Mendes, H. F, Andersen, T. & Sæther O. A

ZOOTAXA

594

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

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

Magnolia Press Auckland, New Zealand 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 ISBN 1-877354-48-1 (Paperback) ISBN 1-877354-49-X (Online edition)

FIRST PUBLISHED IN 2004 BY Magnolia Press P.O. Box 41383 Auckland 1030 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) Zootaxa 594: 1–82 (2004) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 594 Copyright © 2004 Magnolia Press ISSN 1175-5334 (online edition)

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

1HUMBERTO F. MENDES, 2TROND ANDERSEN & 2OLE A. SÆTHER 1Departamento 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 ...... 4 Introduction ...... 5 Material and Methods ...... 6 List of abbreviations ...... 6 Phylogeny ...... 6 Key to males of some orthoclad genera ...... 22 Antillocladius Sæther ...... 23 Key to the males of Antillocladius Sæther ...... 25 Key to known females of Antillocladius Sæther ...... 26 Key to known pupae of Antillocladius Sæther ...... 27 Key to known larvae of Antillocladius Sæther ...... 27 Antillocladius antecalvus Sæther ...... 27 Antillocladius arcuatus Sæther ...... 29 Antillocladius biota new species ...... 30 Antillocladius calakmulensis new species ...... 32 Antillocladius folius new species ...... 34 Antillocladius herradurus new species...... 39 Antillocladius musci new species ...... 41 Antillocladius pluspilalus Sæther ...... 48 Antillocladius scalpellatus Wang and Sæther ...... 49 Antillocladius skartveiti Andersen and Contreras-Ramos ...... 49 Antillocladius sooretama new species ...... 50 Antillocladius ubatuba new species ...... 52 Antillocladius venequatoriensis new species ...... 54 Antillocladius zempoalensis new species ...... 57 Antillocladius zhengi Wang and Sæther ...... 59 Compterosmittia Sæther ...... 59 Key to the males of Compterosmittia Sæther ...... 61

Accepted by A. Whittington: 10 Jul. 2004; published: 6 Aug. 2004 3 ZOOTAXA Compterosmittia aberrans new species ...... 62 594 Compterosmittia berui new species ...... 64 Compterosmittia claggi (Tokunaga)...... 66 Compterosmittia croizati new species ...... 67 Compterosmittia dentispina Sæther ...... 69 Compterosmittia nerius (Curran) ...... 69 Compterosmittia pectinata (Freeman) ...... 69 Compterosmittia pittieri new species ...... 70 Compterosmittia tuberculifera Tokunaga ...... 72 Compterosmittia virga Wang ...... 72 Litocladius new genus ...... 72 Litocladius mateusi new species ...... 74 Distribution ...... 79 Biology of the Antillocladius, Compterosmittia and Litocladius species ...... 79 Acknowledgements ...... 80 References ...... 81

Abstract

A parsimony analysis of recently collected species sharing several features such as scalpellate acrostichals, often setae apically on the wing membrane and often strongly extended costa, together with 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 Antillocladius are described and figured as male imagines: A. calakmulensis, A. herradurus and A. zempoalensis from 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, the pupae of A. antecalvus Sæther, A. folius, and A. musci, and the larvae of A. folius and A. musci are also 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 and larvae 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 the British 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 all three genera are terrestrial or associated with phytotelmata, and notes on their biology and larval habitats are included.

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

4 © 2004 Magnolia Press MENDES ET AL. Introduction ZOOTAXA 594 The Neotropical chironomid fauna contains numerous species that are difficult to place amongst the previously described, mostly Holarctic genera. A number of recently collected species shares several features such as scalpellate acrostichals, often setae apically on the wing membrane, often strongly extended costa etc. Most of these features are found in the genus Antillocladius Sæther. However, also Paralimnophyes Brundin, and Compterosmittia Sæther, combine scalpellate acrostichals with well-developed anal point and 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, for instance, is quite similar to the pupa of Antillocladius pluspilalus Sæther, but has lateral expansions of the anal lobe. Another lacks the characteristic thorn-like macrosetae on the anal 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 analysis entering all known species of Antillocladius and Compterosmittia separately together with these 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 by Sæther (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 and emended the genus. Wang and Sæther (1993) described two additional new species from Palaearctic and Oriental China. Recently Andersen and Contreras-Ramos (1999) described a 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. zhengi Wang and Sæther, 1993 from the Hainan Province in China and A. skartveiti Andersen and Contreras-Ramos, 1999 from Ecuador. Spies and Reiss (1996) recorded A. pluspilalus from Nicaragua. The female of A. antecalvus was described by Sæther (1981), while the immatures of A. pluspilalus were described by Sæther (1984), indicating that the genus is related to Bryophaenocladius Thienemann and Gymnometriocnemus Goethgebuer. The genus Compterosmittia originally was described from the British West Indies based 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 synonym of C. nerius (Curran, 1930). Outside the New World the genus has been found in Australia [C. pectinata (Freeman, 1961)], Oceania [C. claggi (Tokunaga, 1964), C. tuberculifera (Tokunaga, 1964)] and the Oriental Region (Cranston & Martin 1989; Cranston & Kitching 1995; Wang 1998). Compterosmittia virga Wang, 1998 is a doubtful member of the genus since it has a virga consisting of numerous spines.

ORTHOCLADIINAE © 2004 Magnolia Press 5 ZOOTAXA Material and Methods 594 The material examined was mounted on slides following the procedure outlined by Sæther (1969). The general terminology follows Sæther (1980). Measurements are given as ranges, followed by the mean when 4 or more specimens are measured, followed by the number of specimens measured in parenthesis. The holotypes of the new Brazilian species are housed in the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP), except for A. sooretama, which is 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 Instituto de Zoología Agrícola “Francisco Fernández Yépes” (MIZA), Faculdad de Agronomia, Universidad Central de Venezuela, Maracay, Venezuela. Other holotypes are in the Museum of Zoology, Bergen, Norway (ZMBN). Paratypes are deposited in the museum as cited 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 158 characters in 38 genera and 25 species, each species entered separately for Antillocladius and Compterosmittia as well as the new preliminary unassigned species (Tables 1, 2). The immatures of Compterosmittia have not been assigned to species with certainty. It is here assumed that the Australian immatures described by Cranston and Kitching (1995) belong to the only described Australian species, C. pectinata (Freeman); and that the somewhat different immatures from Hong Kong belong to C. claggi (Tokunaga) which again appear to be a junior synonym of C. nerius (Curran). Diplocladius Kieffer, Propsilocerus Kieffer and Brillia Kieffer combined were used as outgroup. Data was analysed under parsimony with 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.

6 © 2004 Magnolia Press MENDES ET AL. TABLE 1 Characters and states used in the phylogenetic analyses. For some characters polymor- ZOOTAXA phies are scored as apomorphies as they otherwise would be uninformative or appear constant. 594

Imagines 1. 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 palpomeres or palpomeres strongly reduced in length. 7. Sensilla clavata of palpomeres: (0) palpomere 3 at most with few sensilla in one group, palpomere 4 without sensilla; (1) at least female with sensilla in more than one group on palpomere 3 and palpomere 4 usually also with at least sensilla clavata, or, in Prodiamesa and Chaetocladius, 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 sometimes female like (more than 7 times as long as wide). 11. Antepronotal lobes: (0) broad, collar like, at most slightly narrowed medially; (1) distinctly narrowed 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

594 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 on M1+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 posterior on tergite. 45. Anal point: (0) not with spatulate microtrichiae- free apex; (1) often with small or large spatulate, microtrichiae-free apex. 46. Anal point: (0) not very broad and rounded to bluntly triangular; (1) conspicuously broad and rounded to bluntly triangular. 47. Anal point: (0) not long, when moderately sized triangular; (1) moderately sized, triangular with 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 and plate-like. 51. Inferior volsella: (0) at least some species with anterior dorsal, often triangular or digitiform part and a usually more rounded, small to large ventral, posterior part, sometimes separated from dorsal part, occasionally more than 2 parts simple, well developed to reduced; (1) different...... continued on the next page

8 © 2004 Magnolia Press MENDES ET AL. TABLE 1 (continued) ZOOTAXA 52. Inferior volsella: (0) without microtrichia-free digitiform projection; (1) with anterior microt- 594 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-shaped or digitiform, sometimes hooked, occasionally spiniform. 54. Inferior volsella: (0) not adpressed to gonocoxite; (1) simple, elongate, adpressed to gonocoxite, sometimes pediform with free end. 55. Inferior volsella: (0) not consisting of long posteromedially directed lobe; (1) consisting of a posteromedially 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 additional 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 triangular. 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 consisting 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 lamellae. 73. Virga: (0) virga not consisting of long spines without lamellae; (1) in some species consisting of 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 gonapophysis; (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 two setigerous protrusions.

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ORTHOCLADIINAE © 2004 Magnolia Press 9 ZOOTAXA TABLE 1 (continued) 594 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 dorsomesal lobe 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 with microtrichiae; (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 elongate 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 species with 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 and species 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 2–3 dorsocentrals 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) some with spines or tubercles. 101. Sternites or sternal conjunctives: (0) without caudal hooklets; (1) sometimes with...... continued on the next page

10 © 2004 Magnolia Press MENDES ET AL. TABLE 1 (continued) ZOOTAXA 102. Male sternite VIII: (0) without posterior spine or tubercle row(s); (1) with. 594 103. 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 double 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 hooklets 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 tergite VIII with in most species. 118. Anal lobe: (0) not extended into projections; (1) extended posteriorly into cylindrical projections with macrosetae at apex. 119. Anal lobe: (0) not with apical projections or extensions; (1) with apical spurs or extended distal 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.

Larva 128. 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

ORTHOCLADIINAE © 2004 Magnolia Press 11 ZOOTAXA TABLE 1 (continued) 594 131. 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 indistinct. 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 conspicuously long. 156. Supraanal setae: (0) weak or absent, shorter than 1/2 length of anal setae when these well developed or shorter than 2/3 when anal setae short; (1) well developed, longer than 1/2 length 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 digitiform. 158. Anal tubules: (0) at least 1/2 length posterior parapods; (1) shorter than 1/2 length posterior parapods or these absent; (2) conspicuously long and narrow. 159. Anal segment: (0) not projecting posterior over posterior parapods or bent at right angles to axis of rest of body; (1) projecting posterior over posterior parapods or bent at right angles to axis of rest of body.

12 © 2004 Magnolia Press MENDES ET AL. TABLE 2 Character states for characters 1–159 in some genera of Orthocladiinae. Polymorphies: A = 0&1, B = ZOOTAXA 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. 594 Character no.: 123456789111111111122222222 012345678901234567 Aagaardia 1000000100000AA000011A00011 Apometriocnemus 000?00000000000110101000411 Botryocladius A0000001000020012200100000A Brillia 001100A000010A11?0101100410 Bryophaenocladius 00A0000A00000AAA00A110000AA Chaetocladius A1A0A0AA01002AA100AA100001A Corynoneura A100000010100001?0011000002 Cricotopus A1AA100AA000AAA100AAAA110AA Diplocladius 000010011000000100011000000 Doithrix 001?001100000100001111000A0 Eukiefferiella A101A00AA0000001E0011000000 Georthocladius s.str0010001100000AA00011AA000A0 Gymnometriocnemus 0010000100100?A110111000N11 Hanocladius 110?00000000000122011000012 Heleniella 0000100100000111?011011100A Heterotanytarsus 000000001000000120010000A01 Heterotrissocladius 0001000000000AAA00A11A00KA0 Hydrobaenus 0100010A00000A0110A1AA000AA Krenosmittia 1100000110100001?0011000002 Limnophyes A1000001A0012AA112100010011 Lopescladius A10?100111101001?0011000001 Metriocnemus A1AA000A010001100010A100NAA Orthocladius 000000010000E?A100AAAA0100A Parachaetocladius 0010001100000A01?011A0000A0 Parakiefferiella A10?000110100001?0011000001 Paralimnophyes 001?00?10000011022001000011 Parametriocnemus A10000A000000AA000A11A00NAB Parasmittia 001?00?0000001A100111100301 Propsilocerus 010001A1A0000AA1?0A1AA00010 Psectrocladius 000A000100000AAA00A11A00000 Pseudorthocladius 0010001A00000AA000111100L01 Pseudosmittia A0AA00A1A0A00 0A1 20AA1 0000 0B Rheocricotopus A0001001A000CA01A0A110000AA Smittia 0011100100000001?01A100000A Stictocladius A001100111A0CAA1?0A11A1100A Tvetenia 01000001A0000001?001100000A Unniella 000000010000000122011000001 Zalutschia 0000000A0000000110A1A00000B aberrans 010000010000000122011000102 antecalvus 000000010000000121011000101 arcuatus 000000010000000101011000000 berui 000000010000000122011000A02 biota 000000010000000102011000002 calakmulensis 000000010000000112011000100 croizati 000000010000000112011000002 dentispina 1?0000010000000112001000102 folius 000000010000000101011000002 herradurus 000000010000000101011000101 mateusi 000000010000000101001000001 musci 000000010000000112011000100 nerius 0?0000010000000112001000A01 pectinata 0000000100000001220?1000101 pittieri 000000010000000122001000202 pluspilalus 000000010000000101011000300 scalpellatus 000000010000000101011000200 skartveiti 000000010000000112011000302 sooretama 000000010000000102011000001 tuberculifera 1000000100000001?20?1000001 ubatuba 000000010000000102011000000 venequatoriensis 000000010000000102011000401 virga 000000010000000112011000002 zempoalensis 000000010000000101011000002 zhengi 000000010000000101011000100

ORTHOCLADIINAE © 2004 Magnolia Press 13 ZOOTAXA TABLE 2 (continued) 594 Character no.: 223333333333444444444455555 890123456789012345678901234 Aagaardia C0A101112110010130000100000 Apometriocnemus 202100201010011120000100000 Botryocladius 000111101010011110000000000 Brillia 000100201100010000000001001 Bryophaenocladius B001001A00AA0A11F0A00A00111 Chaetocladius A0A00AAA1010100A30AHE100000 Corynoneura 01A2130101C0A11100000010000 Cricotopus A0011BC0B0C0010AJ0000A00010 Diplocladius 000001111100011130000101000 Doithrix 0011101011110110200G1101010 Eukiefferiella 0A01ADA010C00AA100000100010 Georthocladius s.str00A10A1011A0001020001100010 Gymnometriocnemus G0A10021211001112000A100000 Hanocladius 302C13010000011130010100000 Heleniella 1021111111A001A1F0000100010 Heterotanytarsus A00C00C0101101A130000A00010 Heterotrissocladius 000100C0011001A1310IC100000 Hydrobaenus A00A0A10C11000013000A100000 Krenosmittia 31C1110121100111G0000100010 Limnophyes C0110CAA211001A120000100010 Lopescladius 01010301?11001100000010?010 Metriocnemus CA01002001A0A011300BA100000 Orthocladius A0010A1001000A0130010A00000 Parachaetocladius 0011001011A00A1020001100010 Parakiefferiella C02111112110A11130100A00110 Paralimnophyes C0010110?10??10130110001001 Parametriocnemus C01100C011100111310B0100010 Parasmittia 20011021?10??11130110100000 Propsilocerus 0000011011010A01F00A0A00100 Psectrocladius 00010A1001B0A000F00BA100010 Pseudorthocladius A011A0C011100110E0001100010 Pseudosmittia 0AB21DA12110AAA1A0000000000 Rheocricotopus 0001011A0110A100300C0010010 Smittia 201101111100011130120100010 Stictocladius AAA10HAAB0EA0A11F0000101001 Tvetenia 0A011110111000A1300C0100010 Unniella 002101101110010130100000000 Zalutschia A001011001100001300A0100000 aberrans 20011?011110011130011100100 antecalvus 100C11A01110011130021101000 arcuatus 101C1?001110011130021101000 berui 30021?011110011130010100100 biota 200C1?A01110011130032101010 calakmulensis 00011?001110011130021100000 croizati 301C11111110011130010100010 dentispina 302211111110011130010100100 folius 200211A01110011130032101010 herradurus 00011?001110011130021101000 mateusi 100211001110011130020100000 musci 100C11A01110011130021100000 nerius 302211111110011130010100010 pectinata 302211111110011130011100010 pittieri 300210111110011130010100100 pluspilalus 100110201110011130021100000 scalpellatus 000110201110011130021100000 skartveiti 200C1A201110011130021101000 sooretama 000C1?001110011130032101000 tuberculifera 300211111110011130010100010 ubatuba 10011?A01110011130032101000 venequatoriensis 100C11201110011130022100000 virga G02C11111110011130021100010 zempoalensis 10011?011110011130021101000 zhengi 10011?001110011130021100000

