Allocladius nanseni (Kieffer) Ferrington Jr & Saether, 2011

Ferrington Jr, Leonard C. & Saether, Ole A., 2011, A revision of the genera Pseudosmittia Edwards, 1932, Allocladius Kieffer, 1913, and Hydrosmittia gen. n. (Diptera: Chironomidae, Orthocladiinae) 2849, Zootaxa 2849 (1), pp. 1-314 : 66-72

Allocladius nanseni (Kieffer) comb. n.

( Figs. 15–17 )

Camptocladius nanseni Kieffer, 1926: 85 .

Prosmittia nanseni (Kieffer) ; Oliver, 1963: 177; Saether, Sublette & Willassen, 1984: 270.

Smittia longitibia Goetghebuer, 1933: 29 ; Saether & Ferrington, 2003: 5.

Pseudosmittia mediocarinata Caspers et Reiss, 1989: 132 ; Saether & Ferrington, 2003: 5.

Pseudosmittia nanseni (Kieffer) ; Cranston & Oliver, 1988: 451; Makarchenko & Makarchenko 2008: 221.

Material examined: CANADA: Nunavut: Ellesmere Island, Indre Eide , holotype male, 4 vii 1902 , 2 nd Fram Expedition ( ZMO); Ellesmere Island , Hazen Camp, 5 males, 15.vii.1961 , D.R. Oliver , 28.vi.1966 , P. Kewan; Ellesmere Island, Grise Fjord , 1 male, 24.vi.1967 , M. Freeman; Heiberg Island, Wolf River , 1 male, 19.viii.1963 , H.K. Rutz; Baffin Island , 1 larva, 23.viii.1984 , L.I. Bernier; Southampton Island, Coral Harbor , 2 males, 11.vii.1948 , G.E. Shewell; Repulse Bay , 1 male, 26.vii.1950 , J.E.H. Martin. Yukon Territory: Herschel Island , 2 males, 29.vi, 24–28.vii.1971 , D.R. Oliver. Whitehorse , 1 male, 6.vii.1974 , N.L.H. Kreuss. Alberta: Edmonton, Wagner National Area , 1 male, 17– 24.vi.1985 , Finnmere & Thormin. Manitoba: Churchill, Bird Cove, Bluff C netting, 3 m a.s.l., N 58.76554 0 W 93.86815 0, 1 male, 12.vii.2006 , E. Stur ( VMT); Delta Marsh, 10 males, 6 females (some of which may belong to A. arenarius ), 10.viii.1980 , 28.viii & 3–9.ix.1981, 11.v.1984, D. Wrubleski; 9. & 16.vi.1981, H. Murkin, 17.v.1982, H. Neckles. Ontario: Waterloo, 1 larva, 11.vii.1972 , D. Williams; Ottawa, Britannia Filtration Plant , 1 male, 12.iv.1971 , D.R. Oliver; Green Creek , 1 male with pupal exuviae, 20.vi.1967 , L. Haig Smillie & R. MacDonald ; 1 pupal exuviae, 3 larvae, 22.v.1985 , M.E. Dillon, D.R. Oliver & P.S. Cranston . New Brunswick: Kouchibouguac National Park , 3 males, 19.v.1977 , W.P. Hanley , 8.vii.1978 , D.R. Oliver & M.E. Roussel . USA: Alaska, Point Barrow , 1 male, 7.vii.1952 , P.D. Hurst; Savonski, Naknok Lake , 1 male, 25.vii.1914 , no collector. South Dakota: Nelson's Pond, 4 mi N., 4 mi E. of Yankton , 2 males, 13.vii.1972 , P.L. Hudson; Beaver Creek , 8 mi N, 2 mi W of Yankton, 1 male, 5.iv.1973 , P.L. Hudson; Yankton, 1003 Douglas, 1 male, 1973, P.L. Hudson; Emmanuel Creek , 2 mi W Springfield, sweeping near creek and spring, 3 males, 21.iv.1973 , P.L. Hudson; Platte County, Platte Bay, Lake Francis Case , 1 male, 25.v–1.vi.1967 , P.L. Hudson, Vermillion , Missouri River, Clay County Park , 1 male, 18.vii.1971, 22.v.1972 , P.L. Hudson; Missouri River, Running Water , 2 males, 8.vi.1972 , P.L. Hudson; Missouri River, 2 mi E., 6 mi S. Gayville, 1 male, 4.x.1972 , P.L. Hudson. Minnesota: Wabashe County , Lake City, Handshaws Coulee, Handshaws Spring , 1 male, 30.vii.1981 , P.L. Hudson. Michigan: Saint Clair