Zootaxa, Diptera, Chironomidae · 1981 and Litocladius new genus (Chironomidae, Orthocladiinae) 1HUMBERTO F. MENDES, 2TROND ANDERSEN & 2OLE A. SÆTHER 1Departamento de Biologia-FFCLRP,

ZOOTAXA

A review of Antillocladius Sæther, 1981; Compterosmittia Sæther, 1981 and Litocladius new genus (Chironomidae, Orthocladiinae)

HUMBERTO F. MENDES, TROND ANDERSEN & OLE A. SÆTHER

Magnolia PressAuckland, New Zealand

594

HUMBERTO F. MENDES, TROND ANDERSEN & OLE A. SÆTHER

A review of Antillocladius Sæther, 1981; Compterosmittia Sæther, 1981 and Litocla-

dius new genus (Chironomidae, Orthocladiinae)

(Zootaxa 594)

82 pp.; 30 cm.

6 August 2004

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Accepted by A. Whittington: 10 Jul. 2004; published: 6 Aug. 2004 3

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2004 Magnolia Press

Zootaxa 594: 1–82 (2004) www.mapress.com/zootaxa/

A review of Antillocladius Sæther, 1981; Compterosmittia Sæther, 1981 and Litocladius new genus (Chironomidae, Orthocladiinae)

1HUMBERTO F. MENDES, 2TROND ANDERSEN & 2OLE A. SÆTHER1Departamento de Biologia-FFCLRP, Universidade de São Paulo, Av. Bandeirantes, 3900, 14040-901 Ribeirão Preto, SP, Brazil ([email protected])2Museum of Zoology, Bergen Museum, University of Bergen, Muséplass 3, N-5007 Bergen, Norway ([email protected], [email protected])

Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6List of abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Key to males of some orthoclad genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22Antillocladius Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23

Key to the males of Antillocladius Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25Key to known females of Antillocladius Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26Key to known pupae of Antillocladius Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27Key to known larvae of Antillocladius Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27

Antillocladius antecalvus Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27Antillocladius arcuatus Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29Antillocladius biota new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30Antillocladius calakmulensis new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32Antillocladius folius new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34Antillocladius herradurus new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39Antillocladius musci new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .41Antillocladius pluspilalus Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48Antillocladius scalpellatus Wang and Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .49Antillocladius skartveiti Andersen and Contreras-Ramos . . . . . . . . . . . . . . . . . . . . . . . .49Antillocladius sooretama new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .50Antillocladius ubatuba new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .52Antillocladius venequatoriensis new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54Antillocladius zempoalensis new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .57Antillocladius zhengi Wang and Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

Compterosmittia Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59Key to the males of Compterosmittia Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .61

MENDES ET AL.4 © 2004 Magnolia Press

594ZOOTAXA Compterosmittia aberrans new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .62

Compterosmittia berui new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .64Compterosmittia claggi (Tokunaga) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66Compterosmittia croizati new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .67Compterosmittia dentispina Sæther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .69Compterosmittia nerius (Curran) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .69Compterosmittia pectinata (Freeman) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .69Compterosmittia pittieri new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .70Compterosmittia tuberculifera Tokunaga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .72Compterosmittia virga Wang . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

Litocladius new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .72Litocladius mateusi new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .74

Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .79Biology of the Antillocladius, Compterosmittia and Litocladius species . . . . . . . . . . . . . . . . . . .79Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .80References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

Abstract

A parsimony analysis of recently collected species sharing several features such as scalpellateacrostichals, often setae apically on the wing membrane and often strongly extended costa, togetherwith 38 genera show that the collected species can be assigned to Antillocladius Sæther, 1981,Compterosmittia Sæther, 1981, and one new genus, Litocladius. Nine new species of Antillocladiusare described and figured as male imagines: A. calakmulensis, A. herradurus and A. zempoalensisfrom Mexico; A. venequatoriensis from Ecuador and Venezuela; A. ubatuba from Brazil and Vene-zuela; and A. biota, A. folius, A. musci, and A. sooretama from Brazil. The female of A. musci, thepupae of A. antecalvus Sæther, A. folius, and A. musci, and the larvae of A. folius and A. musci arealso described and figured. New records of A. antecalvus Sæther from Brazil and Venezuela; of A.arcuatus Sæther from Brazil, Mexico and Venezuela; A. pluspilalus Sæther from Ecuador and Mex-ico and of A. zhengi Wang and Sæther from Thailand are given. The genus Antillocladius Sæther,1981, originally described from the British West Indies, now includes 15 species from North, Cen-tral and South America, Russia, China and Thailand. Keys to all known males, females, pupae andlarvae are given. Four new species of Compterosmittia are described and figured as male imagines:C. aberrans from Costa Rica; C. croizati from Brazil and Venezuela; C. pittieri from Venezuela;and C. berui from Brazil. The genus Compterosmittia Sæther, 1981, originally described from theBritish West Indies, now includes 8 to 10 species from North, Central and South America, Austra-lia, Oceania and Southeast Asia. A key to male imagines is given. The new genus, Litocladius,includes a single species, L. mateusi, described as male, female and pupa. The immatures of allthree genera are terrestrial or associated with phytotelmata, and notes on their biology and larvalhabitats are included.

Key words: Chironomidae, Orthocladiinae, Antillocladius, Compterosmittia, Litocladius, Nearctic,Neotropical, Oriental, Palaearctic, systematics

© 2004 Magnolia Press 5ORTHOCLADIINAE

594ZOOTAXAIntroduction

The Neotropical chironomid fauna contains numerous species that are difficult to placeamongst the previously described, mostly Holarctic genera. A number of recently col-lected species shares several features such as scalpellate acrostichals, often setae apicallyon the wing membrane, often strongly extended costa etc. Most of these features are foundin the genus Antillocladius Sæther. However, also Paralimnophyes Brundin, andCompterosmittia Sæther, combine scalpellate acrostichals with well-developed anal pointand extended costa. At least two of the mentioned species appears to belong in Compteros-mittia, but lacks the characteristic comb-like megaseta. Others clearly belong in Antillo-cladius and a few even have associated immatures confirming the placement. However,also among the immatures the generic positions warrant examination. One pupa, forinstance, is quite similar to the pupa of Antillocladius pluspilalus Sæther, but has lateralexpansions of the anal lobe. Another lacks the characteristic thorn-like macrosetae on theanal lobe making it more similar to Gymnometriocnemus Goetghebuer s.str. Nevertheless,most of the species appear to belong in, or at least near, Antillocladius or Compterosmittia.In order to ensure the phylogenetic position of these species we did a parsimony analysisentering all known species of Antillocladius and Compterosmittia separately together withthese species in a data matrix containing a high number of orthoclad genera.

The genus Antillocladius Sæther, 1981, is distributed in the Nearctic, Neotropical,Palaearctic (East Asia) and Oriental biogeographical regions. The genus was erected bySæther (1981) based on A. antecalvus Sæther, 1981, from the British West Indies. Subse-quently, Sæther (1982) described two new species from the southeastern United States andemended the genus. Wang and Sæther (1993) described two additional new species fromPalaearctic and Oriental China. Recently Andersen and Contreras-Ramos (1999) describeda sixth species from Ecuador. The genus thus comprises A. antecalvus from St. Vincent; A.arcuatus Sæther, 1982 and A. pluspilalus Sæther, 1982 both from South Carolina in USA;A. scalpellatus Wang and Sæther, 1993 from the Guangdong Province in China; A. zhengiWang and Sæther, 1993 from the Hainan Province in China and A. skartveiti Andersen andContreras-Ramos, 1999 from Ecuador. Spies and Reiss (1996) recorded A. pluspilalusfrom Nicaragua. The female of A. antecalvus was described by Sæther (1981), while theimmatures of A. pluspilalus were described by Sæther (1984), indicating that the genus isrelated to Bryophaenocladius Thienemann and Gymnometriocnemus Goethgebuer.

The genus Compterosmittia originally was described from the British West Indiesbased on C. dentispina Sæther, 1981. C. clavigera later was described from South Carolina(Sæther 1982). The species was shown by Cranston and Oliver (1988) to be a junior syn-onym of C. nerius (Curran, 1930). Outside the New World the genus has been found inAustralia [C. pectinata (Freeman, 1961)], Oceania [C. claggi (Tokunaga, 1964), C. tuber-culifera (Tokunaga, 1964)] and the Oriental Region (Cranston & Martin 1989; Cranston &Kitching 1995; Wang 1998). Compterosmittia virga Wang, 1998 is a doubtful member ofthe genus since it has a virga consisting of numerous spines.


594ZOOTAXA Material and Methods

The material examined was mounted on slides following the procedure outlined by Sæther(1969). The general terminology follows Sæther (1980). Measurements are given asranges, followed by the mean when 4 or more specimens are measured, followed by thenumber of specimens measured in parenthesis.

The holotypes of the new Brazilian species are housed in the Museu de Zoologia daUniversidade de São Paulo, São Paulo, Brazil (MZUSP), except for A. sooretama, whichis in the Entomological Collection of the Instituto Oswaldo Cruz, Rio de Janeiro, Brazil(FIOCruz). Holotypes of the Venezuelan species are deposited at the Museo del Institutode Zoología Agrícola “Francisco Fernández Yépes” (MIZA), Faculdad de Agronomia,Universidad Central de Venezuela, Maracay, Venezuela. Other holotypes are in theMuseum of Zoology, Bergen, Norway (ZMBN). Paratypes are deposited in the museum ascited in the type material section of the description.

List of abbreviations

FIOCruz: Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil.MIZA: Museo del Instituto de Zoología Agrícola “Francisco Fernández Yépes”, Fac-

uldad de Agronomia, Universidad Central de Venezuela, Maracay, Venezuela.MZUSP: Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil.UFSCar: Laboratório de Entomologia Aquática da Universidade Federal de São Carlos,

São Carlos, Brazil.ZMBN: Museum of Zoology, Bergen Museum, University of Bergen, Norway.ZSM: Zoologische Staatssammlung, Munich, Germany.

Phylogeny

Assessments of phylogenetic relationship involved compilation of a data matrix for 158characters in 38 genera and 25 species, each species entered separately for Antillocladiusand Compterosmittia as well as the new preliminary unassigned species (Tables 1, 2). Theimmatures of Compterosmittia have not been assigned to species with certainty. It is hereassumed that the Australian immatures described by Cranston and Kitching (1995) belongto the only described Australian species, C. pectinata (Freeman); and that the somewhatdifferent immatures from Hong Kong belong to C. claggi (Tokunaga) which again appearto be a junior synonym of C. nerius (Curran). Diplocladius Kieffer, Propsilocerus Kiefferand Brillia Kieffer combined were used as outgroup. Data was analysed under parsimonywith PAUP 4.0b10 (Swofford 1998) operating on a Macintosh, and employing 1,000 ran-dom addition sequence replicates. Only for the multistate characters 17, 25, 27, 28, 30, 33,34, 128 and 138 can a sound rationale be given for ordering the states.


594ZOOTAXATABLE 1 Characters and states used in the phylogenetic analyses. For some characters polymor-

phies are scored as apomorphies as they otherwise would be uninformative or appear constant.