14 © 2004 Magnolia Press MENDES ET AL. TABLE 2 (continued) ZOOTAXA Character no.: 555556666666666777777777788 594 567890123456789012345678901 Aagaardia 00000 1 0 1E 000000000012102111 Apometriocnemus 0000000100010000000???????? Botryocladius 000000111000011000000011110 Brillia 000010012101000000010004111 Bryophaenocladius 10000 0 0C1 000000000010004111 Chaetocladius 0000AA0 1BA0100000000C1 1 4 110 Corynoneura 00000 0 2 2E 001000000001110211 Cricotopus 0000A0 0 1B00110000000 2BB 3 11A Diplocladius 100010012100000000010004100 Doithrix 1000000220011000000???????? Eukiefferiella 00000 0 0AB001100000000000A11 Georthocladius s.str00000 1A0C001000000010004111 Gymnometriocnemus 00000 0ACB001000110010114111 Hanocladius 0000000120010100000???????? Heleniella 00000 0 0A2 001000110002004110 Heterotanytarsus 00000AA1 2 001000000012010010 Heterotrissocladius 00000A01000000000001 2A0 2 110 Hydrobaenus 00000A01200000000001C1 1M111 Krenosmittia 000000012001000110002111110 Limnophyes 00000001000101100001B1 1 4 110 Lopescladius 00000 0AC2 1010000000???????? Metriocnemus 00000 0 0AA0000000000AAAA 4 11A Orthocladius 00000 0 0 1A0000000000A0 0A 3 11A Parachaetocladius 000001012001000000000004111 Parakiefferiella 00000 0 0 1C000000110002111110 Paralimnophyes 00000010000100000001B1 1 4 110 Parametriocnemus 00000001000100000001211M110 Parasmittia 000000010001000000000103210 Propsilocerus 1100101010010000000AC0 0 4 200 Psectrocladius 00000 0 0AE 0010100000A2A1MC1A Pseudorthocladius 00000 0 0 1C00010011111 0A1 4 111 Pseudosmittia 00000 0ACC00101100000E 1A 4 210 Rheocricotopus 10000 0 0AA00100000000E 1 1 4 111 Smittia 000000100000101000000103210 Stictocladius 00001 0AC2 10100000001 2CC 4 21A Tvetenia 00000 0 0 1E 000000000012114110 Unniella 00000001000110000001 0 1 ? 4 110 Zalutschia 00000110200000000001B1 1 4 110 aberrans 0000000100010000000???????? antecalvus 001000010001000000001114110 arcuatus 0011000100010000000???????? berui 0000000100010000000???????? biota 0000000100010000011???????? calakmulensis 0000001100000000000???????? croizati 0000000100010000000???????? dentispina 000000010011000000012114111 folius 0000001100010000011???????? herradurus 1100000100010010000???????? mateusi 000000110001000110002114110 musci 000000010001000001102114110 nerius 0000000100110000000???????? pectinata 00000 0 0 1 ? 0110000000???????? pittieri 0000001100010000000???????? pluspilalus 0000000100010000011???????? scalpellatus 0000000100010000010???????? skartveiti 0010000100010000000???????? sooretama 1100000100010000000???????? tuberculifera 00000 0 0 1 ? 0110000000???????? ubatuba 1000000100010000010???????? venequatoriensis 0000000100010000011???????? virga 00000 0 1 1 ? 0100000010???????? zempoalensis 0011000100010000000???????? zhengi 0000000100010000010????????

ORTHOCLADIINAE © 2004 Magnolia Press 15 ZOOTAXA TABLE 2 (continued) Character no.: 888888889999999999111111111 594 234567890123456789000000000 012345678 Aagaardia 000010000000000000000000000 Apometriocnemus ??????????????????????????? Botryocladius ???A11001110001A011A0100111 Brillia 00000A000011010000000000000 Bryophaenocladius 0A0 0 1 0 1 1 001 1 0 A0 0 000 0 0 00 0 00 0 Chaetocladius 00011A00000100100010A000000 Corynoneura 01 0 0 ? 0 0 1 000 0 0 0 A0 001 1 1 00 1 11 1 Cricotopus 01 0 AA0 AA100 AA0 0 0 000 0 0 00 A00 0 Diplocladius 011011000011000000000000000 Doithrix ?? ? ? ? 0 1 1 000 0 0 0 1 0 001 0 1 00 0 00 0 Eukiefferiella 1100101A1100001001A1A00A110 Georthocladius s.str000010110000001000101000000 Gymnometriocnemus 01 0 0 1 0 A1 00A1 0 AA0 000 0 0 00 0 00 0 Hanocladius ??????????????????????????? Heleniella 010010A00000001000101100000 Heterotanytarsus 010001000010001000000000000 Heterotrissocladius 010110000010A00010001000000 Hydrobaenus 010A1100000A000000000000000 Krenosmittia 010110000010001000101000000 Limnophyes 01 A1 1 0 0 1 000 A0 A1 0 010 0 1 11 0 00 0 Lopescladius ?? ? ? ? 0 1 1 00A0 0 0 1 0 001 0 1 00 0 00 0 Metriocnemus 110110A1001A011000A0A000000 Orthocladius 0A0 AAAA0 10AA0 AA1 000 0 0 0A0 00 0 Parachaetocladius 000010110000001000101000000 Parakiefferiella 01 0 1 A0 0 A100 A0 0 0 0 000 0 0 00 0 00 0 Paralimnophyes 01 A1 1 0 0 1 000 ? 0 1 1 0 010 0 1 10 0 00 1 Parametriocnemus 01 0 1 1 A1 0 00AA1 0 1 0 101 0 1 10 0 00 0 Parasmittia 0?101?11000?01100000??00?11 Propsilocerus 000000A000A1000000000000000 Psectrocladius 01 0 0 A1 A0 00AA0 A1 1 01A0 0 0A0 00 0 Pseudorthocladius 00001011000A001010101100000 Pseudosmittia 0A0 A1 AA1 000 A0 1 0 0 000 0 0 00 0 00 0 Rheocricotopus 10 0 0 1 0 A0 000 0 0 A1 1 00A0 0 01 0 00 0 Smittia 0?1010110001000000000000000 Stictocladius 00 0 0 A0 1 A00A0 0 A1 0 101 0 A11 0 00 0 Tvetenia 0100100001A01010101010111A1 Unniella 000000000000001000100010000 Zalutschia 0AAA1 0 0 0 001 A0 0 0 0 000 0 0 00 0 00 0 aberrans ??????????????????????????? antecalvus 000110110000001000001000000 arcuatus ??????????????????????????? berui ??????????????????????????? biota ??????????????????????????? calakmulensis ??????????????????????????? croizati ??????????????????????????? dentispina 01101? ? ? ??? ? ? ? ? ? ??? ? ? ?? ? ?? ? folius ?? ? ? ? 0 1 1 000 1 0 0 1 0 000 0 1 00 0 00 0 herradurus ??????????????????????????? mateusi 0? 0 ? ? 0 1 1 000 1 0 0 0 0 000 0 0 00 0 00 0 musci 000110110001001000001000000 nerius ?????001000?0?1001001110000 pectinata ?????001000?0?1001001110000 pittieri ??????????????????????????? pluspilalus ?? ? ? ? 0 1 1 000 0 0 0 1 0 000 0 1 00 0 00 0 scalpellatus ??????????????????????????? skartveiti ??????????????????????????? sooretama ??????????????????????????? tuberculifera ??????????????????????????? ubatuba ??????????????????????????? venequatoriensis ??????????????????????????? virga ??????????????????????????? zempoalensis ??????????????????????????? zhengi ???????????????????????????

16 © 2004 Magnolia Press MENDES ET AL. ZOOTAXA TABLE 2 (continued) Character no.: 111111111111111111111111111 594 011111111112222222222333333 901234567890123456789012345 Aagaardia 1A1110000000000000200101000 Apometriocnemus ??????????????????????????? Botryocladius 111110000100010000CA110A110 Brillia 000011000000000000120110000 Bryophaenocladius 1111A01A0001010002A10100A10 Chaetocladius 111110100000010100A10100A00 Corynoneura 111100000000001000020201010 Cricotopus 00001110000001000BAC0A0A010 Diplocladius 000011000100010000110100000 Doithrix 111100100010110012110101100 Eukiefferiella A11100A0000001000EAC0100010 Georthocladius s.str1111A0100000010A1A110101100 Gymnometriocnemus 1111A010000101000CA10100A10 Hanocladius ??????????????????????????? Heleniella 111010100100110000110110100 Heterotanytarsus 0A0000000000000000220200100 Heterotrissocladius 000000001000000000201C01A00 Hydrobaenus 000000000000000000A00100000 Krenosmittia 1111001000101100000201010A0 Limnophyes 111110100000010000110101A00 Lopescladius 111100110100010000011211110 Metriocnemus 11AAA010000001000BA1010AAA0 Orthocladius AAAAAA1 0 00A0A1 0 0 0EAC0100010 Parachaetocladius 111110100000010010110100100 Parakiefferiella 100A10100010110000A11101010 Paralimnophyes 1111101100000100001A0101A00 Parametriocnemus 1000001000001A0000B10100000 Parasmittia ? ? ? ?1? ? ??000010002?20000101 Propsilocerus 000AA00000000000001C0101000 Psectrocladius 0A0A1A0000A000000EA10101001 Pseudorthocladius 11110010000011000CC1010AA00 Pseudosmittia 1111AA11000001000B120001110 Rheocricotopus 0A001A0000000A00001101000A0 Smittia 111100110000010002C20100000 Stictocladius 11AAA0110100A1001AA11C111A0 Tvetenia 111100100000111000110100A00 Unniella 000011000000000000110101000 Zalutschia 01AA00001000000000A00100A00 aberrans ??????????????????????????? antecalvus 1111001000010101001???????? arcuatus ??????????????????????????? berui ??????????????????????????? biota ??????????????????????????? calakmulensis ??????????????????????????? croizati ??????????????????????????? dentispina ??????????????????????????? folius 111100100000010100210001001 herradurus ??????????????????????????? mateusi 1111001000000100021???????? musci 111100100000010100110001101 nerius 111110100000010000110101100 pectinata 111110100000010000110101000 pittieri ??????????????????????????? pluspilalus 11110010000001010A110001101 scalpellatus ??????????????????????????? skartveiti ??????????????????????????? sooretama ??????????????????????????? tuberculifera ??????????????????????????? ubatuba ??????????????????????????? venequatoriensis ??????????????????????????? virga ??????????????????????????? zempoalensis ??????????????????????????? zhengi ???????????????????????????

ORTHOCLADIINAE © 2004 Magnolia Press 17 ZOOTAXA TABLE 2 (continued) 594 Character no.: 111111111111111111111111 333344444444445555555555 678901234567890123456789 Aagaardia 001001100011001010000000 Apometriocnemus ???????????????????????0 Botryocladius 1010?A01000100011000?000 Brillia 000011110011110010000010 Bryophaenocladius 0120011002A2A10011120211 Chaetocladius 00A0A10A01AC0A00100000A0 Corynoneura 012011010E1101001?010000 Cricotopus AA20AA10A0010100100000A0 Diplocladius 000011110011000100000010 Doithrix 001001100112010010100120 Eukiefferiella 0120001A10AC01001A0AA010 Georthocladius s.str0010?01001021100011C0120 Gymnometriocnemus 012001100212A10011120211 Hanocladius ???????????????????????? Heleniella 002001100111110000001010 Heterotanytarsus 00101110001100000000A000 Heterotrissocladius 0010011A01A1000010000000 Hydrobaenus 00100110001100001000A000 Krenosmittia 0A200110000101001?030000 Limnophyes 00200100001101001A0010A0 Lopescladius 00200001000101001?031000 Metriocnemus 0AA0010101A1010011000000 Orthocladius 1020AAA00A0A010AA00000A0 Parachaetocladius 0010001001A2110011030120 Parakiefferiella 01C00A1000A100000?0000A0 Paralimnophyes 00C00A01001101001A0010A0 Parametriocnemus 0010A1100A11001001001000 Parasmittia 002011100001010011120010 Propsilocerus 00001111001A00000000A000 Psectrocladius 002000100EA10001010A0000 Pseudorthocladius 002000100112A100010301E0 Pseudosmittia 1020110A0E0C01001A1C0200 Rheocricotopus A020001000A100010000A000 Smittia 0020?1000001010000120010 Stictocladius 0A20101A01AC11000A031110 Tvetenia 0020A01010A1010011001000 Unniella 0020?1110001000010000000 Zalutschia 0010111000110001000000E0 aberrans ???????????????????????? antecalvus ???????????????????????? arcuatus ???????????????????????? berui ???????????????????????? biota ???????????????????????? calakmulensis ???????????????????????? croizati ???????????????????????? dentispina ???????????????????????? folius 002111110001110011120001 herradurus ???????????????????????? mateusi ???????????????????????? musci 002111110001110011120001 nerius 00200101001101001?0000?? pectinata 00200101001101001?0000?? pittieri ???????????????????????? pluspilalus 00211111000111001112000? scalpellatus ???????????????????????? skartveiti ???????????????????????? sooretama ???????????????????????? tuberculifera ???????????????????????? ubatuba ???????????????????????? venequatoriensis ???????????????????????? virga ???????????????????????? zempoalensis ???????????????????????? zhengi ????????????????????????

ORTHOCLADIINAE © 2004 Magnolia Press 19 ZOOTAXA FIGURE 1. Parsimony analysis of some genera of the Orthocladiinae with Diplocladius Kieffer, 594 Propsilocerus Kieffer and Brillia Kieffer combined as outgroup. A, strict consensus of shortest trees. Bremer supports for each branch are given as numbers above the branches. B, strict consensus 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 consensus 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 the basal three genera, 81 % for A. arcuatus plus A. zempoalensis, 78 % for Lopescladius plus Stictocladius, 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 the weights 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 show that croizati, aberrans, berui and pittieri all should be placed in Compterosmittia. The immatures of Compterosmittia are practically not separable from those of Limnophyes Eaton and Paralimnophyes Brundin. Immatures are, however, tentatively known for two species only. The imagines, on the other hand, differ in many respects. Probable synapomorphies as compared to Limnophyes and Paralimnophyes are the finely punctated wings with apical setae in most males and the known female, the bare squama, and the strongly extended costa. All previously described species of the genus have a comb-like megaseta while none of the new species have. The divisions in the reweighted cladogram reflect this. However, it may be more appropriate for C. croizati and C. dentispina to change position 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. The remaining species all have inferior volsella with a dorsal, bare, digitiform anterior projection and a ventral, posterior, rounded lobe. However, any subdivisions within the genus are premature considering the lacking knowledge of females and immatures. Also whether Hanocladius should remain as a separate genus as indicated by the not reweighted cladogram, or should be incorporated in Compterosmittia as suggested by the reweighted cladogram will have to await further material. It also is possible that C. virga may deserve a separate genus primarily based on the unique virga. Particularly the long virga with lateral lamellae and the pupa without thorn-like macrosetae, without posterior spines on the tergites and short male genital sac makes mateusi more similar to Gymnometriocnemus than to species of Antillocladius. We thus propose a separate genus, Litocladius new genus, with the single included species Litocladius mateusi new species. L. mateusi also differs from the remaining species by having a supraalar and a normally developed anal point with normal hair-like lateral setae.

20 © 2004 Magnolia Press MENDES ET AL. The remaining species are maintained within the genus Antillocladius. The many- ZOOTAXA toothed pecten epipharyngis of the known larvae constitute a uniquely derived synapomor- 594 phy. Also the thorn-like macrosetae of the pupae can be regarded as a unique synapomorphy. Although similar macrosetae occurs in Chaetocladius Kieffer they there are more spine-like. The approximately palmate seta anteriores (S I) also is a good synapomorphy although similar setae anteriores occur in Parasmittia Strenzke. The species of Antillocladius fall into two main groups partly reflected in the strict consensus tree of the reweighted characters; the first (eventually the nominal subgenus) includes antecalvus, skartveiti, arcuatus, zempolaensis and with some uncertainty herradurus; the second group includes the remaining species The first group has a unique, characteristic strongly set off inferior volsella with one circular lobe and with or without another circular posterior projection of the lobe. A. herradurus 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 the other species of the group. The pupa is known of A. antecalvus. It has a laterally expanded anal lobe as in some Gymnometriocnemus and Bryophaenocladius and thus differs from the other known pupae of the genus. The second group can be divided into three more or less distinct subgroups; one subgroup 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 a third subgroup containing the remaining species. A. calakmulensis has a virga consisting of several spines as opposed to the remaining species of the genus which either have no virga or a virga consisting of two spines. A. musci has a very similar hypopygium except for 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 derived synapomorphy. Most members of the last subgroup also have an inferior volsella with a more 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 volsella and two species have a very long inferior lobe apically divided in one species. There also are large variations in the shape of the virga ranging from absence to 2 short or very long spines. The anal point varies from long and parallel-sided with short lateral setae to long and triangular with strong setae. One species lacks setae on squama. However, both pupae and larvae are known for two of the species and the differences appear too slight to warrant any divisions. The characters are coded in ordered to indicate the phylogenetic position of the different species relative to known genera. The relationships among more basally placed genera thus should not be taken for relevant. In fact preliminary parsimony analyses of a wider range 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.

ORTHOCLADIINAE © 2004 Magnolia Press 21 ZOOTAXA Key to males of some orthoclad genera 594 1. Acrostichals scalpellate ...... 2 — Acrostichals not scalpellate, but may be hooked ...... Not keyed (According to Cranston et al. (1989) the acrostichals of Pseudosmittia Goetghebuer may 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 ...... 4 3. 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 Edward 4. 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 usually present, costal extension short to long, megaseta never dentate ...... 6 5. 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 Sæther 6. 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...... 7

7. 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...... 8 8. Wing with coarse punctation, without setae; gonocoxite with 2 lobes, the anteriormost appearing to be superior volsella and dorsal part of inferior volsella combined, extending from gonocoxite base to distal of midpoint; posteriormost lobe extending to close to insertion of gonostylus; virga absent...... Paralimnophyes Brundin — Wing with fine punctation, often with apical setae; volsellae variable, but never as above; 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

22 © 2004 Magnolia Press MENDES ET AL. Antillocladius Sæther ZOOTAXA 594 Antillocladius Sæther, 1981: 4; Coffman et al. (1986: 160); Cranston et al. (1983: 157); Cranston et al. (1989: 176); Sæther (1982: 471; 1984: 1).