County, Saint Clair River, Russel Island , 1 male, 24.v.1983 , P.L. Hudson. Ohio: Franklin County, Scioto Park , soil & moss adjoining spring, 1 male, 12.ix.1987 , M.J. Bolton. Missouri: Saint Louis County, 2 males, 30.v.1990 , C.A. Wright; Saint Louis County, Mattese Creek, 2 pupal exuviae, 10.v.988, N.H. Crisp; Jefferson County , Pomme Creek , 31.v.1992 , C.A. Wright. Wyoming: Carbon County, Bush Creek Area NE of Ryan Peak, 2470 m, viii.1973 , P.H. Arnaud; Yellowstone Park, Madison Junction on Gibbon River , 1 male, 17.vii.1952 . Idaho: Mammet, 1 male, 25.vi.1956 , Knowlton. Kansas: Cheyenne County, stream SW of Saint Francis , 1 male, 3.v.1982 , L.C. Ferrington, Jr. ; Douglas County , Kansas River 2.5 mi NW of Lecompton , 12 males, 5. & 18.v.1989 , D.S. Goldhammer. Utah: Wellsville, 3males 7. & 13.v.1954 , G.E. Bohart; Garfield County, 1 male, 4.vi.1952 , E.B. Hayden; Utah County, Provo, Barbizan Pond , 1 male, 11.ix.1981 , J.E. Sublette & R.W. Bowman ; Utah County, Near Tithing Mountain, Peeteetweet Creek , swarm over scrub oaks, 3males, 15.vii.1982 , C.A. Miles; Box Elder County, Clear Creek Campground, Clear Creek , 1 male, 15.viii.1980 , Baumann & Clark; Emory County, North Hughes Canyon, Huntington Creek , 3.vi.1975 , Winget; Washington Co., 1 male, 8.vi.1952 . Colorado: M. Edith, Navaho River at New Mexico State Line , Malaise trap, 17.vii.1976 . California: Kirk Creek , 2males, 12– 13.viii.1969 , B. Hocking; Riverside County, Riverside , 9males, 1.i.1963 , S. Frommer; Riverside County, Arlington, 6724 Western , 1 male, 2.vi.1967 , R.D. Sjogren; Riverside County, Hidden Lake , 3 mi N. Arlington, 1 male, 28.iv.1967 , R.D. Sjogren; Riverside County, Palm Desert ( White Sun Guest Ranch ), sweeping grass near pond, 1 male, 17.iv.1970 , S. Frommer & L. La Pre; Lasson County, Winterwood , stream margin, 1 male, 16.v.1948 , W.W. Wirth; Inyoe County, Big Pine , 4000', 3 males, vi.1972 , D. Guiliani; Inyoe County, Chise Ranch , 1 male, 30.v.1955 ; Sishigon County , Mount Shasta City, at light, 1 male, 21.vi.1958 , J. Powell; San Bernardino County, 2 mi W. of Devote , 1 male, 28.v.1970 , M.F. Nargo; Meade County, Willow Crests , 1 male, 15.v.1948 , W.W. Wirth; Monterey County, Carmel, mouth of Carmel River , 3 males, 21.iii.1967 , G. Grodhaus; San Diego County, Sentenac Canyon Gaging , 1800–1900, 1 male, 4.v.1968 , E.J. Schlinger; Ventura County, Point Magu , 1 male, 4.iii.1950 ; Sacramento County, Rio Lucinda , 1 male, 15.x.1953 , R.E. Darby; Orange County, Aliso Creek , 1 male, 8.v.1955 , C.L. Hogue. New Mexico: Navajo River at bridge 1/ 4 mi N. of New Mexico state line between Lumberton , New Mexico and Edith , Colorado, 2 males, 9.ix.1974 , M. Beard; Qucy County, Canadian River, 2 mi E. of Logden , Highway 54, Revello Creek , Malaise trap, 2 males, 1.x.1974 ; Quay County, Taylor Springs , sweep net, 1male, 3.x.1974 ; Colfax County, Cimarron River at Taylor Springs , sweep net, 2males, 3.x.1974 ; Eddy County, 3 mi N., 1 mi E. of Loving , Malaise trap, 1 male, 14–15.iv.1973 , A.L. Gennaro; Eddy County, Pecos River, Harroun Crossing , sweep net, 1.vi.1978 , T. Bert; Chaves County, Bitten