Imagines1. Antennal ratio: (0) sometimes higher than 0.9 and always higher than 0.45; (1) sometimes lower

than 0.46 and always lower than 0.9.2. Number of flagellomeres: (0) always13; (1) sometimes fewer.3. Male antennal apex: (0) without subapical seta; (1) with.4. Female antennal apex: (0) without subapical seta; (1) with. 5. Eyes: (0) bare, at most partly pubescent; (1) hairy or pubescent.6. Palpomeres: (0) always 5 palpomeres of normal length; (1) at least sometimes 4 or less pal-

pomeres or palpomeres strongly reduced in length.7. Sensilla clavata of palpomeres: (0) palpomere 3 at most with few sensilla in one group, pal-

pomere 4 without sensilla; (1) at least female with sensilla in more than one group on pal-pomere 3 and palpomere 4 usually also with at least sensilla clavata, or, in Prodiamesa andChaetocladius, numerous sensilla clavata at well developed sensillum coeloconicum.

8. Dorsomedian eye elongation: (0) moderately to well developed; (1) absent or very weak.9. Temporals: (0) inner verticals present or replaced by frontals, usually more outer verticals; (1)

inner verticals absent, outer verticals usually few.10. Tentorium: (0) male tentorium normal (less than 7 times as long as wide); (1) at least some-

times female like (more than 7 times as long as wide).11. Antepronotal lobes: (0) broad, collar like, at most slightly narrowed medially; (1) distinctly nar-

rowed medially.12. Dorsal antepronotals: (0) absent; (1) present.13. Humeral pit: (0) inconspicuous; (1) consisting of several smaller areas; (2) conspicuous, oval.14. Dorsocentrals: (0) uniserial anterior; (1) bi- multiserial anterior.15. Dorsocentrals: (0) uniserial posterior; (1) bi- multiserial posterior.16. Acrostichals: (0) moderately long to long and strong, (1) short, or absent.17. Acrostichals: (0) starting in front; (1) starting some distance from antepronotum; (2) in centre

of scutum. (Absence scored as (?))18. Acrostichals: (0) simple or absent; (1) anterior simple, posterior scalpellate; (2) all scalpellate.19. Prealars: (0) 1–5; (1) 6 or more.20. Supraalar(s): (0) present; (1) absent.21. Setae of preepisternum and/or anepisternum: (0) present; (1) absent.22. Scutellars: (0) uniserial; (1) bi- multiserial.23. Postnotum: (0) bare; (1) sometimes with setae. 24. Wing spots: (0) absent; (1) sometimes present.25. Setae in cells in male: (0) absent in all cells; (1) present in cell r4+5; (2) present in cell r4+5 and

m1+2; (3) present also in cell m3+4; (4) present also in cell cu+cu1.

26. Wing membrane: (0) not to moderately punctated; (1) coarsely punctated.27. Anal lobe: (0) well developed, right-angled or protruding; (1) weak to distinct, but not right-

angled or protruding; (2) reduced or wing cuneiform.28. Costal extension in male: (0) absent or short, less than 0.04 wing length; (1) 0.04–0.06 wing

length; (2) 0.07–0.08 wing length; (3) 0.09 or more of wing length.29. R4+5: (0) ends above or distal to apex of M3+4; (1) ends proximal to apex of M3+4.

.....continued on the next page


594ZOOTAXA TABLE 1 (continued)

30. Cu1: (0) not sinuous; (1) slightly sinuous; (2) strongly sinuous.

31. VR: (0) < 1.06; (1) 1.06–1.40; (2) > 1.40.32. Anal veins: (0) An1 extending well beyond cubital fork and An2 conspicuous; (1) anal veins

shorter.33. R veins: (0) setae present on R, R1 and usually R4+5 in both sexes; (1) setae present on R,

absent on R1 and often R4+5 of male, at most absent on R1 in female; (2) setae absent on R of

male, present in female; (3) setae absent on R and R1 of both sexes, at most 1 apical seta on

R4+5.

34. Setae on veins in male: (0) absent on all except brachiolum and on C- extension and apically onM1+2 when wing hairy; (1) present on R, R1 or R4+5; (2) present on R veins and M, M1+2 or

M3+4.

35. Squama: (0) with setae; (1) bare.36. Leg ratio of male (LR1): (0) sometimes higher than 0.8, and at least higher than 0.5; 1) 0.5–0.8;

(2) sometimes lower than 0.5 and at least lower than 0.8.37. Tibial spurs: (0) often with distinct lateral denticles; (1) denticles always indistinct or absent.38. Inner tibial spur of hind leg: (0) At least 1/2 as long as outer spur; (1) shorter; (2) absent with

also second spur of mid leg absent.39. Hind tibial comb: (0) well developed, occupying full width of tibia,; (1) weak or absent.40. Hind tibial comb: (0) with less than13 setae; (1) conspicuous with 13 or more setae of which

some about as long as longest spur.41. Pseudospurs: (0) present; (1) absent.42. Sensilla chaeticae of tarsi: (0) present; (1) absent.43. Pulvilli: (0) present and distinct; (1) absent or vestigial, less than 1/2 claw length.44. Anal point: (0) absent; (1) small and anterior on tergite; (2) represented by hump-like extension

of tergite or if absent represented by some stronger median setae; (3) well set off and posterioron tergite.

45. Anal point: (0) not with spatulate microtrichiae- free apex; (1) often with small or large spatu-late, microtrichiae-free apex.

46. Anal point: (0) not very broad and rounded to bluntly triangular; (1) conspicuously broad androunded to bluntly triangular.

47. Anal point: (0) not long, when moderately sized triangular; (1) moderately sized, triangularwith concave margins; (2) conspicuously long, triangular with nearly straight sides; (3) long,nearly parallel-sided apically with blunt apex.

48. Setae on anal point: (0) normal, hair-like, relatively few or anal point absent; (1) strong, stiff,bristle-like or lamellate, numerous; (2) short, numerous, often directed anteriorly or laterally.

49. Superior volsella: (0) present; (1) absent.

50. Superior volsella: (0) absent or with setae and/or microtrichiae, not plate-like; (1) bare andplate-like.

51. Inferior volsella: (0) at least some species with anterior dorsal, often triangular or digitiformpart and a usually more rounded, small to large ventral, posterior part, sometimes separatedfrom dorsal part, occasionally more than 2 parts simple, well developed to reduced; (1) differ-ent.



594ZOOTAXATABLE 1 (continued)

52. Inferior volsella: (0) without microtrichia-free digitiform projection; (1) with anterior microt-richia-free, digitiform anterior projection and long rounded, low to prominent posterior lobe.

53. Inferior volsella: (0) not short, simple and projecting; (1) simple, triangular to tongue-shapedor digitiform, sometimes hooked, occasionally spiniform.

54. Inferior volsella: (0) not adpressed to gonocoxite; (1) simple, elongate, adpressed to gonocox-ite, sometimes pediform with free end.

55. Inferior volsella: (0) not consisting of long posteromedially directed lobe; (1) consisting of aposteromedially directed, simple or apically bifid lobe.

56. Inferior volsella: (0) not consisting of long bifid lobe; (1) consisting of a apically bifid lobe.57. Inferior volsella: (0) not conspicuously set off; (1) circular, conspicuously set off, sometimes

without additional rounded posterior extension. 58. Inferior volsella: (0) not conspicuously set off; (1) circular, conspicuously set off, with addi-

tional rounded posterior extension. 59. Gonostylus: (0) simple; (1) double.60. Heel of gonostylus: (0) absent; (1) present.61. Transverse sternapodeme: (0) convex; (1) straight or concave; (2) absent, sternapodeme trian-

gular.62. Oral projections of transverse sternapodeme: (0) strongly developed; (1) weak to moderately

developed; (2) absent. 63. Crista dorsalis: (0) evident, triangular or rounded preapical; (1) elongate, low; (2) not evident/

weak.64. Megaseta: (0) present; (1) absent.65. Megaseta: (0) not dentate; (1) dentate.66. Virga: (0) in some species consisting of cluster of at least 6 short spines or two groups of very

strong spines; (1) virga not consisting of cluster or groups of spines.67. Virga: (0) not consisting of narrow or broad field of scattered spinules; (1) in some species con-

sisting of narrow or broad field of scattered spinules.68. Virga: (0) not consisting of single plate; (1) in some species consisting of single plate.69. Virga: (0) virga not consisting of 2–4 short spines; (1) in some species consisting of 2–4 short,

often partly fused, sometimes plate-like spines without distinct lateral lamellae.70. Virga: (0) virga without lateral lamellae; (1) in some species consisting of 2–12 median, usually

fused, spines, and distinct lateral lamellae. 71. Virga: (0) virga clearly shorter than half the length of phallapodeme or without lateral lamellae;

(1) clearly longer than half apodeme length and with lateral lamellae.72. Virga: (0) virga not consisting of long spines without lamellae; (1) in some species consisting

of spines at least nearly as long as half the length of phallapodeme, but without lateral lamel-lae.

73. Virga: (0) virga not consisting of long spines without lamellae; (1) in some species consistingof spines clearly longer than half the length of phallapodeme, without lateral lamellae.

74. Gonocoxapodeme: (0) absent, short and straight or evenly curved and ending at base of gonap-ophysis; (1) continuing basal of vagina or at least past base of gonapophysis.

75. Female tergite IX: (0) undivided; (1) divided by caudal concavity or notch; (2) divided into twosetigerous protrusions.




76. Female gonocoxite IX: (0) large, projecting; (1) moderately developed to reduced.77. Female gonocoxite IX: (0) with long setae only; (1) with some long and some short setae; (2)

with short setae only.78. Gonapophysis VIII: (0) undivided; (1) divided with ventrolateral lobe much smaller and more

or less brush-like; (2) divided, with lobes of about same size; (3) divided with dorsomesallobe smaller and with anterior rounded projection; (4) divided with dorsomesal lobe narrow,often line-like.

79. Apodeme lobe: (0) not apparent; (1) well developed, but not meeting at mid line and withmicrotrichiae; (2) meeting at mid line and/or with microtrichiae or fully sclerotized.

80. Number of seminal capsules: (0) 3; (1) 2.81. Seminal capsules: (0) spherical to ovoid, small or of normal size; (1) large, spherical to elon-

gate ovoid.82. Seminal capsules: (0) at least partly coloured; (1) often completely pale. Polymorphies are scored as synapomorphies as no genera have all included species with pale

capsules and the character otherwise would be uninformative.83. Opening of spermathecal ducts: (0) separate; (1) common.84. Spermathecal ducts: (0) not fused; (1) partly fused ducts before common opening.85. Bulbs of spermathecal ducts: (0) absent; (1) present.86. Spermathecal ducts: (0) straight; (1) with bend or loop.

Pupa 87. Frontal apotome: (0) without warts or tubercles; (1) with warts or tubercles.88. Frontal setae: (0) present; (1) absent.89. Thoracic horn: (0) present; (1) absent.90. Thoracic horn: (0) not rounded to ovoid; (1) mostly rounded to elongate ovoid.

Polymorphies are scored as synapomorphies as only Botryocladius have all included specieswith rounded thoracic horn and the character otherwise would be uninformative.

91. Thoracic horn: (0) not with bulbous base and narrow apical portion; (1) with. Eukiefferiella is scored as (1) since most species have a thoracic horn with bulbous base andspecies without thoracic horn apparently has this secondarily reduced.

92. Thorax: (0) mostly smooth to wrinkled; (1) mostly tuberculose or spinulose.93. Antennal sheath pearl row: (0) absent; (1) present.94. Wing sheath: (0) without pearls; (1) with.