Type species Antillocladius antecalvus Sæther, 1981, by original designation.

Other included species Antillocladius arcuatus Sæther, 1982; A. biota new species; A. calakmulensis new species; A. folius new species; A. herradurus new species; A. musci new species; A. pluspilalus Sæther, 1982; A. scalpellatus Wang and Sæther, 1993; A. skartveiti Andersen and Contreras-Ramos, 1999; A. sooretama new species; A. ubatuba new species; A. venequatoriensis new species; A. zempoalensis new species and A. zhengi Wang and Sæther, 1993.

Diagnostic characters The imagines can be separated from other orthoclad genera by having scalpellate acrostichals at least in the middle of scutum combined with a moderately to extremely long anal point with lateral setae. (Although A. pluspilalus in the original description is mentioned as apparently having non-scalpellate acrostichals, more recently found specimens show that scalpellate acrostichals are present.) All known females and several males have setae apical on the wing membrane. The combination of absence of thoracic horn and presence of thorn-like macrosetae will separate the pupae from other genera. The larva is distinguished by a palmate S I; pecten epipharyngis divided into about 8–15 teeth; anal segment protruding over posterior parapods; and anal setae absent or perhaps reduced to single seta on minute tubercle.

Generic diagnosis Imago Small 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 antennal ratio 0.50–1.70. Head. Eye naked, with or without minute pubescence between ommatids, without dorsomedial extension. Temporal setae divided into weak inner verticals, stronger outer verticals, 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 antepronotum, some distance from antepronotum or in the middle of scutum; prealars uniserial; supraalar 0–1; scutellars uniserial.

594 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 normal. 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 well developed. Anterior margin of sternapodeme nearly straight to strongly arcuate, oral projections barely indicated to strongly developed. Virga absent or consisting of 2–6 long, strong spines. Inferior volsella highly variable; simple and rounded or triangular; with anterior dorsal triangular or digitiform part and a more rounded ventral, posterior part; posterior lobe low to prominent; consisting of a posteriomedially directed, apically simple or bifid lobe; or circular with or without additional rounded posterior extension and conspicuously 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. Tergite IX divided or undivided with setae concentrated to lateral halves. Gonapophysis IX with large ventrolateral lobe covering apodeme lobe and apex of narrow, line-like dorsomesal lobe. Labia relatively large, pointed. Cerci small. Seminal capsules small to medium-sized, circular, with triangular neck. Spermathecal ducts with bends or loops, with weak bulbs before separate openings. Pupa Small 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 horn absent. Three weak precorneals situated in row or narrow triangle. Apparently two median and 0–1 lateral antepronotals. Dorsocentrals very small, in two groups of two. Thorax and wing sheath smooth. Abdomen. Tergite I without shagreen; tergites II–VIII covered with coarse, shagreen except along anterior margin; tergite IX with strong shagreen except along anterior and posterior margin. Sternites I and II bare; sternite III with median and lateral shagreen; sternites IV–VI with slightly more extensive median and lateral shagreen; sternites VII and VIII with extensive shagreen; sternite IX with anterolateral shagreen. Tergite II without

24 © 2004 Magnolia Press MENDES ET AL. posterior hooklets. Tergites II–VII or VIII with posterior row of weak spines. Conjunctives ZOOTAXA bare. Pedes spurii A and B absent. Apophyses weak, nearly straight. Segment II–VIII with 594 2 weak, hair-like L setae. Anal lobe with 3 thorn-like macrosetae, sometimes laterally expanded; male genital sac overreaching anal lobe, female genital sac not reaching apex of lobe. Larva Small 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, style large. 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 normally scales divided into 2–5 teeth). Chaetulae laterales simple, 3 strong and sclerotised, 4–5 weak, pointed and unsclerotised. Chaetulae basales simple or bifid. Mandible with apical tooth shorter than the combined widths of 4 inner teeth, fourth tooth sometimes joined to mola. Seta subdentalis triangular. Seta interna slender, of 4–5 nearly smooth to strongly serrated branches. Mentum with large, irregular triangular, sometimes medially notched median tooth higher than 5 pairs of lateral tooth, fourth pair smaller than fifth and apparently occasionally completely reduced. Ventromental plates small, visible only below 3 outermost teeth, no beard. Maxilla without pecten galearis, lacinial chaeta reduced to rounded 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. Procercus apparently absent, but a vestigial tubercle with one short seta may represent procercus at least 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 strong apical claws.

Key to the males of Antillocladius Sæther

1. Squama bare (Fig. 16D); Mexico...... A. zempoalensis new species — Squama with setae...... 2 2. Anal point nearly parallel-sided with numerous, weak, partly anterior directed setae; inferior volsella either pointed triangular or consisting of a long apically simple or bifid 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 more rounded ventral lobe or circular with or without additional rounded posterior extension and 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

ORTHOCLADIINAE © 2004 Magnolia Press 25 ZOOTAXA extension of moderate length ...... 5 594 4. Inferior volsellae triangular, leaf-like (Fig. 5D); Brazil...... A. folius new species — Inferior volsella small, sharply pointed (Fig. 3D); Brazil...... A. biota new species 5. Inferior volsella apically bifid (Fig. 13C); Brazil ...... A. sooretama new species — Inferior volsella simple (Fig. 14D); Brazil, Venezuela ...... A. ubatuba new species 6. 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 ...... 11 7. 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...... 8 8. 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 ...... 10 9. 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 Sæther 10. Virga about 2/3 as long as phallapodeme; squama with 5–6 setae; cell m3+4 with several 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...... 13 12. Virga consisting of 6 spines; inferior volsella with digitiform dorsal anterior lobe covering 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, USA A. arcuatus Sæther — Inferior volsella a well set off circular lobe; wing membrane usually with setae...... 14 14. 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

26 © 2004 Magnolia Press MENDES ET AL. Key to known pupae of Antillocladius Sæther ZOOTAXA 594 1. Antennal sheath with pearl row...... 2 — Antennal sheath lacking pearl row ...... 3 2. 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 species 3. 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 ...... 2 2. 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 about 12 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, Malaise trap, M.V. Yamada (UFSCar). VENEZUELA: Aragua, Parque Nacional Henri Pittier, Rancho Grande, 10°21.047'N, 67°41.198'W, 21 males, 16–18.ix.1999, about 1000 m a.s.l., sweep net & 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 the genus by having 0–3 apical setae on the wing membrane; squama with 1–3 setae; few scalpellate to non-scalpellate acrostichals in the middle of scutum; virga absent; and inferior

ORTHOCLADIINAE © 2004 Magnolia Press 27 ZOOTAXA volsella large with anterior bare, strongly set off rounded projection and low posterior 594 lobe. The female differs from other known females by having tergite IX undivided, but tergite setae still in two groups. The pupa has thorn-like macrosetae and anal lobe laterally expanded.

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 from Saint Vincent. Sæther (1981) described the male and female in detail. The males from Venezuela vary slightly from the specimens from Saint Vincent, as they have scalpellate acrostichals (not observed from St. Vincent), the transverse sternapodeme is straight to weakly arcuate, and the shape of gono-

28 © 2004 Magnolia Press MENDES ET AL. stylus varies slightly. The pupa is described below. The specimens from Venezuela were ZOOTAXA all taken in mountainous areas with rainforest. The Brazilian specimen was taken in a Cer- 594 rado formation in the northeast São Paulo State. Pupa (n = 1) Total length 2.04 mm. Exuviae pale, nearly transparent. Cephalothorax. Frontal apotome as in Fig. 2C. Ocular field with three pairs of setae. Antennal sheath smooth. One median antepronotal, 14 µm long. Precorneals in single row (Fig. 2D), each about 20 µm long; distance between anterior and median seta 9 µm, between median and posterior seta 5 µm, between anterior and posterior seta 16 µm, other thoracic setae not measurable. Abdomen (Fig. 2A–B). Numbers of caudal spines on tergites II–VIII as: 23, 24, 22, 22, 23, 23, 20. Length (in µm) of longest caudal spines on tergites II–VIII as: 8, 11, 11, 9, 9, 9, 7. Anal lobe (Fig. 2E) with three thorn-like macrosetae 19, 18 and 17 µm long. Geni- tal sac overreaches anal lobe by 47 µm.

Antillocladius arcuatus Sæther

Antillocladius arcuatus Sæther, 1982: 474, Fig. 5.

Material examined: Type material as in Sæther (1982). BRAZIL: São Paulo, Cajuru, Fazenda Rio Grande, 21°12'S, 47°09'W, 2 males, 18.xii.1999–10.i.2000, Malaise trap, G. Melo & F. Nascimento (MZUSP). MEXICO: Nuevo Leòn, Santiago, 30 km west of Cola Caballo on road to Laguna de Sánchez, 2 males, 19.ix.1998, light trap, T. Andersen & A. Contreras-Ramos (ZMBN); Nuevo Leòn, Allende, Rio Ramos at Raices, 2 km west of highway 85, 2 males, 20.ix.1998, sweep net & light trap, T. Andersen & A. Contreras- Ramos (ZMBN). VENEZUELA, Falcón, Rio Mitare near San Luis, 11°07.930'N, 69°39,184'W, 8 males, 07.vi.2001, 589 m a.s.l., R.W. Holzenthal, R. Blahnik, H. Paprocki & C. Cressa (2 MIZA , 6 ZMBN). Diagnostic characters: The male imago can be separated from other members of the genus by having bare wing membrane; squama with 2–3 setae; acrostichals starting in front, non-scalpellate anteriorly, scalpellate in the middle of scutum; virga absent; anal point thin, tapering to point; and inferior volsella strongly projecting and lobe-like. Remarks: This species previously was known only from the type locality in southeast USA. The present paper adds records from Brazil, Mexico and Venezuela. The specimens have been taken close to fast flowing streams and rivers in Mexico and Venezuela, and close to a swampy area in Brazil.

Type material: Holotype male, BRAZIL: São Paulo State, Parque Estadual Intervales, sede de pesquisa, 16.viii.2000, larvae taken on tree trunks, H.F. Mendes (MZUSP, slide mounted in Euparal). Paratypes: 2 males, as holotype (ZMBN). Diagnostic characters: The male imago can be separated from other members of the genus by having bare wing membrane; squama with a few setae; long costal extension; scalpellate acrostichals starting near antepronotum; long virga; nearly parallel-sided anal point with blunt apex and weak lateral setae; and inferior volsella short and pointed triangular. Etymology: Named after the Biota program supported by FAPESP; the name is a noun in apposition. Male (n = 1–3). Total length 1.91–2.12 mm. Wing length 1.19–1.25 mm. Total length / wing length 1.60–1.74. Wing length / length of profemur 2.45–2.54. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 0.60–0.61, ultimate flagellomere 241–294 µm. Temporal setae 8, including 2–3 inner verticals, 1–2 outer verticals and 3–4 postorbitals. Clypeus with 6–7 setae. Ten- torium, stipes and cibarial pump as in Fig. 3A; tentorium 126–131 µm long, 10 µm wide at sieve pore; stipes 88 µm long. Palpomere lengths in µm: 18–25, 30–38, 56–63, 71–76, 101–103. Third palpomere with 3–5 sensilla clavata subapically, longest 13 µm long. Thorax (Fig. 3B). Antepronotum with 1–3 setae. Dorsocentrals 8–11; acrostichals 11– 12, all scalpellate starting close to antepronotum; prealars 2–3. Scutellum with 6 setae. Wing (Fig. 3C). VR 1.38–1.64. Costal extension 76–88 µm long. Brachiolum with 1 seta, R with 0–3 setae, R1 with 0–1 seta, other veins bare. Wing membrane bare. Squama with 3–6 setae. Legs. Spur of front tibia 38–50 µm long, spurs of middle tibia 17–20 µm and 25–33 µm long, of hind tibia 17–20 µm and 43–51 µm long. Width at apex of front tibia 28–33 µm, of middle tibia 28–30 µm, of hind tibia 33–38 µm. Hind tibial comb with 9–11 setae, shortest seta 13–18 µm long, longest seta 28–33 µm long. Lengths (in µm) and proportions of legs as in Table 3.

TABLE 3. Lengths (in µm) and proportions of legs of Antillocladius biota new species.

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

p1 468–497 517–604 390–420 224–234 127–136 78 49–58 0.69–0.77 2.78–3.18 2.46–4.65 2.5–2.7

p2 497–525 525–556 205–224 107–175 78–88 39–58 29–39 0.39–0.43 3.59–4.85 4.61–5.04 2.4–2.5

p3 546–575 595–643 341 175–205 136–156 68 49–58 0.53–0.57 3.26–3.47 3.34–3.57 3.0–3.5

FIGURE 3. Antillocladius biota new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; F–gonostylus.

ORTHOCLADIINAE © 2004 Magnolia Press 31 ZOOTAXA Hypopygium (Fig. 3D–F). Anal point 56–58 µm long; 20–25 µm wide at base, with 594 13–14 strong lateral setae, void of microtrichia except at base. Laterosternite IX with 7–8 setae. Phallapodeme 68–70 µm long. Transverse sternapodeme slightly arcuate; with rounded oral projections, 63–76 µm long. Virga 58 µm long. Gonocoxite 119–134 µm long; with long, low inferior volsella, with rounded projection orally. Gonostylus 76–81 µm long; megaseta 13–15 µm long. HR 1.57–1.77, HV 2.51–2.79.

Antillocladius calakmulensis new species (Fig. 4A–G)

Type material: Holotype male, MEXICO: Campeche Province, Reserve Biosphere Calak- mul, Campamento Carola, 20.ix.1997, A. Contreras-Ramos et al. (ZMBN Type no. 398, slide mounted in Canada balsam). Diagnostic characters: The male imago can be separated from other members of the genus by having a virga consisting of 6 short spines; apical setae on the wing membrane; squama with a few setae; short costal extension; scalpellate acrostichals in the middle of scutum; anal point long and triangular, with strong lateral setae; and inferior volsella prominent with anterior digitiform projection equally long as and partly covering rounded posterior lobe. Etymology: Named after the Reserva Biosphere Calakmul in SE Mexico, using the Latin suffix, –ensis, denoting the place of origin. Male (n = 1). Total length 2.13 mm. Wing length 1.14 mm. Total length / wing length 1.86. Wing length / length of profemur 2.55. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. (Fig. 4A) AR 1.27, ultimate flagellomere 396 µm long. Temporal setae 9, including 3 inner verticals, 1 outer vertical and 5 postorbitals. Clypeus with 9 setae. Tento- rium, stipes and cibarial pump as in Fig. 4B; tentorium 121 µm long, 22 µm wide at sieve pore; stipes 121 µm long, 35 µm wide. Palpomere lengths in µm: 28, 40, 76, 73, 97. Third palpomere apparently with 1 sensillum clavatum subapically, 10 µm long. Thorax (Fig. 4C). Antepronotum with 2 setae. Dorsocentrals 10; acrostichals 13, all scalpellate starting some distance from antepronotum; prealars 5, supraalar 1. Scutellum with 8 setae. Wing (Fig. 4D). VR 1.32. Costal extension 22 µm long. Brachiolum with 1 seta, M1+2 with 3 setae, other veins bare. Wing membrane with 16 setae in cell r4+5. Squama with 5 setae. Legs. Spur of front tibia 54 µm long, spurs of middle tibia 29 µm and 26 µm long, spurs of hind tibia 35 µm and 19 µm long. Width at apex of front tibia 32 µm, of middle tibia 33 µm, of hind tibia 40 µm. Hind tibial comb with 12 setae, shortest seta 12 µm long, longest seta 35 µm long. Lengths (in µm) and proportions of legs as in Table 4.

FIGURE 4. Antillocladius calakmulensis new species, male imago. A–head; B–cibarial pump, tentorium and stipes; C–thorax; D–wing; E–anal point and tergite IX and dorsal aspect of left gonocoxite; F–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; G–gonostylus.

ORTHOCLADIINAE © 2004 Magnolia Press 33 ZOOTAXA TABLE 4. Lengths (in µm) and proportions of legs of Antillocladius calakmulensis new species. 594 fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR p1 449 507 378 221 147 90 57 0.74 2.59 2.54 2.7 p2 449 466 212 123 87 49 41 0.46 3.83 4.31 2.8 p3 507 580 327 229 139 74 41 0.56 2.93 3.33 6.5

Hypopygium (Fig. 4E–G). Anal point 59 µm long, 33 µm wide at base, with 20 strong lateral setae, void of microtrichia except at base. Laterosternite IX with 5 setae. Phallapo- deme 76 µm long. Transverse sternapodeme arcuate, with rounded oral projections, 69 µm long. Virga consisting of 6 spines, 25 µm long. Gonocoxite 137 µm long, with long, low inferior volsella, with rounded projection orally. Gonostylus 86 µm long, megaseta 12 µm long. HR 1.60, HV 2.48.