Creek , 1 male, 23.v.1983 , P. Sanchez; Chaves County, Bob Crosby Draw , 1 male, 10.iv.1983 , Tucker & Boca; Los Alamos County, Bandelier National Monument , 100 yards from stream, 1 male, 25– 26.viii.1973 , J.E. & M.F. Sublette; Catron County, San Francisco River at Glenwood , Malaise trap, 1 male, 18.xi.1974 ; Hidalgo County, Gila River at State Line near Virdun , Malaise trap, 1 male, 21.ix.1974 ; Soccoro County, Rio Grande near San Marcial , Malaise trap, 1 male, 18.ix.1947 ; Socorro County, Rio Grande near San Marcial , 2 males, 11.viii.1976 , M. Beard; Dona Ana County, Rio Grande at Texas State Line, El Paso, 1 male, 15. & 21.ix.1974 , M. Beard; Santa Fe County, Avari Bridge, Rio Grande near San Ildefanso between Santa Fe & Los Alamos, 3males, 8.x.1974 , M. Beard; Rio Grande, Oton Bridge between Santa Fe & Los Alamos, sweep net, 1 male, 16.viii.1976 ; San Juan River at Bridge 1 mi W. of San Juan Hospital , Malaise trap, 1 male, 10.xi.1974 . Texas: Randall County, Palo Duro Creek at water near Buffalo Stadium 3 mi N. of Canyon , 3 males, 5.xii.1973 , M. Beard; El Paso County, Rio Grande near El Paso, sweep net, 1 male, 8.vii.1964 , J.E. Sublette. Louisiana: East Baton Rouge Parish, Baton Rouge , at light, 2 males, 20.iii.1947 , W.W. Wirth. South Carolina: Pickens County, Clemson, Hudson's garden, wet area, 4 males, 20.iv.1980 , P.L. Hudson; Anderson County, Savannah River, Lake Hartwell Tailwaters , 1 male, 9.vii.1978 , P.L. Hudson. Georgia: Muscogee County, Weracoba Creek at Cherokee Avenue and Camille Street , 5 males, 23.x.1984 , B.A. Caldwell; Whitfield County, Tar Creek, Abutmont Road , 5 males, 28.ii.1985 , B.A. Caldwell; Jefferson County, Mc Quaddy Branch at Georgia Highway 296, 2 males, 8.v.1985 , B.A. Caldwell; Hall County, headwater area of Flat Creek near Indian Boulevard behind Wayne Feeds , swarm, 1 male, 14.xi.1985 , B.A. Caldwell; Walker County, City Creek at confluence with Town Creek , Surber sample, 1 larva, 5.ix.1974 , Caldwell and Peterson; Crawford County, Spring Creek at US 341 about 5 mi SSE Roberta , 1 male, 27.v.1982 , P.L. Hudson; Hart County, Savannah River , 4 mi below Hartwell Dam, 1 male, 18.ix.1980 , P.L. Hudson; Camden County, Saint Marys Road at Shadon Lawn , light trap, 1 male, 8.iv.1977 , M.O. Tooli. MEXICO: Mexico City, Azcapolzalco , 1 male, 5–31.vii.1921 , A. Contreras et al. GREENLAND: Eastern Greenland, male holotype of Smittia longitibia, 1930 , Hoels Expedition , ( ZMD); Thule, 1 male, 4.viii.1952 , C. Vibe; Nedre Midsommer Lake , 1 male, 7.vii.1966 , Commander Perry Land Expedition . NORWAY: No locality, 1 male, as P. mediocarinata , 25.vii.1958 , Clastier. TURKEY: Bolu Province, Yeniçaga , male holotype P. mediocarinata , 24.ix.1986 ( ZMBN) . CHINA: Inner Mongolia, Huhehaote , 1 male, 20.v.1986 , X. Wang; Inner Mongolia, Alex , 1 male, 31.vii.1987 , X. Wang; Hebei, Beidaihe, 1 male, 4.vii.1986 , X. Wang; Ningxia, Mount Liupan , 1 male, 4.vii.1987 , X. Wang; Ningxia, Tongxin, 1 male, 4.viii.1987 , X. Wang; Gagsu, Jinquan , 1 male, 12.vi.1986 , X. Wang. ( BMNH, BYU, CAS, CNN, FLAM, JES, UMSP, UCLA, UMN, USNM, ZMBN, ZMK, ZSM) .

