5). Compared to P. hazelriggi without widening to dorsal in distal half ...... 5
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– Superior volsella usually with near-apical, lateral – Gonostylus widest in midregion ...... lobe or spur (at least with low prominence from ...... varus (Goetghebuer) »P rim of terminal setal pit); if volsellar projection 13. Superior volsella with subapical, fimbriate projec- absent, gonostylus widened to dorsal in distal half tion (fig. 13) ...... sp. Bolton* ...... 9 – Projection of superior volsella never fimbriate ..14 5. Superior volsella and anal point elongate, the lat- 14. Gonostylus in dorsal aspect mostly narrower than ter without apical knob or hook ...... 6 ½ gonocoxite width, curving or bending to medi- – Superior volsella and anal point short, the latter an and/or dorsal (figs. 2, 11, 12) ...... 16 apically widened and/or curving to ventral ...... 7 – If gonostylus rather narrow, then nearly straight 6. Superior volsella with median setal pit in distinct- ...... 15 ly proximal position, its rim often slightly pro- 15. Anal point setose to far distal, the bare part hook- truding from volsellar stem. Gonostylus with ing to ventral. Superior volsella with large, subtri- basal ½ moderately expanded, distal ½ curving angular, apico-lateral projection. Gonostylus toward median ...... abortivus (Malloch)* rather robust, its midsection not bare and strong- – Superior volsella with both setal pits in apical, ly widened to dorsal. Lehmann (1970: fig. 16) ... roughly parallel positions (Townes 1945: fig...... subalpinus (Goetghebuer)* »P 177) ...... elodeae (Townes) – Not with above combination ...... Unresolved* 7. Anal tergite with at most few setae antero-dorsal 16. Anal point in total longer than superior volsella, of anal point, the latter at apex with widened in dorsal aspect with elongate conical base. Anal hook to ventral. Gonostylus proximally strong tergite with some caudal setae lateral and dorsal and straight, distally narrower and curving to me- of anal point origin; alveoli (sockets) of portal se- dian (Spies et al. 1994: figs. 28, 31a, b) tae closely adjacent. Inferior volsella lacking me- ...... supparilis (Edwards) dian or posterior lobe. Figs. 5, 11 ...... var. centralis Spies, Fittkau & Reiss »S ...... hazelriggi sp. n. – Not with above combination ...... 8 – Anal point in total at most as long as superior 8. Gonostylus widest subapically. Anal point apical- volsella, basal widening in dorsal aspect not elon- ly with widened hook to ventral (Lehmann 1970: gate. Anal tergite caudal setae mostly restricted to fig. 20) ...... vitiosus (Goetghebuer). »P base of anal point; alveoli of portal setae not adja- – Gonostylus of mostly even circumference. Anal cent. Inferior volsella with a distal lobe at least to point without apical widening, gradually curving median. Figs. 2, 4, 12 ...... 17 to ventral ...... schneideri Beck & Beck 17. Anal point about as long as superior volsella, dis- 9. Gonostylus with bare base followed by extensive, tal bare section at least ½ anal point length. Infe- expanded section bearing numerous strong setae, rior volsella without elongate microtrichia. distal part of gonostylus narrower and curving or Gonostylus expanding to dorsal just beyond mid- bending to median. Superior volsella with large dle. Fig. 12 ...... gillespieae sp. n. apico-lateral lobe or subapical projection, setal – Anal point in total shorter than superior volsella, pits positioned symmetrically (or nearly so) at distal bare section about ⅓ anal point length. In- volsellar apex. Anal point long, strongly tapering ferior volsella with elongate microtrichia distally. from wide, dorsally setose base, with apical hook Gonostylus expansion to dorsal limited to distal to ventral ...... 10 ⅓. Figs. 2, 4 ...... tenuicaudatus (Malloch) »P – Not with above combination ...... 13 18. Superior volsella apicolaterally with large, blunt 10. Superior volsella with apico-lateral, lamellar lobe projection ...... potamogeti (Townes)* ...... 11 – Superior volsella at most with low projection – Superior volsella with subacute projection dis- from lateral rim of terminal setal pit ...... 19 tinctly proximal of setal pit region ...... 12 19. Superior volsella straight, strong, distally often 11. Gonostylus in dorsal aspect considerably narrow- considerably inflated; distal setal pit covering en- ing from wide proximal expansion. Inferior tire terminal surface, median subapical pit small- volsella with distal lobe to posterior ...... er, usually in distinctly proximal position ...... parilis (Walker)* »P ...... chaetoalus (Sublette)* – Gonostylus distally nearly as wide as in proximal – Superior volsella slightly sinuous, mostly slender; section. Inferior volsella projecting only to medi- setal pits about equal in size, both near volsellar an ...... ‘sp. B’ Sæther (1977: 172)* apex ...... hirtalatus (Beck & Beck) 12. Gonostylus widest in proximal half, narrowing 20. Anal point long and broadened, lined with setae and bending to median near middle to far distal ...... 21 ...... forceps (Townes) – Not with above combination ...... 22