Antillocladius folius new species (Figs 5A–F, 6A–E, 7A–F)

Type material: Holotype male with pupal and larval skins, BRAZIL: São Paulo State, Ribeirão Preto, 28.v.1997, H.F. Mendes (MZUSP, slide mounted in Euparal). Paratypes: 2 males with pupal exuviae, 2 larvae, as holotype (ZMBN); São Paulo State, Ribeirão Preto, 2 males, 5.iii.1997, H.F. Mendes (FIOCruz); São Paulo State, Parque Estadual Intervales, Boca gruta Minotauro, 1 male, 27–28.x.2001, CDC, E.A. Galati (MZUSP); São Paulo State, Parque Estadual Intervales, Boca gruta Colorida, 1 male, 27–28.x.2001, CDC, E.A. Galati (MZUSP); São Paulo State, Estação Biológica Boracéia, córrego Venerando, 3 males, 12.vii.2001, light trap, C.G. Froehlich et al. (MZUSP); São Paulo State, Parque Estadual Jaraguá, local-2 E, 1male, 14.xii.2000, M.V. Yamada (UFSCar); São Paulo State, Parque Estadual Jaraguá, Pico do Jaraguá, 1 male, 14.xii.2000, M.V. Yamada (UFSCar); São Paulo State, Parque Estadual Jaraguá, Alojamento, 1 male, 14.xii.2000, M.V. Yamada (UFSCar); São Paulo State, Parque Estadual Jaraguá, 1 male, 1999, M.V. Yamada (UFS- Car); São Paulo State, Teodoro Sampaio, Parque Estadual Morro do Diabo, foz Córrego Taquara, 7 males, 27.vii.2000, sweep net, H.F. Mendes (3 ZSM, 4 ZMBN); São Paulo State, Ubatuba, Picinguaba, Córrego da Sede, 23º21'502''S, 44º49'128''W, 1 male, 17.ii.2002, Malaise trap, F.O. Roque (UFSCar); Santa Catarina State, São Bento do Sul, 26°19'25.6''S, 48°18'26.5''W, 1 male,13–16.x.2001, Malaise trap, M.V. Yamada (UFSCar). Diagnostic characters: The male imago can be separated from other members of the genus by having bare wing membrane and squama with a few setae; acrostichals starting in front, anterior ones hair-like, posterior ones scalpellate; virga well developed; anal point only slightly tapering to blunt apex, with weak lateral setae; and inferior volsella consisting of triangular, pointed lobe. The pupa differs from that of other known species by having antennal sheath with pearl row, and thorn-like anal macrosetae less than half as long as

34 © 2004 Magnolia Press MENDES ET AL. the long male gonopodal sheaths. The larva has mentum with 5 lateral teeth and antennal ZOOTAXA blade subequal in length to flagellum. 594 Etymology: From Latin, folium, meaning leaf, referring to the shape of the inferior volsellae. Male (n = 10, except when otherwise stated). Total length 1.93–2.40, 2.12 mm. Wing length 1.19–1.36, 1.27 mm. Total length / wing length 1.49–1.77, 1.66. Wing length / length of profemur 2.57–2.97, 2.71. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 0.87–1.13, 1.00, ultimate flagellomere 348–402, 375 µm long. Temporal setae 8–9, 8, including 3 inner verticals, 2–3, 2 outer verticals and 3 postorbitals. Clypeus with 6 setae. Tentorium, stipes and cibarial pump as in Fig. 5A; tentorium 121–126 (2) µm long, 33–38 (2) µm wide at sieve pore; stipes 76 (2) µm long, 38 (1) µm wide. Palpomere lengths in µm: 22–25, 24; 38–46, 40; 86–88, 87; 94–114, 107 (7); 115–139, 125 (5). Third palpomere with 6–7, 6 sensilla clavata subapically; longest 15–18, 17 µm long. Thorax (Fig. 5B). Antepronotum with 3–4, 4 setae. Dorsocentrals 9–12, 10; acrostichals 14–20, 18, anterior six hair-like, posterior all scalpellate, starting some distance from antepronotum; prealars 3. Scutellum with 4–6, 5 setae. Wing (Fig. 5C). VR 1.50–1.54 (2). Costal extension 63–88, 83 µm long. Brachiolum with 1 seta, R with 0–2, 1 setae, other veins bare. Wing membrane bare. Squama with 4–7, 6 setae. Legs. Spur of front tibia 40–51 (2) µm long; spurs of middle tibia 18–20, 19 µm and 28–33, 30 µm long; spurs of hind tibia 15–25, 21 µm and 40–48, 43 µm long. Width at apex of front tibia 33 (2) µm; of middle tibia 28–33, 31 µm; of hind tibia 36–40, 39 µm. Hind tibial comb with 11–13, 12 setae; shortest seta 15–20, 16 µm long; longest seta 25– 35, 30 µm long. Lengths (in µm) and proportions of legs as in as in Table 5.

TABLE 5. Lengths (in µm) and proportions of legs of Antillocladius folius new species, (n=2–10).

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR p1 439–441 507 400–408 224–244 136–185 78 49 0.79–0.81 2.53–2.67 2.39–2.31 2.3–2.5 p2 439–517, 458–527, 195–224, 107–117, 76–84, 49–58, 36–42, 0.37–0.49, 4.00–4.50, 4.52–5.30, 2.5–2.56, 480 487 203 109 78 51 39 0.42 4.23 4.99 2.5 p3 507–546, 536–634, 292–331, 156–185, 127–136, 49–68, 38 0.49–0.62, 3.25–3.73, 3.15–3.70, 4.5 523 579 312 129 129 58 0.54 3.51 3.53

Hypopygium (Fig. 5D–F). Anal point 58–66, 61 µm long; 23–33, 28 µm wide at base; with 10–17, 15 weak lateral setae; void of microtrichia except at base. Laterosternite IX with 6 setae. Phallapodeme 76–96, 80 µm long. Transverse sternapodeme nearly straight, with rounded oral projections; 63–73, 68 µm long. Virga 50–66, 63 µm long. Gonocoxite 144–177, 154 µm long, with well set–off, leaf–like inferior volsella. Gonostylus 70–76, 73 µm long; megaseta 13–15, 14 µm long. HR 1.93–2.31, 2.10; HV 2.63–3.29, 2.90.

FIGURE 5. Antillocladius folius new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; F–gonostylus.

36 © 2004 Magnolia Press MENDES ET AL. Pupa (n = 1–3). ZOOTAXA Total length 2.37 mm. Colour brownish yellow, legs, antennae and wings dark brown; 594 exuviae pale transparent. Cephalothorax. Frontal apotome as in Fig. 6C. Ocular field with 3 pairs of setae. Antennal sheath with pearl row. One prealar, about 23 µm long. Precorneals in narrow triangle (Fig. 6D). Abdomen (Fig. 6A–B, E). Numbers of caudal spines on tergites II–VIII as: 25–33, 20– 30, 23–31, 24–31, 27–29, 23–26, 33. Length of longest caudal spine on tergites II–VIII (in µm) as: 7, 7, 7, 7–9, 8–11, 12–13, 10–13. Anal lobe with three thorn-like macrosetae 28– 36, 30–34, and 34–40 µm long. Genital sac overreaches anal lobe with 71–86 µm.

FIGURE 6. Antillocladius folius new species, pupa. A–tergites I–IX; B–sternites I–IX; C–frontal apotome; D–thorax, anterior part, lateral view; E–detail of anal lobe.

FIGURE 7. Antillocladius folius new species, larva. A–antenna; B–S I setae; C–mentum; D–mandible; E–premandible; F–anal end.

Fourth instar larva (n = 1–2). Total length 3.0–3.1 mm. Head capsule 281 µm long. Postmentum 98 µm long. Colour of thoracic segments brown, head and abdomen amber yellow. Head. Antenna as in Fig. 7A; segment lengths in µm: 28–30, 15–17, 5–7, 2–3, 2–3. Blade 28 µm long, as long as flagellum; apical style of second segment 4 µm long. Seta I nearly palmate (Fig. 7B), other setae simple, labral lamella inconspicuous. Chaeta media serrate, chaetulae laterales simple, chaetulae basales bifid. Pecten epipharyngis with about 12 teeth. Mandible (Fig. 7D) 122–127 µm long, with apical tooth and four inner teeth; seta

38 © 2004 Magnolia Press MENDES ET AL. subdentalis triangular; seta interna with four branches. Premandible as in Fig. 7E, 45–66 ZOOTAXA µm long. Mentum (Fig. 7C) 88–95 µm wide, with irregular 27 µm long, median tooth and 594 five lateral teeth. Ventromental plates small. Seta submenti 43 µm long Abdomen without distinct setae. Anterior parapods fused, with numerous claws; posterior parapods well developed, with strong apical claws, about 105 µm long (Fig. 7F). Procerci absent. Anal tubules 89 µm long, 28 µm wide.

Antillocladius herradurus new species (Fig. 8A–G)

Type material: Holotype male, MEXICO: Campeche Province, Reserva Biosphere Calak- mul, Zona Arqueològica, Aguada Grande, 18°07'26.7''N, 89°48'56.7''W, 19.ix.1996, 265 m a.s.l., light trap, A. Contreras-Ramos et al. (ZMBN, type no. 399, slide mounted in Can- ada balsam). Diagnostic characters: The male imago can be separated from other members of the genus by having a few setae apically on wing membrane; squama with a few setae; costa barely extended; acrostichals starting in front, anterior ones hair-like, posterior ones scalpellate; virga short; anal point triangular, tapering to blunt apex, with strong lateral setae; and inferior volsella strongly projecting posteriomedially, ending in two rounded lobes. Etymology: From Spanish, herradura, meaning horseshoe, referring to the shape of the virga. Male (n = 1). Total length 1.89 mm. Wing length 1.06 mm. Total length / wing length 1.78. Wing length / length of profemur 2.32. Coloration brown, thorax pale with brown vittae, tarsi of all legs light brown. Head. (Fig. 8A) AR 1.13, ultimate flagellomere 372 µm. Temporal setae 10, including 3 inner verticals, 4 outer verticals and 3 postorbitals. Clypeus with 5 setae. Tentorium, stipes and cibarial pump as in Fig. 8B; tentorium 113 µm long, 18 µm wide at sieve pore; stipes 99 µm long. Palpomere lengths in µm: 15, 35, 87, 76, 102. Third palpomere with 2 sensilla clavata subapically, longest 12 µm long. Thorax (Fig. 8C). Antepronotum with 4 setae. Dorsocentrals 8; acrostichals 14, of which the 2 anterior broadly hair-like, the 12 posterior scalpellate, starting close to antepronotum; prealars 3. Scutellum with 6 setae. Wing (Fig. 8D). VR 1.38. Costal extension 28 µm long. Brachiolum with 1 seta, other veins bare, wing membrane with 2 setae in cell r4+5. Squama with 4 setae. Legs. Spur of front tibia 44 µm long, spurs of middle tibia 28 µm and 21 µm long, spurs of hind tibia 40 µm and 21 µm long. Width at apex of front tibia 29 µm, of middle tibia 29 µm, of hind tibia 37 µm. Hind tibial comb with 12 setae, shortest seta 19 µm long, longest seta 29 µm long. Lengths (in µm) and proportions of legs as in Table 6.

FIGURE 8. Antillocladius herradurus new species, male imago. A–head; B–cibarial pump, tentorium and stipes; C–thorax; D–wing; E–anal point and tergite IX and dorsal aspect of left gonocoxite; F–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; G–gonostylus.

40 © 2004 Magnolia Press MENDES ET AL. TABLE 6. Lengths (in µm) and proportions of legs of Antillocladius herradurus new species. ZOOTAXA 594 fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR p1 458 498 335 188 114 74 49 0.67 3.04 2.85 3.1 p2 453 449 184 106 82 49 41 0.41 3.91 4.91 4.0 p3 523 539 302 163 131 65 41 0.56 3.41 3.51 7.3

Hypopygium (Fig. 8E–G). Anal point 55 µm long, 35 µm wide at base, with 18 strong lateral setae, void of microtrichia except at base. Laterosternite with 6 setae. Phallapodeme 81 µm long. Transverse sternapodeme arcuate, with weak, rounded oral projections; 57 µm long. Virga apparently consisting of 2 spines, 18 µm long. Gonocoxite 137 µm long, with well developed, set-off inferior volsella. Gonostylus 68 µm long, megaseta 17 µm long. HR 2.02, HV 2.78.

Antillocladius musci new species (Figs 9A–F, 10A–I, 11A–F, 12A–F)

Type material: Holotype male with larva and pupal exuviae, BRAZIL: São Paulo State, Ribeirão Preto, Lake Monte Alegre, 21°11'S, 47°51'W, 5.iii.1997, H.F. Mendes (MZUSP, slide mounted in Euparal). Paratypes: 4 males, 2 females with pupal exuviae, 1 female, 5 larvae, as holotype (ZMBN); 1 male, 7 males with pupal exuviae, as holotype except 28.v.1997 (MZUSP), 4 males, 7 females, 9 pupal exuviae, as holotype except 9.iv.1998 (MZUSP); 3 males, 1 female with larval and pupal exuviae, as holotype except 4.xii.1999 (MZUSP); 3 males, São Paulo State, Ubatuba, 2.i.2001, H.F. Mendes (ZSM); 2 males with larval and pupal exuviae, 7 males with pupal exuviae, 1 male, São Paulo State, São Carlos, 9.xii.1999, H.F. Mendes (FIOCruz, UFSCar); 5 males, São Paulo, Pedregulho, Furna São Pedro, 7.vii.2000, S. Mateus (MZUSP); 2 males, 1 male with pupal exuviae, 3 larvae, 1 pupal exuviae, Minas Gerais State, Uberaba, Peirópolis, Estação Paleontológica, 6.xii.1998, H.F. Mendes (MZUSP); 2 males, São Paulo State, Ubatuba, Praia das Tonin- has, 9.xi.2000, sweep net, H.F. Mendes (MZUSP); 5 males, 1 male with larval and pupal exuviae, São Paulo State, Santa Rosa de Viterbo, Fazenda Cascata, 4.vi.2000, reared from larvae collected on tree trunks, H.F. Mendes (MZUSP). Diagnostic characters: The male imago can be separated from other members of the genus by having a few apical setae on the wing membrane; setose squama; acrostichals all scalpellate, starting some distance from antepronotum; long virga; anal point tapering, with strong lateral setae; and inferior volsella rectangular with anterior digitiform projection. The female has divided tergite IX and a wing length of about 2.0 mm. The pupa differs from that of other known species by having antennal sheath with pearl row, and thorn- like anal macrosetae more than half as long as the distance by which the male genital sac is

ORTHOCLADIINAE © 2004 Magnolia Press 41 ZOOTAXA overreaching the anal lobe. The larva has about 15 teeth in pecten epipharyngis, mentum 594 with 5 lateral teeth and antennal blade longer than flagellum. Etymology: From Latin, muscus, moss, indicating the habitat of the larvae.

FIGURE 9. Antillocladius musci new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; F–gonostylus.

42 © 2004 Magnolia Press MENDES ET AL. Male (n = 10). ZOOTAXA Total length 2.30–2.71, 2.48 mm. Wing length 1.32–1.55, 1.45 mm. Total length / 594 wing length 1.65–1.75, 1.71. Wing length / length of profemur 2.26–2.50, 2.42. Coloration brown, thorax dark brown. Head. AR 1.39–1.65, 1.55; ultimate flagellomere 428–544, 531 µm long. Temporal setae 8–16, 12; including 2–4, 3 inner verticals; 1–6, 4 outer verticals; and 4–7, 5 postorbitals. Clypeus with 6–10, 8 setae. Tentorium, stipes and cibarial pump as in Fig. 9A; tentorium 137–145, 142 µm long; 15–18, 17 µm wide at sieve pore. Stipes 142–156, 147 µm long; 35–47, 41 µm wide. Palpomere lengths in µm: 20–39, 28; 44–59, 51; 94–113,107; 90–101, 98 (8); 120–141, 134 (8). Third palpomere with 2–3, 3 (5) sensilla clavata subapically, longest 14–17, 16 µm long. Thorax (Fig. 9B) Antepronotum with 3–6, 5 setae. Dorsocentrals 8–15, 12; acrostichals 12–28, 18, all scalpellate starting close to antepronotum; prealars 3–5, 4. Scutellum with 7–14, 10 setae. Wing (Fig. 9C). VR 1.32–1.44, 1.39. Costal extension 41–61, 51 µm long. Brachiolum with 1 seta, R with 0–4, 2 setae; other veins bare. Wing membrane with 0–11, 6 setae in cell r4+5; 0–1, 0 seta in cell r2+3. Squama with 9–16, 12 setae. Legs. Spur of front tibia 55–65, 61 µm long; spurs of middle tibia 28–35, 31 and 19– 29, 25 µm long; of hind tibia 50–63, 59 and 17–36, 23 µm long. Width at apex of front tibia 37–43, 39 µm; of middle tibia 33–41, 38 µm; of hind tibia 45–55, 50 µm. Hind tibial comb with 12–15, 13 setae; shortest seta 17–23, 19 µm long; longest seta 36–48, 42 µm long. Lengths (in µm) and proportions of legs as in Table 7.

TABLE 7. Lengths (in µm) and proportions of legs of Antillocladius musci new species, male (n=2–10).