Diagnostic characters: The virga consisting of a weak median spine and lateral groups of spinules combined with the presence of 4–16 acrostichals and a supraalar seta will separate the male imago from other members of the genus. The female imago is at present not separable from A. arenarius and A. aizaiensis . These species are separable from other known females of the group by having at most 1 non-marginal seta on R between FR and the apex of R 4+5. The pupa is characterized by even shagreen on tergites, presence of setae on the anal segment, tergal conjunctive VII/VIII at most with 2 spinules, II/III with 0–17 spinules and sternal conjunctive VII/VIII with 50–103 spinules. The larva is characterized by having 4 inner teeth on the mandible, 5 pairs of lateral teeth on the mentum, about 40 longer claws on each anterior parapod, and 11 claws on each posterior parapod.

Male imago (n = 15–17, except when otherwise stated)

Total length 1.97–3.08, 2.33 mm. Wing length 1.08–1.82, 1.32 mm (57). Total length/wing length 1.69–1.90, 1.79. Wing length/length of profemur 2.80–3.16, 2.98. Coloration dark brown to black.

Head ( Fig. 15 B ). AR 1.00–1.75, 1.21 (59). Terminal flagellomere 333–639, 427 µm (59) long, more or less apically pointed. Temporal setae 5–12, 7 (59), consisting of 1–6, 3 (59) inner verticals, 2–4, 2 (59) outer verticals and 1–3, 2 (59) postorbitals. Clypeus with 5–10, 7 setae. Tentorium 112–135, 128 µm (13) long, 23–43, 30 µm (14) wide. Cibarial pump and stipes as in Fig. 15 D . Stipes 105–130, 117 µm (13) long, 23–50, 30 µm (14) wide. Palpomere lengths (in µm): 21–27, 26; 39–57, 46; 64–94, 79; 62–94, 80; 91–137, 107. Third palpomere ( Fig. 15 C ) with 1–2, 2 lanceolate sensilla clavata.

Thorax. Median antepronotal lobes well developed, antepronotum with 1–2, 1 (12) lateral setae. Dorsocentrals 5– 12, 8, acrostichals 4–16, 8 (54), prealars 2–4, 3, supraalar 1. Scutellum with 4–8, 5 (13) setae.

Wing ( Fig. 15 A ). VR 1.31–1.43, 136. Anal lobe slightly projecting. Costal extension 0–14, 7 µm long. R 4+5 ending proximal to apex of M 3+4; Cu 1 curved. Brachiolum with 1 (59) seta, R with 0–7, 3 (59) setae, R 4+5 with 0–1, 0 (59) setae, other veins bare.

Legs. Spur of front tibia 32–48, 40 µm long, spurs of middle tibia 16–21, 19 µm (13) and 14–18, 15 µm (13) long, of hind tibia 34–46, 40 µm and 11–18, 16 µm long. Width at apex of front and middle tibiae both 25–37, 28 µm, of hind tibia 32–46, 38 µm. Comb of 13–16, 14 setae, shortest seta 17–23, 20 µm long, longest seta 32–37, 34 µm long. Lengths and proportions of legs as in Table 9.

Hypopygium ( Fig. 15 F ). Anal point 9–46, 28 µm (18) long, apex free of microtrichia; tergite IX with 14–33, 18 setae; laterosternite IX with 4–13, 7 setae. Phallapodeme 55–101, 75 µm long; transverse sternapodeme 46–62, 54 µm long, with weak or nearly absent oral projections. Virga ( Fig. 15 E ) consisting of weak median plate recognizable in only 2 specimens, 11–18 µm long, and 8–50, 25 (57) lateral spinules. Gonocoxite 139–215, 162 µm long; superior volsella present as dorsal swelling of inner margin of gonocoxite, reaching to 0.41–0.48, 0.45 gonocoxite length; inferior volsella and accessory lobe well developed, reaching to 0.60–0.64, 0.62 and 0.71–0.94, 0.79 gonocoxite length, respectively. Accessory lobe usually free at apex. Gonostylus 54–87, 63 µm long; megaseta 9–11, 10 µm long. HR 2.30–2.77, 2.55; HV 3.25–4.39, 3.77.

Female imago

The female imago ( Fig. 16 ) appears to be not separable from that of A. arenarius redescribed below and the probable female imagines of A. nanseni are included in that description. Perhaps all Nearctic and at least the female from Green Creek, Ontario belong to A. nanseni . However, while the female from Ontario has an LR 1 of 0.47, which is higher than all the other specimens, one of the females from Manitoba has a LR 1 of 0.41, at the lower end of the variation. The last mentioned specimen is also the longest with a wing length of 1.65 mm. Two additional females from Manitoba have LR 1 which are in the middle of the range of variation of A. arenarius .

Pupa (n = 5, except when otherwise stated)

Total length 2.48–4.19, 2.75 mm. Exuviae with yellowish tinge on margins of cephalothorax and anal segment.