95. Dorsocentrals: (0) anterior 2 and posterior 2(3) paired, anterior 3 grouped, all in row or 2–3dorsocentrals only; (1) posterior 3 grouped or all 4 together.

96. Tergites II–VIII: (0) without posterior spine, or tubercle row(s), but may have caudal hooklets;(1) some with spines or tubercles.

97. Median field of tergite IV: (0) without discrete spine patch(es) or row(s); (1) with.98. Tergite I: (0) without posterior spine row(s); (1) sometimes with.99. Tergal armament: (0) lacks elongate, needle-like spines; (1) includes elongate, parallel-sided,

needle-like spines in some species.

100. Sternites II–VII: (0) without posterior spine row(s), but may have caudal hooklets; (1) somewith spines or tubercles.

101. Sternites or sternal conjunctives: (0) without caudal hooklets; (1) sometimes with......continued on the next page



102. Male sternite VIII: (0) without posterior spine or tubercle row(s); (1) with.103. Sternite II or II and III: (0) without anterior spine group; (1) sometimes with.104. Tergites and sternites: (0) with single or no posterior row of spines; (1) at least some with dou-

ble to multiple row of spines.105. Tergite III: (0) without caudal hooklets; (1) with minute or conspicuous caudal hooklets.

Cricotopus absurdus type without thoracic horn and with pearl row, another type with hook-lets on T III and PSB on II and III.

106. Tergite IV: (0) without caudal hooklets; (1) with caudal hooklets.107. Tergite V: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-

like anteriorly directed; (1) with minute or conspicuous caudal hooklets.108. Tergite VI: (0) without caudal hooklets, although rows of conjunctival spinules may be hook-

like anteriorly directed; (1) sometimes with caudal hooklets.109. Caudal hooklets of tergite II: (0) present, (1) absent.110. Pedes spurii B: (0) present; (1) absent.111. Pedes spurii A on sternite IV: (0) present; (1) absent.112. Pedes spurii A on sternite VI: (0) present; (1) absent.113. Tergal conjunctives or posterior of tergal spines: (0) without spinules, but may have hooklets

in single row; (1) with spinules which may be hooklet-like recurved.114. Spinules on tergal conjunctives: (0) absent or not hooklet-like recurved or anteriorly directed;

(1) hooklet-like recurved or anteriorly directed spinules in about 3 rows.115. Taeniate L setae: (0) present; (1) absent.116. Segment VIII: (0) with more than 1 L seta; (1) with at most 1 L seta.

117. Posterolateral corners of tergites VI–VIII: (0) without embedded spines; (1) at least tergiteVIII with in most species.

118. Anal lobe: (0) not extended into projections; (1) extended posteriorly into cylindrical projec-tions with macrosetae at apex.

119. Anal lobe: (0) not with apical projections or extensions; (1) with apical spurs or extended dis-tal of macrosetae.

120. Anal lobe: (0) without lateral expansions; (1) sometimes with.121. Apical spines of anal lobe: (0) absent; (1) present.122. Anal lobe: (0) with fringe of setae; (1) without fringe of setae.123. Inner margin of anal lobe: (0) without long seta; (1) with.124. Anal macrosetae: (0) not short and spine-like, but may be short and hair-like or absent; (1)

short and spine- or thorn-like.

125. Width of anal macrosetae or apical spines: (0) < 5 µm; (1) > 4 µm.126. No. of anal macrosetae: (0) 3 or more; (1) 2; (2) 0–1.127. Genital sac: (0) not reaching apex of anal lobe; (1) reaching somewhat beyond lobe; (2)

reaching much beyond anal lobe.

Larva128. Antenna: (0) with 6–7 segments; (1) with 5; (2) with 3–4.129. Ultimate antennal segment: (0) normal; (1) whip- or thread-like.130. Antenna: (0) reduced, less than half mandible length; (1) 1/3 as long as head capsule or

shorter, but not reduced; (2) longer. .....continued on the next page



131. Second antennal segment: (0) undivided, fully sclerotised; (1) divided or partly unsclerotised.132. Lauterborn organs: (0) moderately large to well developed; (1) weak or absent.133. Antennal blade: (0) short to moderate length, shorter than flagellum when antenna of normal

length; (1) conspicuous, longer than flagellum except when flagellum extremely long.134. S I: (0) plumose, branched, toothed or palmate; (1) bifid or simple.135. S I: (0) not palmate; (1) palmate.136. S I: (0) not bifid; (1) bifid.137. S I: (0) not simple; (1) simple.138. Labral lamella: (0) with pectinate, plumose or rugose apex, mostly well developed; (1) weak,

no apical teeth or plumosity; (2) absent.139. Pecten epipharyngis: (0) consisting of 3 simple scales or single scale; (1) of several teeth.140. Chaetulae laterales: (0) simple or reduced; (1) at least one serrated or plumose.141. Premandible: (0) simple; (1) with 2 or more teeth.142. Premandibular brush: (0) present; (1) absent.143. Mandible: (0) with 2–3 inner teeth; (1) with 4 or more.144. Mola of mandible: (0) smooth; (1) with teeth or spines.145. Seta interna of mandible: (0) with smooth, slightly plumose laterally or apically, or serrate

branches; (1) branches conspicuously branched; (2) seta interna absent.146. Median tooth of mentum: (0) single; (1) double, bifid or with several median teeth.147. Lateral teeth of mentum: (0) 7 or more pairs; (1) 5–6 pairs; (2) fewer.148. Lateral teeth of mentum: (0) outer tooth not larger or higher than one of the inner teeth; (1)

clearly larger or higher.149. Ventromental plates: (0) well developed, extending past lateral teeth on flattened mentum,

sometimes double; (1) reduced or weak.150. Ventromental plates: (0) never double; (1) at least sometimes double.151. Ventromental plates: (0) without setae (beard) underneath; (1) with setae underneath.152. Setae submenti: (0) situated at level of base of outer lateral tooth or higher; (1) lower.153. Claws of anterior parapods: (0) with relatively distinct teeth; (1) smooth or teeth very indis-

tinct.154. Procercus: (0) well developed; (1) reduced or absent.155. Anal setae: (0) 5 or more setae, none conspicuously long; (1) 3–4 setae, none conspicuously

long; (2) 0–2 not conspicuously long anal setae; (3) 2 or more setae with 1 or 2 conspicuouslylong.

156. Supraanal setae: (0) weak or absent, shorter than 1/2 length of anal setae when these welldeveloped or shorter than 2/3 when anal setae short; (1) well developed, longer than 1/2length of anal setae when anal setae long, longer than 2/3 when anal setae short.

157. Posterior parapods: (0) well developed; (1) small, digitiform; (2) absent to weak but not digi-tiform.

158. Anal tubules: (0) at least 1/2 length posterior parapods; (1) shorter than 1/2 length posteriorparapods or these absent; (2) conspicuously long and narrow.

159. Anal segment: (0) not projecting posterior over posterior parapods or bent at right angles toaxis of rest of body; (1) projecting posterior over posterior parapods or bent at right angles toaxis of rest of body.


594ZOOTAXATABLE 2 Character states for characters 1–159 in some genera of Orthocladiinae. Polymorphies: A = 0&1, B =

0&1&2, C = 1&2, D = 1&2&3, E = 0&2, F = 0&3, G = 2&3, H = 0&1&2&3, I = 1&3, J = 0&1&3, K = 0&1&2&3&4, L = 0&4, M = 2&4, N

= 3&4.Character no.: 1 2 3 4 5 6 7 8 9 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2

0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7Aagaardia 1 0 0 0 0 0 0 1 0 0 0 0 0 A A 0 0 0 0 1 1 A 0 0 0 1 1Apometriocnemus 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 1 0 0 0 4 1 1Botryocladius A 0 0 0 0 0 0 1 0 0 0 0 2 0 0 1 2 2 0 0 1 0 0 0 0 0 ABrillia 0 0 1 1 0 0 A 0 0 0 0 1 0 A 1 1 ? 0 1 0 1 1 0 0 4 1 0Bryophaenocladius 0 0 A 0 0 0 0 A 0 0 0 0 0 A A A 0 0 A 1 1 0 0 0 0 A AChaetocladius A 1 A 0 A 0 A A 0 1 0 0 2 A A 1 0 0 A A 1 0 0 0 0 1 ACorynoneura A 1 0 0 0 0 0 0 1 0 1 0 0 0 0 1 ? 0 0 1 1 0 0 0 0 0 2Cricotopus A 1 A A 1 0 0 A A 0 0 0 A A A 1 0 0 A A A A 1 1 0 A ADiplocladius 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0Doithrix 0 0 1 ? 0 0 1 1 0 0 0 0 0 1 0 0 0 0 1 1 1 1 0 0 0 A 0Eukiefferiella A 1 0 1 A 0 0 A A 0 0 0 0 0 0 1 E 0 0 1 1 0 0 0 0 0 0Georthocladius s.str 0 0 1 0 0 0 1 1 0 0 0 0 0 A A 0 0 0 1 1 A A 0 0 0 A 0 Gymnometriocnemus0 0 1 0 0 0 0 1 0 0 1 0 0 ? A 1 1 0 1 1 1 0 0 0 N 1 1Hanocladius 1 1 0 ? 0 0 0 0 0 0 0 0 0 0 0 1 2 2 0 1 1 0 0 0 0 1 2Heleniella 0 0 0 0 1 0 0 1 0 0 0 0 0 1 1 1 ? 0 1 1 0 1 1 1 0 0 AHeterotanytarsus 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 2 0 0 1 0 0 0 0 A 0 1Heterotrissocladius 0 0 0 1 0 0 0 0 0 0 0 0 0 A A A 0 0 A 1 1 A 0 0 K A 0Hydrobaenus 0 1 0 0 0 1 0 A 0 0 0 0 0 A 0 1 1 0 A 1 A A 0 0 0 A AKrenosmittia 1 1 0 0 0 0 0 1 1 0 1 0 0 0 0 1 ? 0 0 1 1 0 0 0 0 0 2Limnophyes A 1 0 0 0 0 0 1 A 0 0 1 2 A A 1 1 2 1 0 0 0 1 0 0 1 1Lopescladius A 1 0 ? 1 0 0 1 1 1 1 0 1 0 0 1 ? 0 0 1 1 0 0 0 0 0 1Metriocnemus A 1 A A 0 0 0 A 0 1 0 0 0 1 1 0 0 0 1 0 A 1 0 0 N A AOrthocladius 0 0 0 0 0 0 0 1 0 0 0 0 E ? A 1 0 0 A A A A 0 1 0 0 AParachaetocladius 0 0 1 0 0 0 1 1 0 0 0 0 0 A 0 1 ? 0 1 1 A 0 0 0 0 A 0Parakiefferiella A 1 0 ? 0 0 0 1 1 0 1 0 0 0 0 1 ? 0 0 1 1 0 0 0 0 0 1Paralimnophyes 0 0 1 ? 0 0 ? 1 0 0 0 0 0 1 1 0 2 2 0 0 1 0 0 0 0 1 1Parametriocnemus A 1 0 0 0 0 A 0 0 0 0 0 0 A A 0 0 0 A 1 1 A 0 0 N A BParasmittia 0 0 1 ? 0 0 ? 0 0 0 0 0 0 1 A 1 0 0 1 1 1 1 0 0 3 0 1Propsilocerus 0 1 0 0 0 1 A 1 A 0 0 0 0 A A 1 ? 0 A 1 A A 0 0 0 1 0Psectrocladius 0 0 0 A 0 0 0 1 0 0 0 0 0 A A A 0 0 A 1 1 A 0 0 0 0 0Pseudorthocladius 0 0 1 0 0 0 1 A 0 0 0 0 0 A A 0 0 0 1 1 1 1 0 0 L 0 1Pseudosmittia A 0 A A 0 0 A 1 A 0 A 0 0 0 A 1 2 0 A A 1 0 0 0 0 0 BRheocricotopus A 0 0 0 1 0 0 1 A 0 0 0 C A 0 1 A 0 A 1 1 0 0 0 0 A ASmittia 0 0 1 1 1 0 0 1 0 0 0 0 0 0 0 1 ? 0 1 A 1 0 0 0 0 0 AStictocladius A 0 0 1 1 0 0 1 1 1 A 0 C A A 1 ? 0 A 1 1 A 1 1 0 0 ATvetenia 0 1 0 0 0 0 0 1 A 0 0 0 0 0 0 1 ? 0 0 1 1 0 0 0 0 0 AUnniella 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 2 0 1 1 0 0 0 0 0 1Zalutschia 0 0 0 0 0 0 0 A 0 0 0 0 0 0 0 1 1 0 A 1 A 0 0 0 0 0 Baberrans 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 2 0 1 1 0 0 0 1 0 2antecalvus 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 1 0 1 1 0 0 0 1 0 1arcuatus 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 0 0 0berui 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 2 0 1 1 0 0 0 A 0 2biota 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 2 0 1 1 0 0 0 0 0 2calakmulensis 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 2 0 1 1 0 0 0 1 0 0croizati 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 2 0 1 1 0 0 0 0 0 2dentispina 1 ? 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 2 0 0 1 0 0 0 1 0 2folius 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 0 0 2herradurus 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 1 0 1mateusi 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 0 1 0 0 0 0 0 1musci 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 2 0 1 1 0 0 0 1 0 0nerius 0 ? 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 2 0 0 1 0 0 0 A 0 1pectinata 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 2 0 ? 1 0 0 0 1 0 1pittieri 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 2 2 0 0 1 0 0 0 2 0 2pluspilalus 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 3 0 0scalpellatus 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 2 0 0skartveiti 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 2 0 1 1 0 0 0 3 0 2sooretama 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 2 0 1 1 0 0 0 0 0 1tuberculifera 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 ? 2 0 ? 1 0 0 0 0 0 1ubatuba 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 2 0 1 1 0 0 0 0 0 0venequatoriensis 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 2 0 1 1 0 0 0 4 0 1virga 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 2 0 1 1 0 0 0 0 0 2zempoalensis 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 0 0 2zhengi 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 1 0 0