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR p1 515–637, 637–735, 392–515, 229–278, 131–172, 90–115, 49–73, 0.61–0.71, 2.91–3.30, 2.77–3.02, 2.2–3.3 621 719 449 261 155 101 65 0.66 3.04 2.86 p2 523–662, 564–678, 229–278, 114–147, 82–106, 49–65, 41–57, 0.40–0.44, 4.18–4.61, 4.53–5.07, 2.5–3.2, 645 637 253 139 95 57 47 0.42 4.40 4.83 2.7 p3 621–711, 686–817, 351–498, 188–229, 131–172, 74–82, 49–65, 0.51–0.61, 3.48–3.81, 3.06–3.72, 3.2–3.5 686 784 425 212 162 77 57 0.58 3.67 3.30

Hypopygium (Fig. 9D–F). Anal point 70–85, 79 µm long; 41–48, 46 µm wide at base; with 14–20, 17 strong, lateral setae, void of microtrichia except at base. Laterosternite with 4–8, 6 setae. Phallapodeme 94–108, 103 µm long. Transverse sternapodeme arcuate with rounded oral projections; 103–110, 106 µm long. Virga 80–90, 84 µm long. Gono- coxite 160–172, 167 µm long; with low inferior volsella. Gonostylus 88–119, 100 µm long, megaseta 10–13, 11 µm long. HR 1.53–1.92, 1.65; HV 2.28–2.77, 2.51.

FIGURE 10. Antillocladius musci new species, female imago. A–head; B–cibarial pump, tentorium and stipes; C–thorax; D–wing; E–genitalia ventral view; F–genitalia dorsal view; G–dorsomesal lobe of gonapophysis VIII; H–ventrolateral lobe of gonapophysis IX; I–apodeme lobe of gonapophysis VIII.

44 © 2004 Magnolia Press MENDES ET AL. Female (n = 8–10). ZOOTAXA Total length 2.04–2.55, 2.32 mm. Wing length 1.39–1.43, 1.41 mm. Total length / 594 wing length 1.44–1.77, 1.59. Wing length / length of profemur 2.17–2.33, 2.27. Colour amber–yellow, antennae brown. Head (Fig. 10A). Pedicel length 44–52, 47 µm, width 64–76, 69 µm. Flagellomeres length / width in µm: 68–80, 74 / 32–36, 35; 52–60, 55 / 24–28, 25; 52–64, 56 / 24–28, 26; 52–64, 58 / 24–28, 26; 108–132, 116 / 24; 20–24, 23. AR 0.43–0.50, 0.46. Temporal setae 10–15, 12; including 2–4, 3 inner verticals; 2–5, 3 outer verticals; and 3–7, 5 postorbitals. Clypeus with 6–12, 9 setae. Tentorium, stipes and cibarial pump as in Fig. 10B; tentorium 124–142, 135 (6) µm long; 11–14, 12 (5) µm wide at sieve pore; 18–26, 21 (5) µm wide at posterior tentorial pit. Stipes 141–145, 142 (5) µm long; 27–35, 33 (5) µm wide. Pal- pomere lengths in µm: 22–28, 25; 44–51, 49; 69–99, 88; 90–97, 93 (6); 116–120, 117 (5). Thorax (Fig. 10C). Antepronotum with 4–7, 5 setae. Dorsocentrals 12–20, 16; acrostichals 18–23, 20, all scalpellate starting close to antepronotum; prealars 3–5, 4. Scutellum with 10–13, 11 setae. Wing (Fig. 10D). VR 1.33–1.44, 1.37. Brachiolum with 1 setae, R with 14–19, 16 setae; R1 with 7–13, 10 setae; R4+5 with 19–24, 22 setae; M1+2 with 30–37, 35 setae; M3+4 with 15–25, 18 setae; Cu with 10–14, 12 setae; Cu1 with 8–13, 10 setae; An with 13–20,

15 setae; other veins bare. Cell r4+5 with 109–137, 129 setae; cell m1+2 with 91–151, 112 setae; cell m3+4 with 44–78, 56 setae; cells cu and an together with 36–72, 56 setae. Squama with 8–12, 10 setae. Legs. Spur of front tibia 39–47, 44 µm long; spurs of middle tibia 29–33, 32 and 25– 28, 26 µm long; of hind tibia 59–65, 61 and 22–25, 23 µm long. Width at apex of front tibia 44–48, 46 µm; of middle tibia 44–47, 46 µm; of hind tibia 57–62, 59 µm. Hind tibial comb with 11–12, 12 setae; shortest seta 19–24, 22 µm long; longest seta 41–49, 47 µm long. Lengths (in µm) and proportions of legs as in Table 8.

TABLE 8. Lengths (in µm) and proportions of legs of Antillocladius musci new species, female (n=3–10).

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR p1 605– 667– 425– 212– 147– 82– 49– 0.61– 3.16– 2.85– 1.7–2.3 653, 622 711, 692 458, 443 237, 227 163, 153 106, 94 65, 57 0.68, 0.64 3.44, 3.32 3.04, 2.95 p2 612– 629– 245– 123– 82–106, 49–74, 40– 0.38– 4.58– 4.82– 1.9–2.3, 678, 642 670, 645 269, 255 131, 129 98 59 43, 41 0.43, 0.40 5.05, 4.72 5.23, 5.08 2.2 p3 645– 735– 400– 196– 147– 65–74, 41– 0.51– 3.47– 3.25– 1.8–2.4, 711, 681 809, 765 433, 418 204, 201 196, 163 71 49, 47 0.57, 0.55 4.05, 3.86 3.65, 3.42 2.0

Genitalia (Fig. 10E–I). Gonocoxite with 6–10, 8 setae. Tergite IX divided, each side with 11–17, 15 setae. Cercus 65–97, 81 µm long; with 22–25, 23 setae. Seminal capsule 146–157 (3) µm long, including 39–47 (3) µm long neck. Notum 106–116, 112 µm long.

FIGURE 11. Antillocladius musci new species, pupa. A–tergites I–IX; B–sternites I–IX; C–frontal apotome; D–thorax, anterior part, lateral view; E–detail of anal lobe, female, ventral view; F–detail of anal lobe, male, dorsal view.

46 © 2004 Magnolia Press MENDES ET AL. Cephalothorax. Frontal apotome as in Fig. 11C. Ocular field with three pairs of seta. ZOOTAXA Antennal sheath with pearl row. Antepronotals 2, about 49 (1) µm long. One prealar, about 594 46 (1) µm long. Precorneals in single row to narrow triangle (Fig. 11D), each about 48 (1) µm long; distance between anterior and median seta 7 (1) µm, between median and posterior seta 17 (1) µm, between anterior and posterior seta 22 (1) µm. Abdomen (Fig. 11A–B). Numbers of caudal spines on tergites II–VIII as: 20–32, 27; 18–36, 27; 23–37, 30; 17–35, 27; 23–37, 30; 19–30, 24; 19–34, 26. Length of longest caudal spine on tergites II–VIII as: 7–10, 8; 8–11, 9; 8–11, 10; 8–12, 10; 11–17, 13; 11–15, 13; 11–18, 14. Anal lobe (Fig. 11E–F) with three thorn-like macrosetae 41–47, 44; 33–55, 43 and 41–50, 47 µm long. Male genital sac overreaches anal lobe by 51–72 µm. Fourth instar larva (n = 6, if not otherwise stated). Total length 3.3 (1) mm. Head capsule 0.34–0.36 (2) mm long. Postmentum 135–139 (2) µm long. Thoracic segments brown, abdomen and head amber yellow. Head. Antenna as (Fig. 12A), segment lengths in µm: 28–32, 31; 13–15,15; 3–4, 3; 2– 3, 2; 2–3, 2. Blade longer than flagellum, 26–29, 28 µm long; apical style of second segment 4–5, 4 µm long. Seta I irregular, nearly palmate (Fig. 12B); other setae simple; labral lamella inconspicuous. Chaeta media serrate, chaetulae laterales simple, chaetulae basales bifid. Pecten epipharyngis with about 15 teeth. Mandible (Fig. 12D) 112–124, 117 µm long, with apical tooth and four inner teeth; seta subdentalis triangular; seta interna with 4 strongly serrated branches. Premandible (Fig. 12E), 48–59, 53 µm long. Mentum (Fig. 12C) 85–94, 89 µm wide; with irregular median tooth 24–28, 25 µm long and 5 lateral teeth. Ventromental plates small. Seta submenti 33–43, 41 µm long. Abdomen without distinct setae. Anterior parapods well developed, with strong apical claws. Posterior parapods about 206 (1) µm long, with 33–36 (2) µm long claws (Fig. 12F). Procerci absent. Longer anal tubules 90–94 (2) µm long, 34 (2) µm wide; shorter anal tubules 64–71 (2) µm long, 23–26 (2) µm wide. Remarks: This species is common and widespread in the São Paulo State, with specimens from Ribeirão Preto, São Carlos, Pedregulho and Ubatuba, and it has also been collected in Uberaba, in the Minas Gerais State. The larvae were collected in moss growing on tree trunks and branches of Bauhinia sp., Tabebuia sp., Mangifera indica, Eugenia jam- bolana and Caesalpinia sp., and in Ubatuba also in lichens growing on rocks.

FIGURE 12. Antillocladius musci new species, larva. A–antenna; B–S I setae; C–mentum; D– mandible; E–premandible; F–anal end.

Antillocladius pluspilalus Sæther

Antillocladius pluspilalus Sæther, 1982: 474, Fig. 6 A–E; 1984. Antillocladius pluspilalus Sæther; Spies and Reiss 1996: 75.

48 © 2004 Magnolia Press MENDES ET AL. Material examined: Type material as in Sæther (1982). MEXICO: Campeche Province, ZOOTAXA Reserva de la Biosfera Calakmul, Zona Arqueológica, 18°07'26.7''N, 89°48'56.7''W, 1 594 male, 20.ix.1997, 265 m a.s.l., light trap, A. Contreras-Ramos et al. (ZMBN). ECUADOR: Provincia Pichincha, Setor Palmeras, unión Rios Guajalito y Bricador, Bosque Protector Rio Guajalito, 00°14.925'N, 78°48.193'W, 1 male, 4–10.ii.2000, 1900 m a.s.l., light trap, F.M. Quesada (ZMBN). Diagnostic characters: The male imago can be separated from other members of the genus by having numerous apical setae on the wing membrane; setose squama; acrostichals in the middle of scutum, all more or less scalpellate; long virga; anal point long, tapering, with moderately strong lateral setae; and inferior volsella prominent with anterior digitiform projection equally long as and partly covering rounded posterior lobe. The pupa has thorn-like macrosetae, caudal spines on tergites, no pearl row on antennal sheath, and anal lobe not extended laterally. Remarks: This species was originally described from southeast USA by Sæther (1982) and the immatures by Sæther (1984). Later, Spies and Reiss (1996) recorded the species from Nicaragua. The present paper adds records from Ecuador and Mexico. The original description states that all acrostichals are simple, in the present material the posterior acrostichals are clearly scalpellate.

Antillocladius scalpellatus Wang and Sæther

Antillocladius scalpellatus Wang and Sæther, 1993: 228, Fig. 1. Antillocladius scalpellatus Wang and Sæther; Wang 2000: 634. Antillocladius scalpellatus Wang and Sæther; Tiunova et al. 2003: 66.

Material examined: Type material as in Wang and Sæther (1993). Diagnostic characters: The male imago can be separated from other members of the genus by having several apical setae on the wing membrane; setose squama; costa barely extended; acrostichals starting in front, anterior ones hair-like, posterior ones scalpellate; short virga; anal point long, tapering, with strong lateral setae; and inferior volsella prominent, rectangular. Remarks: The species is described in detail by Wang and Sæther (1993). The type locality is in the Province Guangdong in China (Wang & Sæther 1993). Tiunova et al. (2003) added records from the far east of Russia.

Antillocladius skartveiti Andersen and Contreras-Ramos

Antillocladius skartveiti Andersen and Contreras-Ramos, 1999: 150, Figs 1–6.

Material examined: Type material as in Andersen and Contreras-Ramos (1999).

ORTHOCLADIINAE © 2004 Magnolia Press 49 ZOOTAXA Diagnostic characters: The male imago can be separated from other members of the 594 genus by having several apical setae on the wing membrane; setose squama; costa moderately extended; acrostichals scalpellate, in the middle of scutum; no virga; anal point long, tapering, with strong lateral setae; and inferior volsella strongly set off rectangular to rounded. Remarks: The species is described in detail by Andersen and Contreras-Ramos (1999). It is known only from the type locality in Ecuador.

Antillocladius sooretama new species (Fig. 13A–E)

Type material: Holotype male, BRAZIL: Espírito Santo State, Parque de Sooretama, Cupido, ii–iii.1948, L. Travassos, Freitas & H. Travassos (FIOCruz, slide mounted in Euparal). Paratypes: 6 males as holotype (FIOCruz). Diagnostic characters: The male imago can be separated from other members of the genus by having bare wing membrane and wing veins; setose squama; costa barely extended; acrostichals starting in front, all scalpellate; no virga; anal point long, nearly parallel-sided, with blunt apex and weak lateral setae; and inferior volsella long, posterior directed, well set-off and apically bifid. Etymology: Named after the Parque Sooretama, the type locality of the species; the name is a noun in apposition. Male (n = 6). Total length 2.03 (1) mm. Wing length 1.20–1.32, 1.26 mm. Total length / wing length 1.68 (1). Wing length / length of profemur 2.35 (1). Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 1.35–1.44, 1.39, ultimate flagellomere 428–464, 450 µm long. Temporal setae 8–12 (2), including 1–3 inner verticals, 4 outer verticals and 3–5 postorbitals. Clypeus with 4–6 (2) setae. Tentorium, stipes and cibarial pump as in Fig. 13A; tentorium 124 (1) µm long, 13 (1) µm wide at sieve pore; stipes 128 (2) µm long. Palpomere lengths (2) in µm: 25, 55–59, 95–109, 91–92, 112–116. Third palpomere with 3 sensilla clavata subapically, longest 15 µm long. Thorax. Antepronotum with 3 setae. Dorsocentrals 6–10, 8; acrostichals 7–10, 9, all scalpellate starting close to antepronotum; prealars 3. Scutellum with 6–8, 7 setae. Wing (Fig. 13B). VR 1.30–1.41, 1.35. Costal extension 32–48, 42 µm long. Brachi- olum with 1 seta, other veins and cells bare. Squama with 4–5 setae. Legs. Spur of front tibia 33–48 (2) µm long; spurs of middle tibia 25–38, 30 µm and 21–25, 23 µm long; of hind tibia 44–51, 49 µm and 21–26, 24 µm long. Width at apex of front tibia 32–42 (2) µm; of middle tibia 32–39, 35 µm; of hind tibia 43–51, 46 µm. Hind tibial comb with 11–13, 12 setae; shortest seta 15–18, 16 µm long; longest seta 34–44, 39 µm long. Lengths (in µm) and proportions of legs as in table 9.

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

p1 536 544–580 232 128 88 56 48 0.43–0.68 4.10 4.65 3.1

p2 532– 548– 204– 104– 92–96, 56– 36– 0.37–0.41 4.01–4.46 4.84–5.29 3.1–3.2 544, 536 560, 550 230, 219 128, 110 95 60, 58 44, 41

p3 568– 604– 328–332 168– 124– 64– 44– 0.50– 3.47– 3.47– 3.0– 632, 602 656, 639 330 204, 186 140, 133 72, 67 52, 51 0.56, 0.54 3.54, 3.51 3.88, 3.65 3.5, 3.3

FIGURE 13. Antillocladius sooretama new species, male imago. A–cibarial pump, tentorium and stipes; B–wing; C–anal point and tergite IX and dorsal aspect of left gonocoxite; D–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; E–gonostylus.

ORTHOCLADIINAE © 2004 Magnolia Press 51 ZOOTAXA Hypopygium (Fig. 13C–E). Anal point 88–96, 93 µm long; 44–59, 52 µm wide at 594 base; with 13–17, 15 strong lateral setae, void of microtrichia except at base. Laterostern- ite IX with 3–4, 3 setae. Phallapodeme 77–80, 78 µm long. Transverse sternapodeme nearly straight, with rounded oral projections, 97 (1) µm long. Virga absent. Gonocoxite 179–197, 189 µm long; with long, well set-off, apically bifid inferior volsella. Gonostylus 94–96, 95 µm long; megaseta 13–15, 14 µm long, situated subapically. HR 1.91–207, 1.99; HV 2.10 (1).