Cephalothorax. Frontal apotome wrinkled, without warts, with frontal setae. Frontal setae, longer precorneals, median antepronotals and longer dorsocentral (Dc 3 or Dc 4) all about 55–80 µm long. Shorter precorneal, shorter lateral antepronotal, postorbitals and shorter dorsocentrals all about 30–60 µm long. Distance between Dc 1 and Dc 2 21–94, 53; between Dc 2 and Dc 2 9–39, 24 µm; between Dc 3 and Dc 4 2–21, 8 µm.

Abdomen ( Fig. 17 A–D ). Tergite I bare or with 2–3 posteromedian spinules; T II–VIII with strong anterior, weak median and strong posterior spinules; T IX with extensive median and lateral shagreen. Sternites I–II bare, S III with weak median spinules, S IV with stronger median spinules, S V–VIII with shagreen in anterior 2/3, S IX bare or with anterolateral shagreen. Tergal conjunctive II/III with 0–17, 6 spinules; III/IV with 25–115, 32; IV/V with 90–105, 99; V/VI with 96–140, 116; VI/VII with 90–135, 114: VII/VIII with 0–2 (3) in female, 36–47 (2) in male. Sternal conjunctives III/IV with 8–33 (3) spinules in female, bare in male; IV/V with 46–125, 89; V/VI with 85–150, 114; VI/ VII with 80–107, 97; VII/VIII with 50–103, 77 spinules. Anal segment with 34–59, 44 µm long basal seta and 2–3 apical setae 34–73, 48 µm long. Genital sac of female with even rugulosity and an apical papilla at most as long as wide. Genital sac of male with ventral spinules most extensive at base and apical papilla; sac overreaching anal lobe by 85–91 µm (2).

Fourth instar larva (n = 4–5, except when otherwise stated)

Total length 2.79–5.58, 3.92 mm. Head capsule length 0.27–0.33, 0.30 mm.

Head. Antenna as in Fig. 17 E . Length of antennal segments (in µm): 8.5–13.5, 11; 4–6.5, 5; 2–2.5, 2; 2–3, 2.5. AR 1.00–1.18, 1.12. Basal antennal segment 15.5–19, 17 µm wide; blade 11.5–14.5, 13 µm long; accessory blade 5.5– 6.5, 6 µm long. Sensilla basiconica 2–3.5, 3.5 µm long. Subapical style of second segment 3.5–5.5, 4.5 µm long. Premandible ( Fig. 17 F ) 51–69, 61 µm long. Mandible ( Fig. 17 G ) 107–126, 116 µm long; with 4 inner teeth and apparently 1 dorsal tooth, seta subdentalis 5.5 µm long; seta interna absent. Mentum as in Fig. 17 H , with 5 pairs of lateral teeth, median tooth 25–32, 29 µm wide, ventromental plate 7.0 µm wide in flattened mentum. Postmentum 89– 119, 107 µm long.

Abdomen. Anterior parapods 114 µm (1), each with about 40 larger claws. Posterior parapods 91–114 µm (3) long; with 11 claws, 34–41, 36 µm long. Dorsal anal tubules 21µm (1) wide at base, ventral anal tubules 41 µm (1) wide; lengths not measurable.

Remarks

This species shows a very large variation and more than one species could be involved. However, there appears to be no consistent seasonal or geographic pattern to the variation among the specimens that we examined.

Ecology and distribution

This species appears to be at least truly aquatic in part of its range, but can probably also live in moist sandy substrates and apparently in salt marshes. There are many Nearctic records from Greenland and Alaska to Mexico, while in Europe the species is recorded only from Norway, Sweden, Russia (including Novaya Zemlya and the Far East) and Turkey. The records from China are all within the Palaearctic regions of China.

Figures

FIGURE 15. Allocladius nanseni (Kieffer) comb. n., male. A—wing; B—head; C—third palpomere; D—cibarial pump and stipes; E—virga; F—hypopygium, left dorsal aspect, right ventral aspect.

FIGURE 16. Allocladius nanseni (Kieffer) comb. n., female. A—wing; B—thorax; C—antenna; D—head; E—genitalia, ventral view; F—dorsomesal lobe; G—ventrolateral lobe; H—apodeme lobe; I—genitalia, dorsal (partly lateral) view.

FIGURE 17. Allocladius nanseni (Kieffer) comb. n., immatures. A—D, pupa; A—posterior sternites of male; B—posterior tergites of male; C—posterior tergites of female; D—posterior sternites of female. E—H, larva; E—antenna; F—premandible; G—mandible; H—mentum.