Character no.: 2 2 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 55 5 5 58 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 01 2 3 4

Aagaardia C 0 A 1 0 1 1 1 2 1 1 0 0 1 0 1 3 0 0 0 0 1 00 0 0 0Apometriocnemus 2 0 2 1 0 0 2 0 1 0 1 0 0 1 1 1 2 0 0 0 0 1 00 0 0 0Botryocladius 0 0 0 1 1 1 1 0 1 0 1 0 0 1 1 1 1 0 0 0 0 0 00 0 0 0Brillia 0 0 0 1 0 0 2 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 01 0 0 1Bryophaenocladius B 0 0 1 0 0 1 A 0 0 A A 0 A 1 1 F 0 A 0 0 A 0 0 1 1 1Chaetocladius A 0 A 0 0 A A A 1 0 1 0 1 0 0 A 3 0 A H E 1 0 0 0 0 0Corynoneura 0 1 A 2 1 3 0 1 0 1 C 0 A 1 1 1 0 0 0 0 0 0 10 0 0 0Cricotopus A 0 0 1 1 B C 0 B 0 C 0 0 1 0 A J 0 0 0 0 A 00 0 1 0Diplocladius 0 0 0 0 0 1 1 1 1 1 0 0 0 1 1 1 3 0 0 0 0 1 01 0 0 0Doithrix 0 0 1 1 1 0 1 0 1 1 1 1 0 1 1 0 2 0 0 G 1 1 01 0 1 0Eukiefferiella 0 A 0 1 A D A 0 1 0 C 0 0 A A 1 0 0 0 0 0 1 0 0 0 1 0Georthocladius s.str 0 0 A 1 0 A 1 0 1 1 A 0 0 0 1 0 2 0 0 0 1 1 00 0 1 0 GymnometriocnemusG 0 A 1 0 0 2 1 2 1 1 0 0 1 1 1 2 0 0 0 A 1 00 0 0 0Hanocladius 3 0 2 C 1 3 0 1 0 0 0 0 0 1 1 1 3 0 0 1 0 1 00 0 0 0Heleniella 1 0 2 1 1 1 1 1 1 1 A 0 0 1 A 1 F 0 0 0 0 1 00 0 1 0Heterotanytarsus A 0 0 C 0 0 C 0 1 0 1 1 0 1 A 1 3 0 0 0 0 A 00 0 1 0Heterotrissocladius 0 0 0 1 0 0 C 0 0 1 1 0 0 1 A 1 3 1 0 I C 1 00 0 0 0Hydrobaenus A 0 0 A 0 A 1 0 C 1 1 0 0 0 0 1 3 0 0 0 A 1 00 0 0 0Krenosmittia 3 1 C 1 1 1 0 1 2 1 1 0 0 1 1 1 G 0 0 0 0 1 00 0 1 0Limnophyes C 0 1 1 0 C A A 2 1 1 0 0 1 A 1 2 0 0 0 0 1 00 0 1 0Lopescladius 0 1 0 1 0 3 0 1 ? 1 1 0 0 1 1 0 0 0 0 0 0 1 0 ? 0 1 0Metriocnemus C A 0 1 0 0 2 0 0 1 A 0 A 0 1 1 3 0 0 B A 1 00 0 0 0Orthocladius A 0 0 1 0 A 1 0 0 1 0 0 0 A 0 1 3 0 0 1 0 A 00 0 0 0Parachaetocladius 0 0 1 1 0 0 1 0 1 1 A 0 0 A 1 0 2 0 0 0 1 1 00 0 1 0Parakiefferiella C 0 2 1 1 1 1 1 2 1 1 0 A 1 1 1 3 0 1 0 0 A 00 1 1 0Paralimnophyes C 0 0 1 0 1 1 0 ? 1 0 ? ? 1 0 1 3 0 1 1 0 0 01 0 0 1Parametriocnemus C 0 1 1 0 0 C 0 1 1 1 0 0 1 1 1 3 1 0 B 0 1 00 0 1 0Parasmittia 2 0 0 1 1 0 2 1 ? 1 0 ? ? 1 1 1 3 0 1 1 0 1 00 0 0 0Propsilocerus 0 0 0 0 0 1 1 0 1 1 0 1 0 A 0 1 F 0 0 A 0 A 00 1 0 0Psectrocladius 0 0 0 1 0 A 1 0 0 1 B 0 A 0 0 0 F 0 0 B A 1 00 0 1 0Pseudorthocladius A 0 1 1 A 0 C 0 1 1 1 0 0 1 1 0 E 0 0 0 1 1 00 0 1 0Pseudosmittia 0 A B 2 1 D A 1 2 1 1 0 A A A 1 A 0 0 0 0 0 0 0 0 0 0Rheocricotopus 0 0 0 1 0 1 1 A 0 1 1 0 A 1 0 0 3 0 0 C 0 0 10 0 1 0Smittia 2 0 1 1 0 1 1 1 1 1 0 0 0 1 1 1 3 0 1 2 0 1 00 0 1 0Stictocladius A A A 1 0 H A A B 0 E A 0 A 1 1 F 0 0 0 0 1 0 1 0 0 1Tvetenia 0 A 0 1 1 1 1 0 1 1 1 0 0 0 A 1 3 0 0 C 0 1 00 0 1 0Unniella 0 0 2 1 0 1 1 0 1 1 1 0 0 1 0 1 3 0 1 0 0 0 00 0 0 0Zalutschia A 0 0 1 0 1 1 0 0 1 1 0 0 0 0 1 3 0 0 A 0 1 00 0 0 0aberrans 2 0 0 1 1 ? 0 1 1 1 1 0 0 1 1 1 3 0 0 1 1 1 00 1 0 0antecalvus 1 0 0 C 1 1 A 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 01 0 0 0arcuatus 1 0 1 C 1 ? 0 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 01 0 0 0berui 3 0 0 2 1 ? 0 1 1 1 1 0 0 1 1 1 3 0 0 1 0 1 00 1 0 0biota 2 0 0 C 1 ? A 0 1 1 1 0 0 1 1 1 3 0 0 3 2 1 01 0 1 0calakmulensis 0 0 0 1 1 ? 0 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 00 0 0 0croizati 3 0 1 C 1 1 1 1 1 1 1 0 0 1 1 1 3 0 0 1 0 1 00 0 1 0dentispina 3 0 2 2 1 1 1 1 1 1 1 0 0 1 1 1 3 0 0 1 0 1 00 1 0 0folius 2 0 0 2 1 1 A 0 1 1 1 0 0 1 1 1 3 0 0 3 2 1 01 0 1 0herradurus 0 0 0 1 1 ? 0 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 01 0 0 0mateusi 1 0 0 2 1 1 0 0 1 1 1 0 0 1 1 1 3 0 0 2 0 1 00 0 0 0musci 1 0 0 C 1 1 A 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 00 0 0 0nerius 3 0 2 2 1 1 1 1 1 1 1 0 0 1 1 1 3 0 0 1 0 1 00 0 1 0pectinata 3 0 2 2 1 1 1 1 1 1 1 0 0 1 1 1 3 0 0 1 1 1 00 0 1 0pittieri 3 0 0 2 1 0 1 1 1 1 1 0 0 1 1 1 3 0 0 1 0 1 00 1 0 0pluspilalus 1 0 0 1 1 0 2 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 00 0 0 0scalpellatus 0 0 0 1 1 0 2 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 00 0 0 0skartveiti 2 0 0 C 1 A 2 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 01 0 0 0sooretama 0 0 0 C 1 ? 0 0 1 1 1 0 0 1 1 1 3 0 0 3 2 1 01 0 0 0tuberculifera 3 0 0 2 1 1 1 1 1 1 1 0 0 1 1 1 3 0 0 1 0 1 00 0 1 0ubatuba 1 0 0 1 1 ? A 0 1 1 1 0 0 1 1 1 3 0 0 3 2 1 01 0 0 0venequatoriensis 1 0 0 C 1 1 2 0 1 1 1 0 0 1 1 1 3 0 0 2 2 1 00 0 0 0virga G 0 2 C 1 1 1 1 1 1 1 0 0 1 1 1 3 0 0 2 1 1 00 0 1 0zempoalensis 1 0 0 1 1 ? 0 1 1 1 1 0 0 1 1 1 3 0 0 2 1 1 01 0 0 0zhengi 1 0 0 1 1 ? 0 0 1 1 1 0 0 1 1 1 3 0 0 2 1 1 00 0 0 0



Character no.: 5 5 5 5 5 6 6 6 6 66 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8 85 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1