Antillocladius ubatuba new species (Fig. 14A–E)

Type material: Holotype male, BRAZIL: São Paulo, Ubatuba, Praia das Toninhas, 09.ix.2000, sweep net, H.F. Mendes (MZUSP, slide mounted in Canada balsam). Paratypes: 14 males, as holotype (ZMBN); São Paulo State, São Carlos, 1 male, 9.xii.1999, H.F. Mendes (UFSCar); Rio de Janeiro State, Mangaratiba, Muriquí, 4 males, 19.i.1991, A. Serpa-Fo et al. (FIOCruz). VENEZUELA: Falcón, Rio Mitare near San Luis, 11°07.930'N, 69°39.184'W, 1 male, 7.vi.2001, 589 m a.s.l., R.W. Holzenthal, R. Blahnik, H. Paprocki, C. Cressa (MIZA). Diagnostic characters: The male imago can be separated from other members of the genus by having bare wing membrane; setose squama; costa moderately extended; acrostichals starting in front, all scalpellate; moderately long virga; anal point long, nearly parallel-sided near apex, with blunt apex and weak lateral setae; and inferior volsella long, posterior directed and digitiform. Etymology: Named after the type locality, Ubatuba in São Paulo State, Brazil; the name is a noun in apposition. Male (n = 8). Total length 2.62–2.74, 2.67 mm. Wing length 1.51–1.67, 1.60 mm. Total length / wing length 1.60–1.75, 1.67. Wing length / length of profemur 2.43–2.52, 2.46. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 1.57–1.69, 1.63; ultimate flagellomere 536–595, 573 µm long. Temporal setae 10–11, 8; including 4 inner verticals; 2–3, 3 outer verticals and 3–4, 4 postorbitals. Clypeus with 6–10, 8 setae. Tentorium, stipes and cibarial pump as in Fig. 14A, tentorium 136–139, 137 µm long, 13–15, 14 µm wide at sieve pore; stipes 112–116 (3) µm long. Pal- pomere lengths in µm: 25–30, 28; 45–56, 51; 121–139, 127; 116–131 (3); 134 (1). Third palpomere with 3–4, 3 sensilla clavata subapically, longest 18–20, 19 µm long. Thorax (Fig. 14B). Antepronotum with 2–4, 3 setae. Dorsocentrals 8–11, 9; acrostichals 14–19, 17, all scalpellate starting close to antepronotum; prealars 2–4, 3; supraalar 0–1, 0. Scutellum with 7–9, 8 setae.

FIGURE 14. Antillocladius ubatuba new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite and gonostylus; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect.

ORTHOCLADIINAE © 2004 Magnolia Press 53 ZOOTAXA Wing (Fig. 14C). VR 1.30–1.37, 1.33. Costal extension 67–88, 77 µm long. Brachi- 594 olum with 1–2, 1 setae; R1 with 0–1, 0 seta; other veins bare. Wing membrane bare. Squama with 6–10, 8 setae. Legs. Spur of front tibia 61–66, 63 µm long; spurs of middle tibia 30–33, 31 µm and 38–53, 43 µm long; spurs of hind tibia 28–33, 31 µm and 56–61, 58 µm long. Width at apex of front tibia 35–40, 37 µm; of middle tibia 35–38, 37 µm; of hind tibia 45–51, 48 µm. Hind tibial comb with 9–13, 12 setae; shortest seta 18–25, 21 µm long; longest seta 30–51, 38 µm long. Lengths (in µm) and proportions of legs as in Table 10.

TABLE 10. Lengths (in µm) and proportions of legs of Antillocladius ubatuba new species, (n=2– 8).

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

p1 595– 663– 497– 273– 175– 117– 58– 0.75– 2.72– 2.38– 2.1– 663, 624 761, 714 585, 544 322, 300 195, 180 136, 122 78, 66 0.77, 0.76 3.01, 2.84 2.79, 2.57 2.2, 2.2

p2 595– 634– 283– 146– 88–107, 64– 68 44–48 0.43– 4.16–4.37 4.41–4.47 3.2–3.5 683, 634 712, 666 312, 292 166, 153 101 0.45, 0.44

p3 653– 731– 409– 224– 166– 83–88 53–57 0.54– 3.32–3.69 3.23– 5.2–5.4 741, 699 809, 775 585, 485 253, 239 185, 175 0.72, 0.62 3.58, 3.46

Hypopygium (Fig. 14D–E). Anal point 76–88, 85 µm long; 38–43, 40 µm wide at base; with 13–19, 16 strong lateral setae; void of microtrichia except at base. Laterostern- ite IX with 7 setae. Phallapodeme 83–88, 87 µm long. Transverse sternapodeme slightly arcuate, with rounded oral projections; 109–114, 112 µm long. Virga 43–51, 45 µm long. Gonocoxite 181–192, 189 µm long, with well set–off inferior volsella with acute apex. Gonostylus 96–101, 99 µm long; megaseta 13–15, 14 µm long, situated slightly subapically. HR 1.87–1.95, 1.90; HV 2.65–2.76, 2.70.

Antillocladius venequatoriensis new species (Fig. 15A–F)

Type material: Holotype male, VENEZUELA: Aragua, Parque Nacional Henri Pittier, Ran- cho Grande, 10°21.047'N, 67°41.198'W, 16–18.ix.1999, about 1000 m a.s.l., sweep net & light trap, T. Andersen (MIZA, slide mounted in Canada balsam). Paratypes: 2 males, as holotype (ZMBN); Aragua, Parque Nacional Henri Pittier, Curva Regressiva on road to Choroni, 5 males, 23.ix.1999, 1200 m a.s.l., light trap, T. Andersen (ZMBN). ECUADOR: Provincia Pichincha, Setor Palmeras, unión Rios Guajalito y Bricador, Bosque Protector Rio Guajalito 00°14.925'N, 78°48.193'W, 1 male, 4–10.ii.2000, 1900 m a.s.l., light trap. F.M. Quesada (ZMBN).

FIGURE 15. Antillocladius venequatoriensis new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; F–gonostylus.

ORTHOCLADIINAE © 2004 Magnolia Press 55 ZOOTAXA Diagnostic characters: The male imago can be separated from other members of the 594 genus by having numerous setae in apical half of wing membrane; setose squama; costa weakly extended; acrostichals starting in front, all scalpellate; long virga; anal point long, nearly triangular with pointed apex and moderately strong lateral setae; and inferior volsella relatively small and rounded. Etymology: The name refers to the countries where the species has been taken, Vene- zuela and Ecuador. Male (n = 6). Total length 2.01–2.21, 2.15 mm. Wing length 1.23–1.44, 1.36 mm. Total length / wing length 1.54–1.63, 1.57. Wing length / length of profemur 2.30–2.42, 2.37. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 0.98–1.05, 1.01; ultimate flagellomere 526–572, 557 µm long. Temporal setae 8–11, 9; including 2–5, 3 inner verticals; 2 outer verticals and 3–4, 4 postorbitals. Clypeus with 10 setae. Tentorium, stipes and cibarial pump as in Fig. 15A; tentorium 125– 139, 131 µm long, 8–11, 9 µm wide at sieve pore; stipes 120–126, 122 µm long; 32–40, 36 µm wide. Palpomere lengths in µm: 22–25, 23; 35–38, 37; 90–96, 93; 85–97, 93 (4); 120– 135, 129 (4). Third palpomere with 3–4 sensilla clavata subapically; longest 10–15, 13 µm long. Thorax (Fig. 15B). Antepronotum with 2–3, 3 setae. Dorsocentrals 9–17, 13; acrostichals 9–14, 12, all scalpellate starting close to antepronotum; prealars 4–6, 5. Scutellum with 7–10, 8 setae. Wing (Fig. 15C). VR 1.37–1.61, 1.48. Costal extension 40–68, 54 µm long. Brachi-

olum with 1 seta; R with 11–15, 13 setae; R1 with 11–14, 13; R4+5 with 20–27, 23; M with

4–7, 6; M1+2 with 40–56, 46; M3+4 with 16–26, 21; Cu1 with 9–12, 10; and An with 10–15,

12 setae. Wing membrane with 122–163, 149 setae in cell r4+5; 81–94, 87 setae in m1+2;

39–43, 41 setae in m3+4; and 15–25, 21 in cu and an together. Squama with 5–7, 6 setae. Legs. Spur of front tibia 45–52, 50 µm long; spurs of middle tibia 30–33, 31 µm and 29–33, 30 µm long; spurs of hind tibia 28 µm and 47–54, 51 µm long. Width at apex of front tibia 35 µm; of middle tibia 30–33, 32 µm; of hind tibia 38–40, 39 µm. Hind tibial comb with 12 setae; shortest seta 18–25, 21 µm long; longest seta 27–38, 33 µm long. Lengths (in µm) and proportions of legs as in Table 11.

TABLE 11. Lengths (in µm) and proportions of legs of Antillocladius venequatoriensis new species.

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

p1 526– 568– 508– 260– 166– 60–112, 39– 0.86– 2.62– 2.16– 2.9–3.9, 600, 564 668, 624 572, 539 272, 269 184, 177 107 56, 41 0.91, 0.87 2.91, 2.87 2.22, 2.18 3.3

p2 540– 532– 260– 132– 104– 49–92, 40– 0.47– 3.95– 3.96– 3.4–4.3, 644, 565 636, 556 300, 273 156, 136 125, 120 87 52, 48 0.53, 0.51 4.27, 4.07 4.12, 4.05 3.8

p3 624– 673– 388– 212– 156– 78–92, 39– 0.56– 3.32– 3.38– 4.0–4.7, 664, 645 744, 683 416, 390 234, 224 172, 161 81 56, 47 0.58, 0.57 3.35, 3.34 3.42, 3.39 4.4

56 © 2004 Magnolia Press MENDES ET AL. Hypopygium (Fig. 15D–F). Anal point 58–63, 61 µm long; 40–44, 42 µm wide at ZOOTAXA base; with 15–27, 24 strong lateral setae; void of microtrichia except at base. Laterostern- 594 ite IX with 3–6, 5 setae. Phallapodeme 83–97, 88 µm long. Transverse sternapodeme arcuate, with rounded oral projections; 96–113, 104 µm long. Virga 58–61, 59 µm long. Gonocoxite 163–168, 165 µm long; with long, low inferior volsella. Gonostylus 94–123, 101 µm long; megaseta 7–10, 8 µm long. HR 1.37–1.73, 1.62; HV 1.86–2.27, 2.15.

Antillocladius zempoalensis new species (Fig. 16A–F)

Type material: Holotype male, MEXICO: Morelos province, Parque Nacional Lagunas de Zempoala, 23.ix.1998, about 3000 m a.s.l., sweep net, T. Andersen, A. Contreras-Ramos & R. Barba-Alvarez (ZMBN Type no. 400, slide mounted in Canada balsam). Diagnostic characters: The male imago can be separated from other members of the genus by having bare wing membrane, veins and squama; costa strongly extended; acrostichals starting in front, anterior one hair-like, posterior ones scalpellate; no virga; anal point moderately long, triangular with pointed apex and strong lateral setae; and inferior volsella well developed, set off and rounded. Etymology: Named after Parque Nacional Lagunas de Zempoala in central Mexico, where the species was taken. Male (n = 1). Total length 2.68 mm. Wing length 1.79 mm. Total length / wing length 1.50. Wing length / length of profemur 2.55. Coloration brown. Head. (Fig. 16A) AR 0.71, ultimate flagellomere 384 µm. Temporal setae 6; including 2 inner verticals, 2 outer verticals and 2 postorbitals. Clypeus with 5 setae. Tentorium, stipes and cibarial pump as in Fig. 16B; tentorium 121 µm long, 17 µm wide at sieve pore; stipes 97 µm long, 30 µm wide. Left palp with extended apex on third palpomere, palpomere lengths of left palp in µm: 29, 48, 69, 72, 83; right palp broken, but third palpomere apparently normal, 88 µm long. Third palpomere with 2 sensilla clavata apically to subapically, 11 µm long. Thorax (Fig. 16C). Antepronotum with 3 setae. Dorsocentrals 6; acrostichals 11, of which 2 anterior hair-like and 9 posterior scalpellate, beginning at one fourth from antepronotum; prealars 2. Scutellum with 4 setae. Wing (Fig. 16D). VR 1.33. Costal extension 91 µm long. Brachiolum with 1 seta, other veins and cells bare. Squama bare. Legs. Spur of front tibia 54 µm long, spurs of middle tibia 26 µm and 25 µm long, spurs of hind tibia 43 µm and 23 µm long. Width at apex of front tibia 33 µm, of middle tibia 33 µm, of hind tibia 37 µm. Hind tibial comb well developed, not measurable. Lengths (in µm) and proportions of legs as in Table 12.

FIGURE 16. Antillocladius zempoalensis new species, male imago. A–head; B–cibarial pump, tentorium and stipes; C–thorax; D–wing; E–anal point and tergite IX and dorsal aspect of left gonocoxite; F–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect.

58 © 2004 Magnolia Press MENDES ET AL. TABLE 12. Lengths (in µm) and proportions of legs of Antillocladius zempoalensis new species. ZOOTAXA 594 fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR p1 702 792 514 261 180 114 57 0.65 3.00 2.91 3.3 p2 678 727 310 172 114 65 41 0.43 4.38 4.53 4.9 p3 735 768 417 221 155 74 49 0.54 3.85 3.61 5.8

Hypopygium (Fig. 16E–F). Anal point 50 µm long, 19 µm wide at base, with 17 strong lateral setae, void of microtrichia except at base. Laterosternite IX with 5 setae. Phallapodeme 79 µm long. Transverse sternapodeme arcuate with rounded oral projections, 35 µm long. Virga absent. Gonocoxite 156 µm long, with well developed, set-off inferior volsella. Gonostylus 72 µm long, megaseta 12 µm long. HR 2.17, HV 3.72.

Antillocladius zhengi Wang and Sæther

Antillocladius zhengi Wang and Sæther, 1993: 229, Fig. 2 A–B.

Material examined: Holotype as in Wang and Sæther (1993). THAILAND: Chiang Mai Province, Nankoi waterfall near Samoeng, 1 male, 10.iv.1991, sweep net, T. Andersen (ZMBN). Diagnostic characters: The male imago can be separated from other members of the genus by having a few apical setae on the wing membrane; bare veins; setose squama; costa strongly extended; acrostichals starting in front, anterior one hair-like to lanceolate, posterior ones scalpellate; moderately long virga; anal point moderately long, triangular with pointed apex and strong lateral setae; and inferior volsella well developed, rectangular, with anterior somewhat pointed projection of about the same length as the posterior rounded part. Remarks: The species was described in detail by Wang and Sæther (1993) based on a single male from southeast China. The present paper adds a new record from northern Thailand. The specimen from Thailand varies a little in the shape of gonostylus and the number of lateral setae on the anal point is higher. Most measurements however, are close to the measurements given by Wang and Sæther (1993).

Compterosmittia Sæther

Compterosmittia Sæther, 1981: 20; Cranston et al. (1989: 186); Sæther (1982: 491); Cranston and Oliver (1988: 429); Cranston and Kitching (1995: 229).

Other included species Compterosmittia aberrans new species; C. berui new species; C. claggi (Tokunaga, 1964: 506, as Metriocnemus, possible synonym of C. nerius); C. croizati new species; C. nerius (Curran, 1930: 34, as Camptocladius, syn. clavigera Sæther, 1982); C. pectinata (Freeman, 1961: 662, as Gymnometriocnemus); C. pittieri new species; C. tuberculifera (Tokunaga, 1964; 530, as Smittia); C. virga Wang, 1998.

Diagnostic characters The imagines can be separated from other orthoclad genera by having scalpellate acrostichals in the middle of scutum, or in C. croizati new species and C. virga Wang starting closer to antepronotum, combined with bare squama, very long costal extension, and often absence of virga and dentate megaseta. All known females and several males have setae apical on the wing membrane. The pupa is not separable from Limnophyes Eaton. The only differences between the larvae of Limnophyes and Compterosmittia consist in the mandible which has 4 inner teeth in Compterosmittia, 3 in Limnophyes; and in that the supra-anal seta is 1/3 the length of the anal setae in Compterosmittia, much longer in Lim- nophyes.

Generic diagnosis Imago Small species, wing length 0.7–1.3 mm. Female antenna with 5 flagellomeres. Male antenna with 10–13 flagellomeres, groove beginning at flagellomere 3, sensilla chaetica on flagellomeres 2, 3 and 13. Male antennal ratio 0.5–1.0. Head. Eye naked, with or without minute pubescence between ommatids, without dorsomedial extension. Temporal setae divided into weak inner verticals, stronger outer verticals, postorbitals absent or few. Third palpomere with about 1–3 sensilla clavata apically, longest 8–17 µm long. Thorax. Antepronotum well developed, lobes meeting medially along short suture. Acrostichals, short, all scalpellate in the middle of scutum or starting about 1/3 of scutum length from antepronotum; prealars uniserial; supraalar 0–1; scutellars uniserial. Wing. Anal lobe protruding or absent. Costa strongly extended. R2+3 running and end-

ing midway between R1 and R4+5; R4+5 ending opposite or slightly distal to M3+4; FCu far

distal to RM; Cu1 nearly straight to distinctly sinuous; An ending proximal to FCu. Veins

bare or with setae on R, R1, R4+5, and/or M1+2,. Membrane with apical setae in female;

male without setae, with few weak setae apically in cell r4+5, or with several setae apically

in cells r4+5, and m1+2. Squama bare.

