Megaloptera: Corydalidae), with Description of a New Genus from the Oriental Realm
中国科技论文在线 http://www.paper.edu.cn Systematic Entomology (2006), 31, 652–670 DOI: 10.1111/j.1365-3113.2006.00346.x
Phylogeny of the subfamily Chauliodinae (Megaloptera: Corydalidae), with description of a new genus from the Oriental Realm
XINGYUE LIU1 andDING YANG1,2 1Department of Entomology, China Agricultural University, Beijing, China and 2Key Laboratory of Insect Evolution and Environment Change, Capital Normal University, Beijing, China
Abstract. A new Oriental fishfly genus, Sinochauliodes gen.n., is described, including four species: S. fujianensis (Yang & Yang, 1999) comb.n., S. griseus (Yang & Yang, 1999) comb.n., S. maculosus sp.n. and S. squalidus sp.n. A cladistic analysis based on adult morphological characters clarified the phylogenetic status of the new genus and allowed the reconstruction of the intergeneric relationships of the subfamily Chauliodinae. Two main clades within Chauliodinae were recog- nized from the cladistic analysis. The Asian fishflies, together with the two Nearctic genera, Chauliodes and Nigronia, formed a monophyletic lineage, and the new genus was assigned as the sister group to the genus Parachauliodes. The biogeography of the Asian fishflies is discussed.
Introduction Davis (1903) divided his Sialididae into two subfamilies, Sialidinae and Corydalinae, and subsequently van der Weele The subfamily Chauliodinae, in Corydalidae, is a very rich (1909) divided the Corydalinae into two tribes, Neuromini group in Megaloptera, comprising fifteen valid genera and and Chauliodini. Tillyard (1918) elevated Davis’ subfamilies more than 110 species from all over the world. By contrast Sialinae and Corydalinae to families and van der Weele’s with Corydalinae, the adults of Chauliodinae are character- two tribes to subfamily level as Corydalinae and Chaulio- ized by the following features: degenerative postocular plane dinae. Thus, Neuromini was synonymized with Corydalinae of the head, usually specialized antennae, three crossveins (Lestage, 1927; Glorioso, 1981). This taxonomic scheme was between R1 and Rs, and reduced male ninth gonocoxite and followed by Barnard (1931), and now has been adopted gonostylus. The larvae of Chauliodinae are distinguished generally, although Theischinger (1983) regarded the tribes from those of Corydalinae by a lack of ventral gill tufts. as having full family status. Recently, after a cladistic The fauna of Chauliodinae is richest in the New World, analysis, Contreras-Ramos (2004) noted that the Corydali- with eight genera (Dysmicohermes, Orohermes, Neohermes, dae may be paraphyletic and the relationships between Nothochauliodes, Protochauliodes, Archichauliodes, Chauli- Chauliodinae and Sialidae may be closer. odes and Nigronia), six of which are endemic. In the Major revisions of the worldwide Chauliodinae were made Afrotropical Realm, three endemic genera (Platychauliodes, by van der Weele (1909, 1910), Lestage (1927) and Kimmins Madachauliodes and Taeniochauliodes) are distributed in (1954). Apart from the scattered descriptive works of some early South Africa and Madagascar. In the Australian Realm, authors (van der Weele, 1907; Esben-Petersen, 1924; Barnard, there are only two genera (Archichauliodes and Protochauli- 1931; Nava´ s, 1932), the recent discoveries and revisions, which odes), both of which have been considered to be ‘relatively were focused on the fauna of a certain zoogeographical primitive’ (van der Weele, 1910). In the Oriental Realm, four realm, have mainly been made by Flint (1983), Theischinger endemic genera (Anachauliodes, Ctenochauliodes, Neochau- (1983), Evans (1984), Yang C. K. & Yang D. (1986, 1990, liodes and Parachauliodes) are distributed, with the last two 1992, 1999), Asahina (1987, 1988), Hayashi (1989, 1990a, genera extending far into northern China and Japan, also 1990b), Yang D. & Yang C. K. (1992, 1997), Contreras- representing Palaearctic elements. Ramos (1995), Penny (1999) and Liu & Yang (2005a, b). The intergeneric and interspecific phylogenetic relation- Correspondence: Ding Yang, Department of Entomology, ships of Chauliodinae are poorly known so far. The latest China Agricultural University, Beijing 100094, China. E-mail: key to the worldwide Chauliodinae genera (Penny, 1999) [email protected] implies that the subfamily may consist of two main clades
# 2006 The Authors 652 Journal compilation # 2006 The Royal Entomological Society
转载 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 653
defined by the different antennal morphology. One main status and interspecific relationships within Chauliodinae. clade with subserrate or pectinate antenna comprises all the Although the present analysis did not include all sixteen Asian genera and two Nearctic genera (Chauliodes and fishfly genera, the sampling involved ten genera, with all five Nigronia). The remaining genera form the other main clade Asian genera and five Australian and New World genera with filiform antenna, within which four genera (Proto- (Archichauliodes, Neohermes, Protochauliodes, Nigronia and chauliodes, Neohermes, Nothochauliodes and Taeniochauli- Chauliodes), which represent the main clades of Chauliodi- odes) may be grouped by the first branch of 2A partly fused nae. Because the generic characters are defined clearly for with 1A. However, no phylogenetic analysis has been each genus, two species were selected for coding in each undertaken on Chauliodinae. genus, except for Chauliodes, Nigronia and Sinochauliodes As with all corydalid species, the larvae