Aagaardia 0 0 0 0 0 1 0 1 E 0 0 0 0 0 0 0 0 0 01 2 1 0 2 1 1 1Apometriocnemus 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?Botryocladius 0 0 0 0 0 0 1 1 1 0 0 0 0 1 1 0 0 0 0 0 0 0 1 1 1 1 0Brillia 0 0 0 0 1 0 0 1 2 1 0 1 0 0 0 0 0 0 0 1 0 0 0 4 1 1 1Bryophaenocladius 1 0 0 0 0 0 0 C 1 0 0 0 0 0 0 0 0 0 01 0 0 0 4 1 1 1Chaetocladius 0 0 0 0 A A 0 1 B A 0 1 0 0 0 0 0 0 0 0 C 1 1 4 1 1 0Corynoneura 0 0 0 0 0 0 2 2 E 0 0 1 0 0 0 0 0 0 00 1 1 1 0 2 1 1Cricotopus 0 0 0 0 A 0 0 1 B 0 0 1 1 0 0 0 0 0 0 0 2 B B 3 1 1 ADiplocladius 1 0 0 0 1 0 0 1 2 1 0 0 0 0 0 0 0 0 0 1 0 0 0 4 1 0 0Doithrix 1 0 0 0 0 0 0 2 2 0 0 1 1 0 0 0 0 0 0 ? ? ? ? ? ? ? ?Eukiefferiella 0 0 0 0 0 0 0 A B 0 0 1 1 0 0 0 0 0 00 0 0 0 0 A 1 1Georthocladius s.str 0 0 0 0 0 1 A 0 C 0 0 1 0 0 0 0 0 0 01 0 0 0 4 1 1 1 Gymnometriocnemus 0 0 0 0 0 0 A C B 0 0 1 0 0 0 1 1 0 01 0 1 1 4 1 1 1Hanocladius 0 0 0 0 0 0 0 1 2 0 0 1 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ?Heleniella 0 0 0 0 0 0 0 A 2 0 0 1 0 0 0 1 1 0 00 2 0 0 4 1 1 0Heterotanytarsus 0 0 0 0 0 A A 1 2 0 0 1 0 0 0 0 0 0 0 1 2 0 1 0 0 1 0Heterotrissocladius 0 0 0 0 0 A 0 1 0 0 0 0 0 0 0 0 0 0 0 1 2 A 0 2 1 1 0Hydrobaenus 0 0 0 0 0 A 0 1 2 0 0 0 0 0 0 0 0 0 0 1 C 1 1 M 1 1 1Krenosmittia 0 0 0 0 0 0 0 1 2 0 0 1 0 0 0 1 1 0 0 0 2 1 1 1 1 1 0Limnophyes 0 0 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 1 B 1 1 4 1 1 0Lopescladius 0 0 0 0 0 0 A C 2 1 0 1 0 0 0 0 0 0 0? ? ? ? ? ? ? ?Metriocnemus 0 0 0 0 0 0 0 A A 0 0 0 0 0 0 0 0 0 0 A A A A 4 1 1 AOrthocladius 0 0 0 0 0 0 0 1 A 0 0 0 0 0 0 0 0 0 0 A 0 0 A 3 1 1 AParachaetocladius 0 0 0 0 0 1 0 1 2 0 0 1 0 0 0 0 0 0 0 0 0 0 0 4 1 1 1Parakiefferiella 0 0 0 0 0 0 0 1 C 0 0 0 0 0 0 1 1 0 00 2 1 1 1 1 1 0Paralimnophyes 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 B 1 1 4 1 1 0Parametriocnemus 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 1 2 1 1 M 1 1 0Parasmittia 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 3 2 1 0Propsilocerus 1 1 0 0 1 0 1 0 1 0 0 1 0 0 0 0 0 0 0 A C 0 0 4 2 0 0Psectrocladius 0 0 0 0 0 0 0 A E 0 0 1 0 1 0 0 0 0 0 A 2 A 1 M C 1 APseudorthocladius 0 0 0 0 0 0 0 1 C 0 0 0 1 0 0 1 1 1 1 1 0 A 1 4 1 1 1Pseudosmittia 0 0 0 0 0 0 A C C 0 0 1 0 1 1 0 0 0 0 0 E 1 A 4 2 1 0Rheocricotopus 1 0 0 0 0 0 0 A A 0 0 1 0 0 0 0 0 0 0 0 E 1 1 4 1 1 1Smittia 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 0 3 2 1 0Stictocladius 0 0 0 0 1 0 A C 2 1 0 1 0 0 0 0 0 0 0 1 2 C C 4 2 1 ATvetenia 0 0 0 0 0 0 0 1 E 0 0 0 0 0 0 0 0 0 01 2 1 1 4 1 1 0Unniella 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 1 0 1 ? 4 1 1 0Zalutschia 0 0 0 0 0 1 1 0 2 0 0 0 0 0 0 0 0 0 0 1 B 1 1 4 1 1 0aberrans 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?antecalvus 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 1 1 1 4 1 1 0arcuatus 0 0 1 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?berui 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?biota 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 1 ? ? ? ? ? ? ? ?calakmulensis 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?croizati 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?dentispina 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 1 2 1 1 4 1 1 1folius 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 1 1 ? ? ? ? ? ? ? ?herradurus 1 1 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 0 ? ? ? ? ? ? ? ?mateusi 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 1 1 0 0 0 2 1 1 4 1 1 0musci 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 1 0 2 1 1 4 1 1 0nerius 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?pectinata 0 0 0 0 0 0 0 1 ? 0 1 1 0 0 0 0 0 0 0? ? ? ? ? ? ? ?pittieri 0 0 0 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?pluspilalus 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 1 ? ? ? ? ? ? ? ?scalpellatus 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 ? ? ? ? ? ? ? ?skartveiti 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?sooretama 1 1 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?tuberculifera 0 0 0 0 0 0 0 1 ? 0 1 1 0 0 0 0 0 0 0? ? ? ? ? ? ? ?ubatuba 1 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 ? ? ? ? ? ? ? ?venequatoriensis 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 1 ? ? ? ? ? ? ? ?virga 0 0 0 0 0 0 1 1 ? 0 1 0 0 0 0 0 0 1 0? ? ? ? ? ? ? ?zempoalensis 0 0 1 1 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ?zhengi 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 ? ? ? ? ? ? ? ?



Character no.: 8 8 8 8 8 8 8 8 9 9 9 9 9 9 9 9 9 9 1 1 1 1 1 1 1 1 12 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 0 0 0 0 0 0 0 0

0 1 2 3 4 5 6 7 8

Aagaardia 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0Apometriocnemus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?Botryocladius ? ? ? A 1 1 0 0 1 1 1 0 0 0 1 A 0 1 1 A 0 1 0 0 1 1 1Brillia 0 0 0 0 0 A 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0Bryophaenocladius 0 A 0 0 1 0 1 1 0 0 1 1 0 A 0 0 0 0 0 0 0 0 0 0 0 0 0Chaetocladius 0 0 0 1 1 A 0 0 0 0 0 1 0 0 1 0 0 0 1 0 A 0 0 0 0 0 0Corynoneura 0 1 0 0 ? 0 0 1 0 0 0 0 0 0 A 0 0 0 1 1 1 0 0 1 1 1 1Cricotopus 0 1 0 A A 0 A A 1 0 0 A A 0 0 0 0 0 0 0 0 0 0 A 0 0 0Diplocladius 0 1 1 0 1 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0Doithrix ? ? ? ? ? 0 1 1 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0Eukiefferiella 1 1 0 0 1 0 1 A 1 1 0 0 0 0 1 0 0 1 A 1 A 0 0 A 1 1 0Georthocladius s.str 0 0 0 0 1 0 1 1 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 Gymnometriocnemus0 1 0 0 1 0 A 1 0 0 A 1 0 A A 0 0 0 0 0 0 0 0 0 0 0 0Hanocladius ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?Heleniella 0 1 0 0 1 0 A 0 0 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 0Heterotanytarsus 0 1 0 0 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0Heterotrissocladius 0 1 0 1 1 0 0 0 0 0 1 0 A 0 0 0 1 0 0 0 1 0 0 0 0 0 0Hydrobaenus 0 1 0 A 1 1 0 0 0 0 0 A 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0Krenosmittia 0 1 0 1 1 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0Limnophyes 0 1 A 1 1 0 0 1 0 0 0 A 0 A 1 0 0 1 0 0 1 1 1 0 0 0 0Lopescladius ? ? ? ? ? 0 1 1 0 0 A 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0Metriocnemus 1 1 0 1 1 0 A 1 0 0 1 A 0 1 1 0 0 0 A 0 A 0 0 0 0 0 0Orthocladius 0 A 0 A A A A 0 1 0 A A 0 A A 1 0 0 0 0 0 0 A 0 0 0 0Parachaetocladius 0 0 0 0 1 0 1 1 0 0 0 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0Parakiefferiella 0 1 0 1 A 0 0 A 1 0 0 A 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0Paralimnophyes 0 1 A 1 1 0 0 1 0 0 0 ? 0 1 1 0 0 1 0 0 1 1 0 0 0 0 1Parametriocnemus 0 1 0 1 1 A 1 0 0 0 A A 1 0 1 0 1 0 1 0 1 1 0 0 0 0 0Parasmittia 0 ? 1 0 1 ? 1 1 0 0 0 ? 0 1 1 0 0 0 0 0 ? ? 0 0 ? 1 1Propsilocerus 0 0 0 0 0 0 A 0 0 0 A 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0Psectrocladius 0 1 0 0 A 1 A 0 0 0 A A 0 A 1 1 0 1 A 0 0 0 A 0 0 0 0Pseudorthocladius 0 0 0 0 1 0 1 1 0 0 0 A 0 0 1 0 1 0 1 0 1 1 0 0 0 0 0Pseudosmittia 0 A 0 A 1 A A 1 0 0 0 A 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0Rheocricotopus 1 0 0 0 1 0 A 0 0 0 0 0 0 A 1 1 0 0 A 0 0 0 1 0 0 0 0Smittia 0 ? 1 0 1 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0Stictocladius 0 0 0 0 A 0 1 A 0 0 A 0 0 A 1 0 1 0 1 0 A 1 1 0 0 0 0Tvetenia 0 1 0 0 1 0 0 0 0 1 A 0 1 0 1 0 1 0 1 0 1 0 1 1 1 A 1Unniella 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 0 0Zalutschia 0 A A A 1 0 0 0 0 0 1 A 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0aberrans ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?antecalvus 0 0 0 1 1 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0arcuatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?berui ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?biota ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?calakmulensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?croizati ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?dentispina 0 1 1 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?folius ? ? ? ? ? 0 1 1 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0herradurus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?mateusi 0 ? 0 ? ? 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0musci 0 0 0 1 1 0 1 1 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0nerius ? ? ? ? ? 0 0 1 0 0 0 ? 0 ? 1 0 0 1 0 0 1 1 1 0 0 0 0pectinata ? ? ? ? ? 0 0 1 0 0 0 ? 0 ? 1 0 0 1 0 0 1 1 1 0 0 0 0pittieri ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?pluspilalus ? ? ? ? ? 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0scalpellatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?skartveiti ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?sooretama ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?tuberculifera ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?ubatuba ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?venequatoriensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?virga ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?zempoalensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?zhengi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?