60 © 2004 Magnolia Press MENDES ET AL. Legs. Pseudospurs, sensilla chaetica and pulvilli lacking. Comb and tibial spurs nor- ZOOTAXA mal. 594 Abdomen. Tergites with irregularly arranged setae. Hypopygium. Anal point moderately long, sharply to bluntly pointed, with mostly strong lateral setae, with microtrichia only at base or nearly to apex. Phallapodeme and aedeagal lobe well developed. Anterior margin of sternapodeme arcuate, oral projections moderately to strongly developed. Virga absent or, in C. virga, consisting of 16–18 spines. Inferior volsella simple, rounded and projecting or with anterior microtrichia-free, digitiform anterior projection and low to prominent posterior lobe. Gonostylus without heel; crista dorsalis absent. Megaseta often dentate. Female genitalia. Gonocoxapodemes curved and meeting anterior of vagina. Gono- coxite relatively well developed, with several long setae and a few short setae. Tergite IX divided. Gonapophysis IX with large ventrolateral lobe covering apodeme lobe and apex of narrow, line-like dorsomesal lobe. Labia small. Cerci normal. Seminal capsules long, with triangular, pale neck. Spermathecal ducts with large loop and common opening. Immatures The immatures of one species are described by Cranston and Kitching (1995). Except for the characters mentioned in the diagnostic characters, the diagnoses for both pupa and larva are identical to those given for Limnophyes by Sæther (1990: 9).

Key to the males of Compterosmittia Sæther

1. Virga consisting of 16–18 spines; China...... C. virga Wang — Virga absent...... 2 2. Inferior volsella a rounded strongly projecting simple lobe ...... 3 — Inferior volsella with an anterior, dorsal, digitiform, bare lobe and a posterior, ventral, more rounded, more or less well developed lobe...... 6 3. Anal point well developed with several strong lateral setae; Australia ...... C. pectinata (Freeman) — Anal point short with few and weak setae...... 4 4. Megaseta not dentate (Fig. 19D); Venezuela, Brazil...... C. croizati new species — Megaseta dentate ...... 5

5. Anal lobe present, Cu1 moderately sinuous; Micronesia, Bonin Is., Guam, Belau...... C. tuberculifera (Tokunaga)

— Anal lobe reduced, Cu1 strongly sinuous ...... C. nerius (Curran) New York, South Carolina; and C. claggi (Tokunaga) Bonin Is., Belau, Micronesia 6. Megaseta strongly dentate; St. Vincent, St. Kitts ...... C. dentispina Sæther — Megaseta apparently simple ...... 7 7. Antennae with 10 flagellomeres; inferior volsella with prominent elongate ventral part

ORTHOCLADIINAE © 2004 Magnolia Press 61 ZOOTAXA (Fig. 17F); Costa Rica ...... C. aberrans new species 594 — Antennae with 13 flagellomeres; inferior volsella either with reduced elongate ventral part or with posterior ventral part short and rounded...... 8 8. Anal point narrow, sharply pointed; posterior ventral part of inferior volsella low and elongate; no supraalar (Fig. 18D); Brazil...... C. berui new species — Anal point moderately long with blunt apex; ventral part of inferior volsella prominent, short and rounded; supraalar present (Fig. 20D); Venezuela ...... C. pittieri new species

Compterosmittia aberrans new species (Fig. 17A–F)

Type material: Holotype male, COSTA RICA: Heredia Province, La Selva Biological Sta- tion, 10º26'N, 83º59'W, 3.iv.1993, Malaise trap, O.A. Sæther (ZMBN Type no. 401, slide mounted in Canada balsam). Diagnostic characters: The male imago can be separated from other members of the genus by having 10 flagellomeres; several apical setae on the wing membrane; inferior volsella with prominent low posterior lobe; and megaseta not dentate. Etymology: From Latin, aberrans, go astray, abnormal, referring to the ten-segmented antenna. Male (n = 1). Total length 1.47 mm. Wing length 0.90 mm. Total length / wing length 1.36. Wing length / length of profemur 2.68. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head (Fig. 17A). Antenna with 10 flagellomeres, nine and ten partly fused, AR 0.97, ultimate flagellomere 260 µm long. Temporal setae 8, including 3 inner verticals, 2 outer verticals and 3 postorbitals. Clypeus with 9 setae. Tentorium, stipes and cibarial pump as in Fig. 17B; tentorium 72 µm long, 14 µm wide at sieve pore; stipes 48 µm long, 11 µm wide. Palp broken, palpomere lengths (1–3) in µm: 12, 26, 48. Third palpomere apparently with 2 sensilla clavata subapically, longest 8 µm long. Thorax (Fig. 17C). Antepronotum with 4 setae. Dorsocentrals 8; acrostichals 6, all scalpellate, in the middle of scutum; prealars 3. Scutellum with 6 setae.

Wing (Fig. 17D). VR 1.36. Costal extension 69 µm long. Brachiolum with 1 seta, M1+2

with 14 setae, other veins bare. Wing membrane with 23 setae in cell r4+5, 5 setae in m1+2. Squama bare. Legs. Spur of front tibia 28 µm long, spurs of middle tibia 19 µm and 28 µm long, of hind tibia 22 µm and 35 µm long. Width at apex of front tibia 23 µm, of middle tibia 23 µm, of hind tibia 26 µm. Hind tibial comb with 4 setae, shortest seta 14 µm long, longest seta 18 µm long. Lengths (in µm) and proportions of legs as in Table 13.

FIGURE 17. Compterosmittia aberrans new species, male imago. A–head; B–cibarial pump, tentorium and stipes; C–thorax; D–wing; E–anal point and tergite IX and dorsal aspect of left gonocoxite and gonostylus; F–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect.

p1 335 366 343 180 98 49 33 0.93 2.91 2.05 2.59

p2 393 327 237 90 57 33 25 0.75 4.68 3.03 3.88

p3 425 393 253 114 98 41 33 0.65 3.74 3.23 6.40

Hypopygium (Fig. 17E–F). Anal point 23 µm long, 11 µm wide at base; with 11 strong, lateral setae, void of microtrichia except at base. Laterosternite IX with 3 setae. Phallapodeme 50 µm long. Transverse sternapodeme arcuate, with rounded oral projections, 55 µm long. Virga absent. Gonocoxite 102 µm long, with long, low inferior volsella, with rounded microtrichia-free projection orally. Gonostylus 59 µm long, megaseta 6 µm long. HR 1.72, HV 2.49.

Compterosmittia berui new species (Fig. 18A–E)

Type material: Holotype male, BRAZIL: São Paulo State, Salesópolis, Estação Biológica Boracéia, córrego Venerando, 12.xii.2001, light trap, C. Froehlich et al. (MZUSP, slide mounted in Euparal). Paratypes: Santa Catarina State, Nova Teotônia, 27°11'S, 52°23'W, 1 male, xii.1970, F. Plaumann (MZUSP); Santa Catarina State, São Bento do Sul, 26°19'25.6''S, 48°18'26.5''W, 4 males, 13–16.x.2001, Malaise trap, M.V. Yamada (1 UFS- Car, 3 ZMBN). Diagnostic characters: The male imago can be separated from other members of the genus by having wing membrane with 0–8 apical setae; thin anal point tapering to point; inferior volsella with anterior knob-like projection and barely set off posterior extension; and megaseta not dentate. Etymology: From the Indian language Tupy, beruì, meaning small midge; the name is a noun in apposition. Male (n = 5). Total length 1.64–1.86, 1.71 mm. Wing length 1.04–1.24, 1.13 mm. Total length / wing length 1.42–1.60, 1.50. Wing length / length of profemur 2.55–2.67, 2.60. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 0.69–0.78, 0.73; ultimate flagellomere 236–304, 271 µm. Temporal setae 6–9, 8 including 2–4, 3 inner verticals, 2–4, 3 outer verticals and 1–2, 1 postorbitals. Clypeus with 6–11, 8 setae. Tentorium, stipes and cibarial pump as in Fig. 18A; tentorium 103–110, 106 µm long, 7–8, 8 µm wide at sieve pore; stipes 69–76, 73 µm long. Pal- pomere lengths in µm: 14–15, 14; 23–25, 24; 50–54, 52; 44–55 (3); 47–55 (3); 75–80 (3). Third palpomere with 2–4, 3 sensilla clavata subapically; longest 13–14, 14 µm long.

FIGURE 18. Compterosmittia berui new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite and gonostylus; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect.

ORTHOCLADIINAE © 2004 Magnolia Press 65 ZOOTAXA Thorax (Fig. 18B). Antepronotum with 2–3, 3 setae. Dorsocentrals 5–6, 5; acrostichals 594 6–7, 6, all scalpellate starting at one third from antepronotum; prealars 2. Scutellum with 4 setae. Wing (Fig. 18C). VR 1.48–1.56, 1.53. Costal extension 103–134, 115 µm long. Bra- chiolum with 1 seta, other veins bare; cell r4+5 with 0–8, 4 setae, other cells bare. Squama bare. Legs. Spur of front tibia 25–30 (3) µm long, spurs of middle tibia 25–32, 27 µm and 16–21, 19 µm long; of hind tibia 29–37, 34 µm and 17–21, 19 µm long. Width at apex of front tibia 22–26, 23 µm; of middle tibia 22–25, 24 µm; of hind tibia 25–29, 27 µm. Hind tibial comb with 5–6 (3) setae, shortest seta 11–17 (3) µm long, longest seta 22–26 (3) µm long. Lengths (in µm) and proportions of legs as in Table 14.

TABLE 14. Lengths (in µm) and proportions of legs of Compterosmittia berui new species, (n=1–3).

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

p1 388–440 436–472 408 412 192–216 96–108 48–60 36–44 0.87–0.94 3.09–3.31 2.02–2.21 3.1–3.3

p2 460 400 248 108 60 36 28 0.62 4.78 3.47 3.5

p3 448–488 452–480 220–284 104–124 80–104 40–44 28–32 0.50–0.61 3.91–4.54 3.31–3.41 4.5–4.8

Hypopygium (Fig. 18D–E). Anal point 41–44, 42 µm long; 32–37, 35 µm wide at base; with 7–12, 10 strong lateral setae, void of microtrichia except at base. Laterosternite IX with 4 setae. Phallapodeme 43–62, 52 µm long. Transverse sternapodeme nearly straight; 50–62, 58 µm long. Virga absent. Gonocoxite 100–153, 124 µm long; with long, low inferior volsella, with rounded projection orally. Gonostylus 62–76, 68 µm long; megaseta 5–8, 7 µm long. HR 1.60–1.95, 1.81; HV 2.16–2.67, 2.38.

Compterosmittia claggi (Tokunaga)

Metriocnemus claggi Tokunaga, 1964: 506. Compterosmittia claggi (Tokunaga); Cranston and Martin 1989: 259.

Diagnostic characters: The male imago cannot be separated from C. nerius (see below). Remarks: The type material in the Bishop Museum, Honolulu, has not been examined, but according to Cranston and Kitching (1995) is of too poor quality for species separa- tion. However, C. claggi appears to be identical to C. nerius. It is known from Bonin Is., Belau and Micronesia.

Type material: Holotype male, VENEZUELA: Aragua, Parque Nacional Henri Pittier, Ran- cho Grande, 10°21.047'N, 67°41.198'W, 16–18.ix.l1999, about 1000 m a.s.l., sweep net & light trap, T. Andersen (MIZA, slide mounted in Canada balsam). Paratypes: Brazil: Santa Catarina State, São Bento do Sul, 26°19'25.6''S, 48°18'26.5''W, 2 males, 13–16.x.2001, 660 m a.s.l., Malaise trap, M.V. Yamada (1 MZUSP and 1 ZMBN). Diagnostic characters: The male imago differs from C. nerius only in having megaseta not dentate. Etymology: Named in honour of the Venezuelan biogeographer L. Croizat. Male (n = 1–3). Total length 1.68–1.85 mm. Wing length 1.05–1.14 mm. Total length / wing length 1.57–1.66. Wing length / length of profemur 2.46–2.62. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 0.66–0.73, ultimate flagellomere 224–280 µm. Temporal setae 5–9, including 1–2 inner verticals, 1–2 outer verticals and 2–4 postorbitals. Clypeus with 12 setae. Tentorium, stipes and cibarial pump as in Fig. 19A; tentorium 110–112 µm long, 7–8 µm wide at sieve pore; stipes 97–98 µm long, 18 µm wide. Palpomere lengths in µm: 15–22, 26–32, 55–62, 48–55, 69. Third palpomere with 2–3 sensilla clavata subapically, 14–17 µm long. Thorax (Fig. 19B). Antepronotum 0–1 seta. Dorsocentrals 7–8; acrostichals 9–11, all scalpellate, starting at one third from antepronotum; prealars 3–4. Scutellum with 4 setae. Wing (Fig. 19C). VR 1.43–1.53. Costal extension 148–168 µm long. Brachiolum with

1 seta, R with 3 setae, R4+5 with 0–4 setae, other veins bare, wing membrane bare. Cu slightly sinuous. Squama bare. Legs. Spur of front tibia 32–36 µm long, spurs of middle tibia 25 µm and 17–21 µm long, spurs of hind tibia 39–41 µm and 18–19 µm long. Width at apex of front tibia 25–32 µm, of middle tibia 21–29 µm, of hind tibia 30–36 µm. Hind tibial comb with 10–12 setae, shortest seta 18–21 µm long, longest seta 36–39 µm long. Lengths (in µm) and proportions of legs as in Table 15.

TABLE 15. Lengths (in µm) and proportions of legs of Compterosmittia croizati new species.

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR p1 428–436 496–492 396 176 112 56 40 0.80 3.44 2.34 2.8 p2 444–464 448–436 280–324 84–88 60–72 32–40 28–32 0.64–0.72 5.09–5.94 2.81–3.14 3.6 p3 440–480 520–528 288–340 123–136 112–116 52–56 40–43 0.59–0.63 3.53–3.96 2.96–3.22 3.6

FIGURE 19. Compterosmittia croizati new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite and gonostylus; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect.

68 © 2004 Magnolia Press MENDES ET AL. Hypopygium (Fig. 19D–E). Anal point 29–34 µm long, 25–27 µm wide at base, with ZOOTAXA 9–16 strong lateral setae, void of microtrichia except at base. Laterosternite IX with 3–4 594 setae. Phallapodeme 59–62 µm long. Transverse sternapodeme nearly straight, with rounded oral projections, 55–62 µm long. Virga absent. Gonocoxite 135–139 µm long, with long, low inferior volsella. Gonostylus 62–65 µm long, megaseta 14–16 µm long. HR 2.13–2.24, HV 2.71–2.98. Remarks: The only significant difference between C. croizati and C. nerius consists in the non-dentate megaseta. This could perhaps be intraspecific variation and C. croizati thus a synonym of the apparently widespread C. nerius.

Compterosmittia dentispina Sæther

Compterosmittia dentispina Sæther, 1981: 20.

Diagnostic characters: The male imago can be separated from other members of the genus by having inferior volsella with an anterior, dorsal, digitiform, bare lobe and a posterior, ventral, more rounded, lobe; presence of supraalar; and megaseta strongly dentate. Remarks: The species is known from St. Vincent and St. Kitts.

Compterosmittia nerius (Curran)

Camptocladius nerius Curran, 1930: 34. Compterosmittia clavigera Sæther, 1982: 491. Compterosmittia nerius (Curran); Cranston and Oliver 1988: 429. ? Metriocnemus claggi Tokunaga, 1964: 506.

Diagnostic characters: The male imago can be separated from other members of the genus by having inferior volsella a rounded strongly projecting simple lobe; presence of supraalar; anal lobe reduced, Cu1 strongly sinuous; anal point weak with few setae; and megaseta dentate. Remarks: The species is known from New York, North and South Carolina and perhaps Bonin Is., Belau and Micronesia (Cranston & Oliver 1988; Caldwell et al. 1997).

Compterosmittia pectinata (Freeman)

Gymnometriocnemus pectinatus Freeman, 1961: 662. Compterosmittia pectinata (Freeman); Cranston and Martin 1989: 259. ? Compterosmittia pectinata (Freeman); Cranston and Kitching 1995: 231.

ORTHOCLADIINAE © 2004 Magnolia Press 69 ZOOTAXA Diagnostic characters: The male imago can be separated from other members of the genus 594 by having inferior volsella a rounded strongly projecting simple lobe; anal point well developed with several strong lateral setae; and megaseta dentate. Remarks: The immatures described from phytotelmata in Australia by Cranston and Kitching (1995) may belong to C. pectinata, as this is the only described Australian species. However, immatures belonging to the genus are widespread in a variety of Australian and Oriental phytotelmata (Cranston & Kitching 1995).

Compterosmittia pittieri new species (Fig. 20A–F)

Type material: Holotype male, VENEZUELA: Aragua, Parque Nacional Henri Pittier, Ran- cho Grande, 10°21.047'N, 67°41.198'W, 16–18.ix.1999, about 1000 m a.s.l., sweep net & light trap, T. Andersen (MIZA, slide mounted in Canada balsam). Diagnostic characters: The male imago can be separated from other members of the genus by having a few apical setae on the wing membrane; anal point short, tapering, with weak lateral setae; inferior volsella relatively low with anterior bare, digitiform projection; and megaseta not dentate. Etymology: Named in honour of the botanist Henry Pittier, who was instrumental in setting up the National Park where the material was collected. Male (n=1). Total length 1.63 mm. Wing length 0.93 mm. Total length / wing length 1.77. Colora- tion brown, thorax dark brown, tarsi of all legs light brown. Head. AR 0.56, ultimate flagellomere 192 µm. Temporal setae 6, including 3 inner verticals, 1 outer vertical and 2 postorbitals. Clypeus with 10 setae. Tentorium, stipes and cibarial pump as in Fig. 20A; tentorium 92 µm long, 7 µm wide at sieve pore; stipes 59 µm long. Palpomere lengths in µm: 12, 26, 48, 51, 72. Third palpomere with 3 sensilla clavata subapically, longest 12 µm long. Thorax (Fig. 20B). Antepronotum with 2 setae. Dorsocentrals 8; acrostichals 7, all scalpellate in the middle of scutum; prealars 2; supraalar 1. Scutellum with 2 setae. Wing (Fig. 20C). VR 1.45. Costal extension 105 µm long. Brachiolum with 1 seta,

M1+2 with 14 setae, R with 3 setae, R4+5 with 1 seta apically, other veins bare. Wing mem-

brane with 21 setae in cell r4+5, 6 setae in m1+2. Squama with 5 setae. Legs. Front legs lost. Spurs of middle tibia 25 µm and 21 µm long, of hind tibia 34 µm and 21 µm long. Width at apex of middle tibia 18 µm, of hind tibia 28 µm. Hind tibial comb with 7 setae, shortest seta 17 µm long, longest seta 23 µm long. Lengths (in µm) and proportions of legs as in Table 16.