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21. Anal point with constriction proximal of apical, tion set forth in the Code of Zoological Nomenclature oval lamella. Gonostylus with basal expansion (International Commission on Zoological Nomencla- limited to ⅓ length ...... frequens (Johannsen) »P ture 1999: Article 8). The present author has exam- – Anal point evenly tapering, without apical lamel- ined Gillespie’s original material (from WFBM). Five of la. Gonostylus expanded over basal ½ (Dendy & her concepts could be matched with previously estab- Sublette 1959: fig. 14) ...... lished species, as identified in the listing below. An- ...... pectinatellae (Dendy & Sublette) other has been described above as P. gillespieae sp. n. 22. Gonostylus with strong proximal expansion, es- The seventh – P. ‘abditus’ (unavailable name) – possi- pecially medioventrally ...... sublettei Beck bly also constitutes a new species, but is here excluded – Basal region of gonostylus at most slightly wider pending further revision. than more distal parts ...... 23 23. Squama with setae. Gonostylus with the distal- abortivus: P. ‘emarginatus’ (unavailable name) of median setae widely spaced and hardly stronger Gillespie (1974: 190) is conspecific. than the more proximal median setae ...... 24 chaetoalus: P. ‘biggami’ (unavailable name) of – Squama bare. Gonostylus with discrete, apical- Gillespie (1974: 180) is conspecific. median group of setae ...... 25 danicus: Nearctic record from Idaho (Gillespie 24. Squama extensively lined with setae. Anal tergite 1974) here confirmed. with dense cluster of strong setae antero-dorsal of digitalis: first Nearctic records (6 adult males) from anal point origin. Superior volsella strong South Carolina, Alabama (both JHE), and Georgia throughout, distally at most gradually widened, (BAC); indistinguishable in all details from Palaearctic often wrinkled; terminal setal pit covering entire specimens (det. Lehmann) at ZSM; juvenile stages un- apical volsellar surface, with short, blunt lateral known from either region. lamellar prominence (Lehmann 1970: fig. 8) ..... guarani: first Nearctic record (1 adult male) from ...... digitalis (Edwards)* »P Alabama (JHE); specimens keying out here should be – Squama with few setae (2-5 seen). Anal tergite checked against Neotropical Parachironomus using rarely with a few stronger setae antero-dorsal to Spies et al. (1994). anal point origin. Superior volsella distinctly nar- parilis: first Nearctic records: Idaho (Gillespie), rower in midsection, typically abruptly widening Ohio (MJB); P. ‘reductus’ and ‘trisetosus’ (unavailable distally, with setal pits roughly subequal and op- names) of Gillespie (1974: 176 and 178, respectively) posite at proximomedian and distal ends of are conspecific. widened volsellar ‘head’ (Dendy & Sublette 1959: potamogeti: wing cell setae – not mentioned by fig. 13) ...... directus (Dendy & Sublette) »C Townes (1945) – found on holotype (BAC and JHE, 25. Longitudinal anal tergite band absent. Anal point pers. comms), and on a paratype male at ZSM; P. ‘barri’ stout throughout. Superior volsella strong, distally (unavailable name) of Gillespie (1974: 181) is conspe- bending to dorsal-lateral, medio-ventrally exten- cific; specimens with indistinguishable hypopygia, but sively lined with microtrichia ...... alatus (Beck) without wing cell setae, key above to ‘Unresolved’; re- – Longitudinal anal tergite band present, usually vision of Townes’ voluminous type series required. embracing bases of several dorsal setae. Anal ‘sp. B’ Sæther: known from pharate males (Ohio) point narrow at least in midsection. Superior reared by MJB; see unique larval mentum in Sæther volsella slender, without microtrichia ...... 26 (1977: fig. 87G). 26. Anal tergite with median longitudinal ridge bear- sp. Bolton: one male from Ohio reared by MJB. ing setae, posteriorly subtruncate due to ‘shoul- subalpinus: reported from Yukon Territories and ders’ lateral of anal point. Superior volsella with ‘N Asia’ by Oliver & Dillon (1997). setal pits subequal and almost parallel at narrow Unresolved: a complex of certainly several species volsellar apex ...... carinatus (Townes) with partially intergrading character states, inseparable – Anal tergite without pronounced ridge or ‘shoul- from the sparse material seen; includes specimens with ders’. Superior volsella apically widened; terminal genitalia close to potamogeti (see comment above), but setal pit larger than the median, subapical one lacking the wing cell setae; also potentially involved is (Spies et al. 1994: figs. 13, 14) ...... the Palaearctic paradigitalis Brundin, 1949...... guarani Spies, Fittkau & Reiss* »S ACKNOWLEDGMENTS Comments In an unpublished Ph.D. thesis, Gillespie (1974) Dedicated to the memory of Dr Friedrich Reiss developed concepts for seven supposed new species, from whom I’ve learned about Parachironomus, and so but their names have remained unavailable because much more. the dissertation does not meet the criteria of publica- This work was performed as part of a doctoral dis-
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sertation supervised by Prof. Ernst Josef Fittkau at the 123: 1-53. Coffman, W. P., 1986. The pupae of Chironomidae University of Munich. Prof. Gerhard Haszprunar, di- (Diptera) of the Holarctic region – Key to subfamilies. – rector, kindly granted full use of ZSM facilities for the Entomologica scandinavica Supplement 28: 9-11. duration of this program. Colburn, E. A., 1988. Factors influencing species diversity Gilbert Challet, Frank Pelsue, and Jack Hazelrigg, in saline waters of Death Valley, USA. – Hydrobiologia district managers, initiated and patiently supported 158: 215-226. the southern California studies. That work was joint- Cranston, P. S. & F. Reiss, 1983. The larvae of Chironomi- dae (Diptera) of the Holarctic region – Key to subfami- ly funded by OCVCD and GLACVCD. lies. – Entomologica scandinavica Supplement 19: 11-15. For help with specimens, discussions, etc., I am Darby, R. E., 1962. 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Dipterologische aanteekenin- Parkin, R. B. & J. B. Stahl, 1981. Chironomidae (Diptera) gen. – Tijdschrift voor Entomologie 17: 109-148. of Baldwin Lake, Illinois, a cooling reservoir. – Hydrobi- ologia 76: 119-128. Pinder, L. C. V. & F. Reiss, 1986. The pupae of Chi- Received: 9 September 1999 ronominae (Diptera: Chironomidae) of the Holarctic re- Accepted: 15 December 1999
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