594ZOOTAXA

TABLE 2 (continued)Character no.: 1 1 1 1 1 1 1 1 11 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

0 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3

9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5

Aagaardia 1 A 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 1 0 1 0 0 0Apometriocnemus ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?Botryocladius 1 1 1 1 1 0 0 0 0 1 0 0 0 1 0 0 0 0 C A 1 1 0 A 1 1 0Brillia 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 2 0 1 1 0 0 0 0Bryophaenocladius 1 1 1 1 A 0 1 A 0 0 0 1 0 1 0 0 0 2 A 1 0 1 0 0 A 1 0Chaetocladius 1 1 1 1 1 0 1 0 0 0 0 0 0 1 0 1 0 0 A 1 0 1 0 0 A 0 0Corynoneura 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 2 0 2 0 1 0 1 0Cricotopus 0 0 0 0 1 1 1 0 0 0 0 0 0 1 0 0 0 B A C 0 A 0 A 0 1 0Diplocladius 0 0 0 0 1 1 0 0 0 1 0 0 0 1 0 0 0 0 1 1 0 1 0 0 0 0 0Doithrix 1 1 1 1 0 0 1 0 0 0 1 0 1 1 0 0 1 2 1 1 0 1 0 1 1 0 0Eukiefferiella A 1 1 1 0 0 A 0 0 0 0 0 0 1 0 0 0 E A C 0 1 0 0 0 1 0Georthocladius s.str 1 1 1 1 A 0 1 0 0 0 0 0 0 1 0 A 1 A 1 1 0 1 0 1 1 0 0 Gymnometriocnemus1 1 1 1 A 0 1 0 0 0 0 1 0 1 0 0 0 C A 1 0 1 0 0 A 1 0Hanocladius ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?Heleniella 1 1 1 0 1 0 1 0 0 1 0 0 1 1 0 0 0 0 1 1 0 1 1 0 1 0 0Heterotanytarsus 0 A 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2 0 2 0 0 1 0 0Heterotrissocladius 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 2 0 1 C 0 1 A 0 0Hydrobaenus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 A 0 0 1 0 0 0 0 0Krenosmittia 1 1 1 1 0 0 1 0 0 0 1 0 1 1 0 0 0 0 0 2 0 1 0 1 0 A 0Limnophyes 1 1 1 1 1 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 1 0 1 A 0 0Lopescladius 1 1 1 1 0 0 1 1 0 1 0 0 0 1 0 0 0 0 0 1 1 2 1 1 1 1 0Metriocnemus 1 1 A A A 0 1 0 0 0 0 0 0 1 0 0 0 B A 1 0 1 0 A A A 0Orthocladius A A A A A A 1 0 0 0 A 0 A 1 0 0 0 E A C 0 1 0 0 0 1 0Parachaetocladius 1 1 1 1 1 0 1 0 0 0 0 0 0 1 0 0 1 0 1 1 0 1 0 0 1 0 0Parakiefferiella 1 0 0 A 1 0 1 0 0 0 1 0 1 1 0 0 0 0 A 1 1 1 0 1 0 1 0Paralimnophyes 1 1 1 1 1 0 1 1 0 0 0 0 0 1 0 0 0 0 1 A 0 1 0 1 A 0 0Parametriocnemus 1 0 0 0 0 0 1 0 0 0 0 0 1 A 0 0 0 0 B 1 0 1 0 0 0 0 0Parasmittia ? ? ? ? 1 ? ? ? ? 0 0 0 0 1 0 0 0 2 ? 2 0 0 0 0 1 0 1Propsilocerus 0 0 0 A A 0 0 0 0 0 0 0 0 0 0 0 0 0 1 C 0 1 0 1 0 0 0Psectrocladius 0 A 0 A 1 A 0 0 0 0 A 0 0 0 0 0 0 E A 1 0 1 0 1 0 0 1Pseudorthocladius 1 1 1 1 0 0 1 0 0 0 0 0 1 1 0 0 0 C C 1 0 1 0 A A 0 0Pseudosmittia 1 1 1 1 A A 1 1 0 0 0 0 0 1 0 0 0 B 1 2 0 0 0 1 1 1 0Rheocricotopus 0 A 0 0 1 A 0 0 0 0 0 0 0 A 0 0 0 0 1 1 0 1 0 0 0 A 0Smittia 1 1 1 1 0 0 1 1 0 0 0 0 0 1 0 0 0 2 C 2 0 1 0 0 0 0 0Stictocladius 1 1 A A A 0 1 1 0 1 0 0 A 1 0 0 1 A A 1 1 C 1 1 1 A 0Tvetenia 1 1 1 1 0 0 1 0 0 0 0 0 1 1 1 0 0 0 1 1 0 1 0 0 A 0 0Unniella 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 1 0 0 0Zalutschia 0 1 A A 0 0 0 0 1 0 0 0 0 0 0 0 0 0 A 0 0 1 0 0 A 0 0aberrans ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?antecalvus 1 1 1 1 0 0 1 0 0 0 0 1 0 1 0 1 0 0 1 ? ? ? ? ? ? ? ?arcuatus ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?berui ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?biota ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?calakmulensis ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?croizati ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?dentispina ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?folius 1 1 1 1 0 0 1 0 0 0 0 0 0 1 0 1 0 0 2 1 0 0 0 1 0 0 1herradurus ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?mateusi 1 1 1 1 0 0 1 0 0 0 0 0 0 1 0 0 0 2 1 ? ? ? ? ? ? ? ?musci 1 1 1 1 0 0 1 0 0 0 0 0 0 1 0 1 0 0 1 1 0 0 0 1 1 0 1nerius 1 1 1 1 1 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 1 0 1 1 0 0pectinata 1 1 1 1 1 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 1 0 1 0 0 0pittieri ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?pluspilalus 1 1 1 1 0 0 1 0 0 0 0 0 0 1 0 1 0 A 1 1 0 0 0 1 1 0 1scalpellatus ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?skartveiti ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?sooretama ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?tuberculifera ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?ubatuba ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?venequatoriensis ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?virga ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?zempoalensis ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?zhengi ? ? ? ? ? ? ? ? ?? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?



Character no.: 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 13 3 3 3 4 4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5

6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9

Aagaardia 0 0 1 0 0 1 1 0 0 0 1 1 0 0 1 0 1 0 0 0 0 0 0 0Apometriocnemus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0Botryocladius 1 0 1 0 ? A 0 1 0 0 0 1 0 0 0 1 1 0 0 0 ? 0 0 0Brillia 0 0 0 0 1 1 1 1 0 0 1 1 1 1 0 0 1 0 0 0 0 0 1 0Bryophaenocladius 0 1 2 0 0 1 1 0 0 2 A 2 A 1 0 0 1 1 1 2 0 2 1 1Chaetocladius 0 0 A 0 A 1 0 A 0 1 A C 0 A 0 0 1 0 0 0 0 0 A 0Corynoneura 0 1 2 0 1 1 0 1 0 E 1 1 0 1 0 0 1 ? 0 1 0 0 0 0Cricotopus A A 2 0 A A 1 0 A 0 0 1 0 1 0 0 1 0 0 0 0 0 A 0Diplocladius 0 0 0 0 1 1 1 1 0 0 1 1 0 0 0 1 0 0 0 0 0 0 1 0Doithrix 0 0 1 0 0 1 1 0 0 1 1 2 0 1 0 0 1 0 1 0 0 1 2 0Eukiefferiella 0 1 2 0 0 0 1 A 1 0 A C 0 1 0 0 1 A 0 A A 0 1 0Georthocladius s.str 0 0 1 0 ? 0 1 0 0 1 0 2 1 1 0 0 0 1 1 C 0 1 2 0 Gymnometriocnemus 0 1 2 0 0 1 1 0 0 2 1 2 A 1 0 0 1 1 1 2 0 2 1 1Hanocladius ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?Heleniella 0 0 2 0 0 1 1 0 0 1 1 1 1 1 0 0 0 0 0 0 1 0 1 0Heterotanytarsus 0 0 1 0 1 1 1 0 0 0 1 1 0 0 0 0 0 0 0 0 A 0 0 0Heterotrissocladius 0 0 1 0 0 1 1 A 0 1 A 1 0 0 0 0 1 0 0 0 0 0 0 0Hydrobaenus 0 0 1 0 0 1 1 0 0 0 1 1 0 0 0 0 1 0 0 0 A 0 0 0Krenosmittia 0 A 2 0 0 1 1 0 0 0 0 1 0 1 0 0 1 ? 0 3 0 0 0 0Limnophyes 0 0 2 0 0 1 0 0 0 0 1 1 0 1 0 0 1 A 0 0 1 0 A 0Lopescladius 0 0 2 0 0 0 0 1 0 0 0 1 0 1 0 0 1 ? 0 3 1 0 0 0Metriocnemus 0 A A 0 0 1 0 1 0 1 A 1 0 1 0 0 1 1 0 0 0 0 0 0Orthocladius 1 0 2 0 A A A 0 0 A 0 A 0 1 0 A A 0 0 0 0 0 A 0Parachaetocladius 0 0 1 0 0 0 1 0 0 1 A 2 1 1 0 0 1 1 0 3 0 1 2 0Parakiefferiella 0 1 C 0 0 A 1 0 0 0 A 1 0 0 0 0 0 ? 0 0 0 0 A 0Paralimnophyes 0 0 C 0 0 A 0 1 0 0 1 1 0 1 0 0 1 A 0 0 1 0 A 0Parametriocnemus 0 0 1 0 A 1 1 0 0 A 1 1 0 0 1 0 0 1 0 0 1 0 0 0Parasmittia 0 0 2 0 1 1 1 0 0 0 0 1 0 1 0 0 1 1 1 2 0 0 1 0Propsilocerus 0 0 0 0 1 1 1 1 0 0 1 A 0 0 0 0 0 0 0 0 A 0 0 0Psectrocladius 0 0 2 0 0 0 1 0 0 E A 1 0 0 0 1 0 1 0 A 0 0 0 0Pseudorthocladius 0 0 2 0 0 0 1 0 0 1 1 2 A 1 0 0 0 1 0 3 0 1 E 0Pseudosmittia 1 0 2 0 1 1 0 A 0 E 0 C 0 1 0 0 1 A 1 C 0 2 0 0Rheocricotopus A 0 2 0 0 0 1 0 0 0 A 1 0 0 0 1 0 0 0 0 A 0 0 0Smittia 0 0 2 0 ? 1 0 0 0 0 0 1 0 1 0 0 0 0 1 2 0 0 1 0Stictocladius 0 A 2 0 1 0 1 A 0 1 A C 1 1 0 0 0 A 0 3 1 1 1 0Tvetenia 0 0 2 0 A 0 1 0 1 0 A 1 0 1 0 0 1 1 0 0 1 0 0 0Unniella 0 0 2 0 ? 1 1 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0Zalutschia 0 0 1 0 1 1 1 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 E 0aberrans ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?antecalvus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?arcuatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?berui ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?biota ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?calakmulensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?croizati ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?dentispina ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?folius 0 0 2 1 1 1 1 1 0 0 0 1 1 1 0 0 1 1 1 2 0 0 0 1herradurus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?mateusi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?musci 0 0 2 1 1 1 1 1 0 0 0 1 1 1 0 0 1 1 1 2 0 0 0 1nerius 0 0 2 0 0 1 0 1 0 0 1 1 0 1 0 0 1 ? 0 0 0 0 ? ?pectinata 0 0 2 0 0 1 0 1 0 0 1 1 0 1 0 0 1 ? 0 0 0 0 ? ?pittieri ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?pluspilalus 0 0 2 1 1 1 1 1 0 0 0 1 1 1 0 0 1 1 1 2 0 0 0 ?scalpellatus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?skartveiti ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?sooretama ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?tuberculifera ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?ubatuba ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?venequatoriensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?virga ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?zempoalensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?zhengi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?