FIGURE 20. Compterosmittia pittieri new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite; E– hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; F– gonostylus.

p1 ------

p2 380452------

p3 452 456 340 116 104 40 28 0.75 4.34 2.67 4.8

Hypopygium (Fig. 20D–F). Anal point 29 µm long, 34 µm wide at base, with 7 strong lateral setae, void of microtrichia except at base. Laterosternite IX with 7 setae. Phallapo- deme 41 µm long. Transverse sternapodeme nearly straight with rounded oral projections, 61 µm long. Virga absent. Gonocoxite 131 µm long, with long, low inferior volsella, with rounded projection orally. Gonostylus 65 µm long, megaseta 4 µm long. HR 2.02, HV 2.51.

Compterosmittia tuberculifera (Tokunaga)

Smittia tuberculifera Tokunaga, 1964: 530. Compterosmittia tuberculifera (Tokunaga); Cranston and Martin 1989: 259.

Diagnostic characters: The male imago can be separated from other members of the genus by having inferior volsella a rounded strongly projecting simple lobe; anal point short with few and weak setae; wing with anal lobe, Cu1 moderately sinuous and megaseta dentate. Remarks: The species is known from Micronesia, Bonin Is., Guam and Belau.

Compterosmittia virga Wang

Compterosmittia virga Wang, 1998: 95.

Diagnostic characters: The male imago can be separated from other members of the genus by having a virga consisting of 16–18 spines. Remarks: The species is known from a subtropical mountain area in southern China.

Litocladius new genus

Type species Litocladius mateusi new species.

72 © 2004 Magnolia Press MENDES ET AL. Diagnostic characters ZOOTAXA The imagines can be separated from other orthoclad genera by having scalpellate 594 acrostichals, long virga with lateral lamellae, short costal extension, setae on squama and long anal point with lateral setae. The male may have a few setae apically on the wing, while the female has numerous setae on the wing membrane. The pupa can be separated from that of Gymnometriocnemus Goetghebuer by lacking frontal setae and by having all thoracic setae reduced. The larva is not known.

Etymology From Greek, litos, meaning plain, simple, referring both to the reduced pupa and the “normal “ inferior volsella; and – cladius, a common ending among orthoclads.

Generic diagnosis Imago Small to medium sized species, wing length about 1.4 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 antennal ratio about 1.5. Head. Eye naked, without dorsomedial extension. Temporal setae divided into weak inner verticals, stronger outer verticals, postorbitals absent or few. Third palpomere with about 3 sensilla clavata apically, longest about 20 µm long. Thorax. Antepronotum well developed, lobes meeting medially along short suture. Acrostichals short, anterior few simple, posterior scalpellate, beginning close to antepronotum, prealars uniserial; supraalar 0–1; scutellars uniserial.

Wing. Anal lobe developed. Costa at most moderately extended. R2+3 running and ending midway between R1 and R4+5; R4+5 ending slightly distal to M3+4; FCu far distal to

RM; Cu1 straight; An ending proximal to FCu. Veins bare in male. Membrane with apical setae in female; male with or without few weak setae apically in cell r4+5, Squama setose. Legs. Pseudospurs, sensilla chaetica and pulvilli lacking. Comb and tibial spurs normal. Abdomen. Tergites with irregularly arranged setae. Hypopygium. Anal point long, pointed, with few lateral setae, without microtrichia near apex. Phallapodeme and aedeagal lobe well developed. Anterior margin of sternapodeme weakly arcuate, oral projections moderately developed. Virga consisting of 2 long, strong spines with lateral lamellae. Inferior volsella with anterior dorsal triangular part and a more rounded ventral, posterior part. Gonostylus without heel; crista dorsalis rounded and well developed. Megaseta normal. Female genitalia. Gonocoxapodemes curved and meeting anterior of vagina. Gono- coxite well developed, with a few long and some short setae. Tergite IX divided. Gonapo-

ORTHOCLADIINAE © 2004 Magnolia Press 73 ZOOTAXA physis IX apparently with large ventrolateral lobe covering apodeme lobe, and narrow, 594 line-like dorsomesal lobe. Labia relatively large, pointed. Cerci small. Seminal capsules small to medium-sized, circular, with triangular neck. Spermathecal ducts not observable on available specimen. Pupa Small, total length 2.3 mm. Cephalothorax. Frontal setae absent. Frontal apotome smooth. Antennal sheath with basal pearl row. Ocular field with 2 reduced postorbitals. Thoracic horn absent. Three weak precorneals situated in narrow triangle. Two weak antepronotals. Dorsocentrals very small, in two groups of two. Thorax and wing sheath smooth. Abdomen. Tergite I–VIII covered with coarse shagreen except along anterior margin; tergite IX with strong shagreen except along posterior margin. Sternites I–IV bare; sternite V with median shagreen; sternites VI–VII with slightly more extensive median and lateral shagreen; sternite VIII with extensive shagreen; sternite IX with anterolateral shagreen. Tergite II without posterior hooklets. Tergites without posterior row of weak spines. Con- junctives bare. Pedes spurii A and B absent. Apophyses weak, nearly straight. Segment II –VIII with 2 weak, hair-like L setae. Anal lobe without macrosetae; male genital sac overreaching anal lobe.

Litocladius mateusi new species (Figs 21A–F, 22A–H, 23A–E)

Type material: Holotype male with pupal exuviae, BRAZIL, São Paulo State, Pedregulho, Furna São Pedro (Cerradão), 7.vii.2000, S. Mateus (MZUSP, slide mounted in Euparal). Paratypes: 1 male, 1 female, as holotype (ZMBN). Etymology: Named after Sidnei Mateus, who collected the type material. Male (n = 1–2). Total length 2.31–2.35 mm. Wing length 1.36–1.38 mm. Total length / wing length 1.70–1.69. Wing length / length of profemur 2.46–2.61. Coloration brown, thorax dark brown, tarsi of all legs light brown. Head. AR 1.46, ultimate flagellomere 508 µm. Temporal setae 6–8, including 1–2 inner verticals, 2 outer verticals and 3–4 postorbitals. Clypeus with 4 setae. Tentorium, stipes and cibarial pump as in Fig. 21A; tentorium 139 µm long, 15 µm wide at sieve pore; stipes not measurable. Palpomere lengths in µm: 25, 40–45, 81–83, 73–81, 91. Thorax (Fig. 21B). Antepronotum with 4–5 setae. Dorsocentrals 7; acrostichals 7–9, of which anterior 3–4 simple, posterior scalpellate, starting close to antepronotum; prealars 3; supraalar 0–1. Scutellum with 4–6 setae. Wing (Fig. 21C). VR 1.45–1.47. Costal extension 50–66 µm long. Brachiolum with 1

seta, other veins bare. Wing membrane with 0–1 setae in cell r4+5. Squama with 4–6 setae.

FIGURE 21. Litocladius mateusi new genus, new species, male imago. A–cibarial pump, tentorium and stipes; B–thorax; C–wing; D–anal point and tergite IX and dorsal aspect of left gonocoxite; E–hypopygium with anal point and tergite IX removed, left dorsal aspect, right ventral aspect; F–gonostylus.

ORTHOCLADIINAE © 2004 Magnolia Press 75 ZOOTAXA Legs. Spur of front tibia 40 µm long, spurs of middle tibia 23 µm and 30–33 µm long, 594 spurs of hind tibia 25 µm and 38–40 µm long. Width at apex of front tibia 33–38 µm, of middle tibia 35 µm, of hind tibia 40 µm. Hind tibial comb with 11 setae, shortest seta 15 µm long, longest seta 33 µm long. Lengths (in µm) and proportions of legs as in Table 17.

TABLE 17. Lengths (in µm) and proportions of legs of Litocladius mateusi new species, male.

fe ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR

p1 507–546 614–644 370–429 224–283 156 107 78 0.60–0.66 2.59 2.77–3.03 2.5

p2 536 575–614 224–244 136 97 58 49 0.39–0.40 4.09 4.72–4.96 2.4

p3 614–634 673–682 390–439 185–214 156–185 78–88 58 0.57–0.65 3.44–3.26 2.98–3.32 4.3

Hypopygium (Fig. 21D–F). Anal point 61–66 µm long, 25–28 µm wide at base, with 13–16 strong lateral setae, void of microtrichia except at base. Laterosternite IX with 5–7 setae. Phallapodeme 73 µm long. Transverse sternapodeme nearly straight with rounded oral projections, 98 µm long. Virga 81–88 µm long. Gonocoxite 157–164 µm long, with long, low inferior volsella, with rounded projection orally. Gonostylus 73–75 µm long, megaseta 13 µm long. HR 2.24–2.07, HV 3.04–3.22. Female (n = 1). Total length 1.38 mm. Wing length 1.02 mm. Total length / wing length 1.35. Wing length / length of profemur 2.55. Coloration yellowish brown, thorax and abdomen amber yellow, antennae brown. Head (Fig. 22A). Pedicel 37 µm long, 50 µm wide. Clypeus / width of pedicel 1.37. Flagellomeres length / width in µm: 90/22, 32/21, 30/21, 39/19, 58/18. AR 0.36. Temporal setae 5, including 2 inner verticals, 2 outer verticals and 1 postorbital. Clypeus with 10 setae. Tentorium, stipes and cibarial pump as in Fig. 22B; tentorium 90 µm long; stipes not measurable. Palpomere lengths in µm: 20, 25, 72, 81, 98. Third palpomere with 6 sensilla clavata subapically in two groups, longest 14 µm long. Thorax (Fig. 22C). Antepronotum apparently with 1 seta. Dorsocentrals 10; acrostichals apparently 4, all simple, beginning close to antepronotum; prealars 3. Scutellum with 6 setae.

Wing (Fig. 22D). VR 1.33. Brachiolum with 1 seta, R with 12 setae, R1 with 11 setae,

R4+5 with 20 setae, M with 17 setae, M3+4 with 16 setae, Cu with 6 setae, Cu1 with 6 setae,

An with 8 setae, other veins bare. Cell r4+5 with 51 setae, cell m1+2 with 87 setae, cell m3+4 with 47 setae, cells cu and an together with 68 setae. Squama with 5 setae. Legs. Spur of front tibia 22 µm long, spurs of middle tibia 25 and 17 µm long, of hind tibia 36 and 19 µm long. Width at apex of front tibia 29 µm, of middle tibia 26 µm, of hind tibia 33 µm. Hind tibial comb with 12 setae, shortest seta 11 µm long, longest seta 21 µm long. Lengths (in µm) and proportions of legs as in Table 18.

FIGURE 22. Litocladius mateusi new genus, new species, female imago. A–head; B–cibarial pump, tentorium and stipes; C–thorax; D–wing; E–genitalia dorsal view; F–dorsomesal lobe of gonapophysis VIII; G–ventrolateral lobe of gonapophysis IX; H–apodeme lobe of gonapophysis VIII.

p1 400 436 244 136 92 56 44 0.56 3.29 3.43 2.1

p2 352392------

p3 408 448 204 112 108 40 36 0.45 3.58 4.20 2.6

FIGURE 23. Litocladius mateusi new genus, new species, pupa. A–tergites I-IX; B–sternites I-IX; C–frontal apotome; D–thorax, anterior part, lateral view; E–detail of anal lobe.

Genitalia (Fig. 22E–H). Sternite VIII with about 14 setae. Gonocoxite with 2 long and strong, 1 of median length and 3 short setae. Tergite IX divided, each side with about 7 setae. Cercus 40 µm long. Seminal capsule 86 µm long, including 25 µm long neck. Notum 86 µm long.

78 © 2004 Magnolia Press MENDES ET AL. Pupa (n = 1). ZOOTAXA Total length 2.34 mm. Exuviae pale, transparent. 594 Cephalothorax. Frontal apotome as in Fig. 23C. Thoracic and cephalic setae small and not measurable. Two weak median antepronotals (Fig. 23D). Three weak precorneals in narrow triangle. Abdomen (Fig. 23A–B). Tergites II–VIII without caudal spines. Anal lobe (Fig. 23E) without macrosetae. Genital sac overreaches anal lobe by 29 µm.

Distribution

Two species of Antillocladius are known from the Oriental region, A. scalpellatus Wang and Sæther, first described from China and recently recorded from far east Russia (Tiunova et al. 2003) and A. zhengi Wang and Sæther, described from China and now recorded from Thailand. Two species are described from the Nearctic region based on material from South Carolina in south-east USA, however both have also been taken in the Neotropical region: A. pluspilalus Sæther in Nicaragua (Spies & Reiss 1996), is now recorded from Mexico and Ecuador, and A. arcuatus Sæther, is now recorded from Mex- ico, Venezuela and Brazil. In Compterosmittia there is one species in China, one species in Australia and two species known from Oceania one of which may be identical to the single species recorded from North America. The genus also has been recorded from Southeast Asia. The remaining species are all found in the Neotropical region including Mexico. All other species are Neotropical-Mexican, including some that are known only from the type locality, L. mateusi new species, C. aberrans new species, C. pittieri new species, A. biota new species, A. calakmulensis new species, A. herradurus new species, A. skartveiti Andersen and Contreras-Ramos, A. sooretama new species and A. zempoalensis new species. The other species are better known, A. antecalvus Sæther originally described from St. Vincent in the West Indies, is now recorded from Venezuela and Brazil. A. venequatoriensis new species is known from Venezuela and Ecuador. C. croizati new species is known from Venezuela and Brazil. C. berui new species, A. folius new species, A. musci new species and A. ubatuba sp. n are known from São Paulo State, Brazil; C. berui new species and A. folius new species are also recorded from Santa Catarina State; A. musci new species is also recorded from Minas Gerais State; A. ubatuba is also recorded from Rio de Janeiro State and Venezuela.

Biology of the Antillocladius, Compterosmittia and Litocladius species

The immatures are poorly known, of the 25 described species only three have the larvae associated, while the pupa is known for five species.

ORTHOCLADIINAE © 2004 Magnolia Press 79 ZOOTAXA The adults were collected by different methods, like sweep nets, light traps and Malaise 594 traps. Larvae of A. folius new species and A. musci new species were first found on the trunk and branches of a Myrtaceous tree near Lake Monte Alegre in March 1997, where they were feeding on jelly-like, dark green lichens. The air temperature this day varied from 22 to 27oC. The larvae moved in the interstitial water of the lichens, but could also be seen moving even if no water was visible. Later larvae have occasionally been found under mosses, but they shy direct exposition to the sun. However, they apparently tolerate some degree of dehydration as they sometimes become active in samples after one to several days of rehydration. In order to rear adults, larvae were transferred to the laboratory. During the pre-pupal period fourth instar larvae build a pupation chamber. The chamber has one opening, through which the insect escapes. On emergence, usually only the cephalothorax protrudes through the opening, but occasionally the whole body leaves the chamber. The adults gen- erally emerge from 06.00 am to 09.00 am. At the sampling site, some pupae have been found under leaves on the ground and then often apparently lacking a pupation chamber; a few pupae have also been encoun- tered in totally bare places like on termite mounds. According to Sæther (1984) larvae of A. pluspilalus were taken in a seep area on the shore of a reservoir in South Carolina, USA. According to Cranston et al. (1989) an asso- ciation with skunk cabbage (Symplocarpus) is indicated. The present observations show larvae of an other Antillocladius species fully adapted to a terrestrial way of life, living in a habitat similar to the larval habitat of many species of the related genera Bryophaenocla- dius and Gymnometriocnemus. The larvae of Compterosmittia all appear to be associated with plant-held waters, phytotelmata. They have been found in pitcher plants, bamboo, rot holes in trees and in the axillary liquid of an aroid (Cranston & Kitching 1995). The pupa of Litocladius was collected among mosses.

Acknowledgements

Special acknowledgements are due to Professor Dr. Claudio Gilberto Froehlich, supervisor of the first author. We are indebted to Gladys Ramirez and Vera L.O. Andrade who made the slide preparations. Dr Marion Kotrba provided the facilities to visit the Zoologische Staatssammlung, Munich. Thanks are also due to Professor Dr. Sebastião J. de Oliveira and Dr. Eliana M. Cancello, curator of the FIOCruz and MZUSP respectively. Gabriel Melo, Sidnei Mateus and Fábio de Oliveira Roque provided material from many different localities in Brazil. The State of São Paulo Research Foundation (FAPESP) (project 00/ 05903-9 and project 98/05073-4) within the BIOTA/FAPESP – The Biodiversity Virtual Institute Program (www.biota.org.br), supported this study.

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