594ZOOTAXA


594ZOOTAXA FIGURE 1. Parsimony analysis of some genera of the Orthocladiinae with Diplocladius Kieffer,

Propsilocerus Kieffer and Brillia Kieffer combined as outgroup. A, strict consensus of shortesttrees. Bremer supports for each branch are given as numbers above the branches. B, strict consen-sus of shortest trees after reweighting according to the rescaled consistency index.

Eighty six trees were obtained each with 849 steps, consistency index (CI) of 0.23,retention index (RI) of 0.50, and rescaled consistency index (RC) of 0.12. The strict con-sensus tree is shown in Fig. 1A with the Bremer supports given above each branch.

Bootstrap values were 95 % for Antillocladius biota plus A. folius, 94 % for the Pseu-dorthocladius group, 88 % for Rheocricotopus plus Psectrocladius, 68 % for all except thebasal three genera, 81 % for A. arcuatus plus A. zempoalensis, 78 % for Lopescladius plusStictocladius, 73 % for Smittia plus Parasmittia, and less than 60% for other groupings.When the results were reweighted according to RC 3 trees each with 857 steps (when theweights were reset to one), CI of 0.33, RI of 0.65 and RC of 0.22 (Fig. 1B) were obtained.

Although the Bremer supports and Bootstrap values are low the results clearly showthat croizati, aberrans, berui and pittieri all should be placed in Compterosmittia. Theimmatures of Compterosmittia are practically not separable from those of LimnophyesEaton and Paralimnophyes Brundin. Immatures are, however, tentatively known for twospecies only. The imagines, on the other hand, differ in many respects. Probable synapo-morphies as compared to Limnophyes and Paralimnophyes are the finely punctated wingswith apical setae in most males and the known female, the bare squama, and the stronglyextended costa. All previously described species of the genus have a comb-like megasetawhile none of the new species have. The divisions in the reweighted cladogram reflectthis. However, it may be more appropriate for C. croizati and C. dentispina to change posi-tion based on the types of inferior volsella. C. nerius, C. pectinata, C. tuberculifera and C.croizati have a simple, projecting, rounded inferior volsella. In fact C. croizati and C.nerius are practically inseparable except for the comb-like megaseta of the latter. Theremaining species all have inferior volsella with a dorsal, bare, digitiform anterior projec-tion and a ventral, posterior, rounded lobe. However, any subdivisions within the genus arepremature considering the lacking knowledge of females and immatures. Also whetherHanocladius should remain as a separate genus as indicated by the not reweighted cla-dogram, or should be incorporated in Compterosmittia as suggested by the reweighted cla-dogram will have to await further material. It also is possible that C. virga may deserve aseparate genus primarily based on the unique virga.

Particularly the long virga with lateral lamellae and the pupa without thorn-like mac-rosetae, without posterior spines on the tergites and short male genital sac makes mateusimore similar to Gymnometriocnemus than to species of Antillocladius. We thus propose aseparate genus, Litocladius new genus, with the single included species Litocladiusmateusi new species. L. mateusi also differs from the remaining species by having asupraalar and a normally developed anal point with normal hair-like lateral setae.


594ZOOTAXAThe remaining species are maintained within the genus Antillocladius. The many-

toothed pecten epipharyngis of the known larvae constitute a uniquely derived synapomor-phy. Also the thorn-like macrosetae of the pupae can be regarded as a unique synapomor-phy. Although similar macrosetae occurs in Chaetocladius Kieffer they there are morespine-like. The approximately palmate seta anteriores (S I) also is a good synapomorphyalthough similar setae anteriores occur in Parasmittia Strenzke.

The species of Antillocladius fall into two main groups partly reflected in the strictconsensus tree of the reweighted characters; the first (eventually the nominal subgenus)includes antecalvus, skartveiti, arcuatus, zempolaensis and with some uncertainty herra-durus; the second group includes the remaining species

The first group has a unique, characteristic strongly set off inferior volsella with onecircular lobe and with or without another circular posterior projection of the lobe. A. her-radurus is scored as having a lobe-like apically bilobed inferior volsella in the data matrix,but could as well have been scored as having two well set off circular lobes, i.e. as theother species of the group. The pupa is known of A. antecalvus. It has a laterally expandedanal lobe as in some Gymnometriocnemus and Bryophaenocladius and thus differs fromthe other known pupae of the genus.

The second group can be divided into three more or less distinct subgroups; one sub-group containing A. calakmulensis and A. musci; another group including A. pluspilalus,known in all stages, the Chinese A. scalpellatus, A. venequatoriensis and A. zhengi; and athird subgroup containing the remaining species. A. calakmulensis has a virga consistingof several spines as opposed to the remaining species of the genus which either have novirga or a virga consisting of two spines. A. musci has a very similar hypopygium exceptfor the virga. The members of the second subgroup all have well-developed anal lobe,hairy wings, relatively long virga and “normal” inferior volsella, but no uniquely derivedsynapomorphy. Most members of the last subgroup also have an inferior volsella with amore or less triangular anterior dorsal part and a more posterior and rounded ventral part,i.e. the “normal” orthoclad type. Two species have a sharply triangular inferior volsellaand two species have a very long inferior lobe apically divided in one species. There alsoare large variations in the shape of the virga ranging from absence to 2 short or very longspines. The anal point varies from long and parallel-sided with short lateral setae to longand triangular with strong setae. One species lacks setae on squama. However, both pupaeand larvae are known for two of the species and the differences appear too slight to war-rant any divisions.

The characters are coded in ordered to indicate the phylogenetic position of the differ-ent species relative to known genera. The relationships among more basally placed generathus should not be taken for relevant. In fact preliminary parsimony analyses of a widerrange of genera have shown that the Corynoneura group of genera belong with Eukiefferi-ella Thienemann and related genera, and that Heleniella Gowin is closely related to Sticto-cladius Edwards and Lopescladius Oliveira.


594ZOOTAXA Key to males of some orthoclad genera

1. Acrostichals scalpellate ................................................................................................. 2— Acrostichals not scalpellate, but may be hooked ........................................... Not keyed

(According to Cranston et al. (1989) the acrostichals of Pseudosmittia Goetghebuermay be scalpellate. This appears to be incorrect.)

2. Anal point small, bare, anterior on anal tergite, consisting of hump-like projection or absent...................................................................................................................................... 3

— Anal point well developed, although sometimes short, posterior on tergite ................ 43. Epimeron, posterior anepisternum II always and preepisternum usually with setae;

wing membrane with coarse punctation.......................... Limnophyes Eaton, pro parte— Epimeron, posterior anepisternum II and preepisternum bare; wing membrane with

fine punctation...................................................... Botryocladius Cranston and Edward4. Squama bare, acrostichals in centre of scutum or some distance from front, costal

extension very long, megaseta can be dentate.............................................................. 5— Squama nearly always with setae, when bare acrostichals starting in front, virga usu-

ally present, costal extension short to long, megaseta never dentate ........................... 65. Virga absent or consisting of numerous spines, with more than 4 acrostichals, leg ratio

lower than 1.0, megaseta often dentate ................................... Compterosmittia Sæther— Virga broad and scale-like, with 4 acrostichals, leg ratio about 1.15..............................

...................................................................................... Hanocladius Wang and Sæther6. Anal point broad, rounded; virga consisting of numerous scattered spines....................

.............................................................................................................. Unniella Sæther— Anal point triangular to parallel-sided; virga absent, consisting of several short spines,

or of few very long spines with or without lateral lamellae......................................... 77. Two acrostichals in centre of scutum, R4+5 ending proximal to above M3+4; virga absent

........................................................................................ Synorthocladius Thienemann— Several acrostichals starting in front or in centre of scutum, R4+5 ending distal to M3+4;

virga present or absent.................................................................................................. 88. Wing with coarse punctation, without setae; gonocoxite with 2 lobes, the anteriormost

appearing to be superior volsella and dorsal part of inferior volsella combined, extend-ing from gonocoxite base to distal of midpoint; posteriormost lobe extending to closeto insertion of gonostylus; virga absent................................. Paralimnophyes Brundin

— Wing with fine punctation, often with apical setae; volsellae variable, but never asabove; virga usually present ......................................................................................... 9

9. Virga long, with lateral lamella; costal extension short ............. Litocladius new genus— Virga absent, short or long, but never with lateral lamellae; costal extension short to

very long...................................................................................... Antillocladius Sæther


594ZOOTAXAAntillocladius Sæther

Antillocladius Sæther, 1981: 4; Coffman et al. (1986: 160); Cranston et al. (1983: 157); Cranston etal. (1989: 176); Sæther (1982: 471; 1984: 1).

Type speciesAntillocladius antecalvus Sæther, 1981, by original designation.

Other included speciesAntillocladius arcuatus Sæther, 1982; A. biota new species; A. calakmulensis new spe-

cies; A. folius new species; A. herradurus new species; A. musci new species; A. pluspila-lus Sæther, 1982; A. scalpellatus Wang and Sæther, 1993; A. skartveiti Andersen andContreras-Ramos, 1999; A. sooretama new species; A. ubatuba new species; A. venequa-toriensis new species; A. zempoalensis new species and A. zhengi Wang and Sæther, 1993.

Diagnostic charactersThe imagines can be separated from other orthoclad genera by having scalpellate

acrostichals at least in the middle of scutum combined with a moderately to extremelylong anal point with lateral setae. (Although A. pluspilalus in the original description ismentioned as apparently having non-scalpellate acrostichals, more recently found speci-mens show that scalpellate acrostichals are present.) All known females and several maleshave setae apical on the wing membrane. The combination of absence of thoracic horn andpresence of thorn-like macrosetae will separate the pupae from other genera. The larva isdistinguished by a palmate S I; pecten epipharyngis divided into about 8–15 teeth; analsegment protruding over posterior parapods; and anal setae absent or perhaps reduced tosingle seta on minute tubercle.

Generic diagnosisImagoSmall to medium sized species, wing length 0.8–2.3 mm. Female antenna with 5 flagellomeres. Male antenna with 13 flagellomeres, groove

beginning at flagellomere 3, sensilla chaetica on flagellomeres 2, 3 and 13. Male antennalratio 0.50–1.70.

Head. Eye naked, with or without minute pubescence between ommatids, without dor-somedial extension. Temporal setae divided into weak inner verticals, stronger outer verti-cals, postorbitals absent or few. Third palpomere with about 3 sensilla clavata apically,longest 8–23 µm long.

Thorax. Antepronotum well developed, lobes meeting medially along short suture.Acrostichals, short, all scalpellate or anterior few simple, beginning close to anteprono-tum, some distance from antepronotum or in the middle of scutum; prealars uniserial;supraalar 0–1; scutellars uniserial.


594ZOOTAXA Wing. Anal lobe protruding to absent. Costal extension moderately to strongly devel-

oped (1/2 to 4 times the length of RM). R2+3 running and ending midway between R1 and

R4+5; R4+5 ending opposite or slightly distal to M3+4; FCu far distal to RM; Cu1 straight to

slightly sinuous; An ending proximal to FCu. Veins bare or with setae on R, R1, R4+5, M1+2,

M3+4, Cu, Cu1 and An. Membrane with apical setae in female; male without setae, with

few weak setae apically in cell r4+5, or with numerous setae apically in cells r4+5, m1+2 and

m3+4. Squama bare or with up to 15 setae.

Legs. Pseudospurs, sensilla chaetica and pulvilli lacking. Comb and tibial spurs nor-mal.

Abdomen. Tergites with irregularly arranged setae.Hypopygium. Anal point long, pointed, with strong lateral setae, with microtrichia

only at base or nearly to apex; apex never with setae. Phallapodeme and aedeagal lobe welldeveloped. Anterior margin of sternapodeme nearly straight to strongly arcuate, oral pro-jections barely indicated to strongly developed. Virga absent or consisting of 2–6 long,strong spines. Inferior volsella highly variable; simple and rounded or triangular; withanterior dorsal triangular or digitiform part and a more rounded ventral, posterior part;posterior lobe low to prominent; consisting of a posteriomedially directed, apically simpleor bifid lobe; or circular with or without additional rounded posterior extension and con-spicuously set off.

Gonostylus with or without heel; crista dorsalis absent to rounded and well developed. Megaseta normal.Female genitalia. Gonocoxapodemes curved and meeting anterior of vagina. Gono-

coxite relatively well developed, with a few long setae and additional short setae. TergiteIX divided or undivided with setae concentrated to lateral halves. Gonapophysis IX withlarge ventrolateral lobe covering apodeme lobe and apex of narrow, line-like dorsomesallobe. Labia relatively large, pointed. Cerci small. Seminal capsules small to medium-sized,circular, with triangular neck. Spermathecal ducts with bends or loops, with weak bulbsbefore separate openings.

PupaSmall to medium sized, total length 2.0 to 3.5 mm.Cephalothorax. Frontal setae absent. Frontal apotome smooth. Antennal sheath with or

without pearl row above pedicel. Ocular field with 2 reduced postorbitals. Thoracic hornabsent. Three weak precorneals situated in row or narrow triangle. Apparently two medianand 0–1 lateral antepronotals. Dorsocentrals very small, in two groups of two. Thorax andwing sheath smooth.

Abdomen. Tergite I without shagreen; tergites II–VIII covered with coarse, shagreenexcept along anterior margin; tergite IX with strong shagreen except along anterior andposterior margin. Sternites I and II bare; sternite III with median and lateral shagreen; ster-nites IV–VI with slightly more extensive median and lateral shagreen; sternites VII andVIII with extensive shagreen; sternite IX with anterolateral shagreen. Tergite II without


594ZOOTAXAposterior hooklets. Tergites II–VII or VIII with posterior row of weak spines. Conjunctives

bare. Pedes spurii A and B absent. Apophyses weak, nearly straight. Segment II–VIII with2 weak, hair-like L setae. Anal lobe with 3 thorn-like macrosetae, sometimes laterallyexpanded; male genital sac overreaching anal lobe, female genital sac not reaching apex oflobe.

LarvaSmall to medium sized, 3.0–3.3 mm long.Head. Antenna 5-segmented, less than half as long as mandible, last three segments

short. Blade as long as or longer than flagellum, striated. Lauterborn organs absent, stylelarge. S I nearly palmate with 5–6 teeth, one of them usually larger. Other S setae normal.No discernible labral lamellae. Pecten epipharyngis of 8–15 teeth (each of the normallyscales divided into 2–5 teeth). Chaetulae laterales simple, 3 strong and sclerotised, 4–5weak, pointed and unsclerotised. Chaetulae basales simple or bifid. Mandible with apicaltooth shorter than the combined widths of 4 inner teeth, fourth tooth sometimes joined tomola. Seta subdentalis triangular. Seta interna slender, of 4–5 nearly smooth to stronglyserrated branches. Mentum with large, irregular triangular, sometimes medially notchedmedian tooth higher than 5 pairs of lateral tooth, fourth pair smaller than fifth and appar-ently occasionally completely reduced. Ventromental plates small, visible only below 3outermost teeth, no beard. Maxilla without pecten galearis, lacinial chaeta reduced torounded lamellae.

Abdomen. Without distinct setae. Anterior parapods fused for most of their lengths,with minute groups of minute claws and some longer mostly smooth claws. Procercusapparently absent, but a vestigial tubercle with one short seta may represent procercus atleast in some species. Anal tubules shorter than half the length of posterior parapods(shape not clear in any available material). Posterior parapods well developed, with strongapical claws.

Key to the males of Antillocladius Sæther

1. Squama bare (Fig. 16D); Mexico.....................................A. zempoalensis new species— Squama with setae......................................................................................................... 22. Anal point nearly parallel-sided with numerous, weak, partly anterior directed setae;

inferior volsella either pointed triangular or consisting of a long apically simple orbifid lobe; wing bare ..................................................................................................... 3

— Anal point tapering with numerous, usually strong and posterolateral directed setae;inferior volsella either with a dorsal anterior triangular to digitiform part and a morerounded ventral lobe or circular with or without additional rounded posterior extensionand conspicuously set off; wing usually with apical setae........................................... 6

3. Inferior volsella pointed triangular, costal extension very long................................... 4— Inferior volsella a long posteromedially directed simple or apically bifid lobe, costal


594ZOOTAXA extension of moderate length ....................................................................................... 5

4. Inferior volsellae triangular, leaf-like (Fig. 5D); Brazil ............... A. folius new species— Inferior volsella small, sharply pointed (Fig. 3D); Brazil ............. A. biota new species5. Inferior volsella apically bifid (Fig. 13C); Brazil ................ A. sooretama new species— Inferior volsella simple (Fig. 14D); Brazil, Venezuela ............ A. ubatuba new species6. Virga at least nearly as long as half phallapodeme length; wing membrane with apical setae

....................................................................................................................................... 7— Virga absent or short; wing membrane with or without apical setae .......................... 117. Anal point with numerous weak setae; inferior volsella weak (Fig. 15D); Venezuela,

Ecuador....................................................................... A. venequatoriensis new species— Anal point with strong, stiff setae; inferior volsella well developed............................ 88. Wing with few setae apically in cell r4+5; AR 1.4–1.7................................................. 9

— Wing with more numerous apical setae present also in cells m1+2 and sometimes m3+4;

AR 1.0–1.6 ................................................................................................................. 109. Virga nearly as long as phallapodeme (Fig. 9E); Brazil.............. A. musci new species— Virga less than half as long as phallapodeme; China, Thailand......................................

........................................................................................... A. zhengi Wang and Sæther10. Virga about 2/3 as long as phallapodeme; squama with 5–6 setae; cell m3+4 with sev-

eral setae; Ecuador, Mexico, Nicaragua, USA............................ A. pluspilalus Sæther — Virga half as long as phallapodeme; squama with 8–13 setae; cell m3+4 bare; China, Russia

................................................................................... A. scalpellatus Wang and Sæther 11. Virga consisting of 2 or 6 very short spines; costal extension short .......................... 12— Virga absent; costal extension long ............................................................................ 1312. Virga consisting of 6 spines; inferior volsella with digitiform dorsal anterior lobe cov-

ering rounded posterior, ventral lobe (Fig. 4F); Mexico A. calakmulensis new species— Virga consisting of 2 spines; inferior volsella apically divided into 2 semicircular lobes

(Fig. 8F); Mexico ................................................................ A. herradurus new species 13. Inferior volsella a well set off circular lobe with additional posterior semicircular

extension; wing membrane bare; Brazil, Mexico, Venezuela, USAA. arcuatus Sæther— Inferior volsella a well set off circular lobe; wing membrane usually with setae...... 1414. Wing membrane with numerous apical setae, present also in cell m3+4; Ecuador..........

................................................................ A. skartveiti Andersen and Contreras-Ramos— Wing membrane with 0–3 setae apically in cell r4+5; Brazil, Saint Vincent, Venezuela

...................................................................................................... A. antecalvus Sæther

Key to known females of Antillocladius Sæther

1. Tergite IX distinctly divided (Fig. 10F) ...................................... A. musci new species— Tergite IX fused............................................................................ A. antecalvus Sæther


594ZOOTAXAKey to known pupae of Antillocladius Sæther

1. Antennal sheath with pearl row.................................................................................... 2— Antennal sheath lacking pearl row ................................................................................ 32. Total length about 2.9–3.2 mm, anal macrosetae longer than half the length by which

the male genital sac overreaches the anal lobe (Fig. 11F)........... A. musci new species— Total length about 2.4 mm, anal macrosetae shorter than half the length by which the

male genital sac overreaches the anal lobe (Fig. 6E) ................... A. folius new species3. Anal lobe expanded laterally (Fig. 2E) ........................................ A. antecalvus Sæther— Anal lobe not expanded laterally................................................. A. pluspilalus Sæther

Key to known larvae of Antillocladius Sæther

1. Antennal blade about as long as flagellum (Fig. 7A)................... A. folius new species— Antennal blade longer than flagellum .......................................................................... 22. Third antennal segment shorter than fourth and fifth; pecten epipharyngis of about 8

teeth; median tooth of mentum notched, fourth lateral tooth apparently sometimes absent..................................................................................................... A. pluspilalus Sæther

— Third antennal segment about as long as fourth and fifth; pecten epipharyngis of about12 teeth; median tooth of mentum not notched, fourth lateral tooth present (Fig. 12A, C)..................................................................................................... A. musci new species

Antillocladius antecalvus Sæther (Fig. 2A–E)

Antillocladius antecalvus Sæther, 1981: 4, Fig. 2 A–G.Antillocladius antecalvus Sæther; Spies and Reiss 1996: 75.

Material examined: Type material as in Sæther (1981). BRAZIL: São Paulo, Pedregulho,Furna São Pedro, 1 male with pupal exuviae, 7.vii.2000, S. Mateus (MZUSP); Santa Cata-rina State, São Bento do Sul, 26°19'25.6''S, 48°18'26.5''W, 1 male, 13–16.x.2001, Malaisetrap, M.V. Yamada (UFSCar). VENEZUELA: Aragua, Parque Nacional Henri Pittier, RanchoGrande, 10°21.047'N, 67°41.198'W, 21 males, 16–18.ix.1999, about 1000 m a.s.l., sweepnet & light trap, T. Andersen (2 MIZA, 19 ZMBN); Falcón, Rio Mitare near San Luis,11°07.930'N, 69°39,184'W, 18 males, 7.vi.2001, 589 m a.s.l., light trap, R.W. Holzenthal,R. Blahnik, H. Paprocki & C. Cressa (2 MIZA, 16 ZMBN).

Diagnostic characters: The male imago can be separated from other members of thegenus by having 0–3 apical setae on the wing membrane; squama with 1–3 setae; few scal-pellate to non-scalpellate acrostichals in the middle of scutum; virga absent; and inferior


594ZOOTAXA volsella large with anterior bare, strongly set off rounded projection and low posterior

lobe. The female differs from other known females by having tergite IX undivided, buttergite setae still in two groups. The pupa has thorn-like macrosetae and anal lobe laterallyexpanded.

FIGURE 2. Antillocladius antecalvus Sæther, 1981, pupa. A–tergites I–IX; B–sternites I–IX; C–

frontal apotome; D–thorax, anterior part, lateral view; E–detail of anal lobe.

Remarks: The species was previously known only f

Documents

Zootaxa, Diptera, Chironomidae · 1981 and Litocladius new genus (Chironomidae, Orthocladiinae) 1HUMBERTO F. MENDES, 2TROND ANDERSEN & 2OLE A. SÆTHER 1Departamento de Biologia-FFCLRP,