of Chauliodinae with all members included. Two dobsonfly species, Chlor- are predaceous, living in clean streams of mountainous oniella peringueyi Esben-Petersen from South Africa and regions and being sensitive to environmental changes. They Neoneuromus ignobilis Nava´ s from China, were selected as may be useful as a biological indicator of water quality the outgroup taxa because Corydalinae is generally consid- (Yang D. & Yang C. K., 1995). The adults emerge from late ered to be ‘more primitive’ than Chauliodinae. A complete spring to early autumn, and are always found near clean list of the taxa used in the phylogenetic analysis is given in water. Adults are readily collected at light traps. Detailed Table 1. studies on the natural history of Chauliodinae are mostly restricted to a few species from temperate regions. Contri- Characters butions to the behaviour have been made by Hayashi (1996, 1999) based on the observation of some Japanese species. Fifty-nine adult morphological characters were selected The present paper describes a new Chauliodinae genus, for coding, with nine cephalic, fourteen thoracic and thirty- Sinochauliodes, from Oriental China. Four species belonging six genitalic characters. The data matrix used in the analysis to the new genus are described and keyed, with two species is shown in Table 2 (0, plesiomorphic state; 1–3, apomor- new to science. The generic status and specific relationships of phic state; ?, state inapplicable). the new genus are discussed based on a phylogenetic analysis of all five Asian and five New World and Australian genera. The biogeography of the Asian fishflies is also discussed. Head
1. Head: (0) flattened; (1) robust. Materials and methods 2. Head: (0) without long setae on ventral lateral portion; (1) with long setae on ventral lateral portion. Taxa and terminology 3. Postocular plane: (0) present; (1) absent. 4. Anteclypeus: (0) articulated to postclypeus; (1) fused to Most specimens examined were obtained from the Ento- postclypeus. mological Museum of China Agricultural University 5. Maxillary palp: (0) five-segmented; (1) four-segmented. (CAU), Beijing, with one paratype of S. maculosus depos- State 1 of the above five characters was considered to be ited in the Institute of Zoology, Chinese Academy of Science synapomorphic for Chauliodinae. (IZCAS), Beijing. The Australian and New World fishfly 6. Male antenna: (0) short, less than one-half of forewing; (1) specimens were obtained through the kind exchange by long, exceeding two-thirds of forewing. Dr O.S. Flint, Jr. from the National Museum of Natural The male antenna is less than one-half of the forewing History (NMNH), Washington DC, U.S.A. in the basal Corydalinae genera and many Chauliodinae The specimens were collected mostly at light traps in genera, whereas it is elongated, exceeding two-thirds mountainous regions, with some of these collected by the of the forewing, in Archichauliodes, Neohermes and authors. Genitalic preparations were made by macerating the Protochauliodes. apex of the abdomen in cold 10% KOH for 8–10 h. The apex 7. Male antenna: (0) filiform; (1) moniliform; (2) subserrate; of the abdomen was then transferred to glycerine for further (3) pectinate. dissection and examination. After examination, it was moved The feature of the antenna in Chauliodinae is an im- to fresh glycerine and stored in a microvial pinned below the portant diagnostic character at the generic level. In most specimen. The habitus photographs of adults were obtained primitive genera of Chauliodinae, the male antenna is with a digital camera (Coolpix 4500, Nikon, Japan). filiform or moniliform. However, in the Asian genera and The terminology of the adult genitalia generally follows two Nearctic genera (Chauliodes and Nigronia), the male Contreras-Ramos (1998, 2004) in order to unify the terms antenna is subserrate or pectinate, which is considered to be with Corydalinae. apomorphic. The pectinate antenna is considered to be derived from the subserrate form. Phylogenetic analysis 8. Male antenna: (0) not pectinate; (1) pectinate with branches not distally widened; (2) pectinate with branches slightly Our present phylogenetic analysis aimed to test the widened distally; (3) pectinate with branches strongly monophyly of Sinochauliodes and to establish its generic widened distally.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
654 X. Liu and D. Yang
Table 1. Taxa studied in the present phylogenetic analysis and Table 1. Continued. their distribution. Taxa Distribution Taxa Distribution Sinochauliodes fujianensis China (south-eastern) Corydalinae (Yang & Yang, 1999), comb.n. Chloroniella Esben-Petersen, Sinochauliodes griseus China (south-eastern) 1924 (Yang & Yang, 1999), comb.n. Chloroniella peringueyi South Africa Sinochauliodes maculosus China (southern) Esben-Petersen, 1924 Liu & Yang, sp.n. Neoneuromus Weele, 1909 Sinochauliodes squalidus Liu & China (southern) Neoneuromus ignobilis China Yang, sp.n. Nava´ s, 1932
Chauliodinae The branches of the pectinate antenna are not widened Anachauliodes Kimmins, 1954 distally in Ctenochauliodes, but are slightly widened distally Anachauliodes sinensis China (south-western) Yang & Yang, 1992 in Anachauliodes, Chauliodes, Neochauliodes and Sinochau- Anachauliodes tonkinicus Vietnam liodes. The branches are strongly shortened and widened Kimmins, 1954 distally in Nigronia fasciatus. Archichauliodes Weele, 1909 9. Female antenna: (0) filiform; (1) subserrate; (2) pectinate. Archichauliodes chilensis Chile In Chauliodinae, the antenna usually shows distinct sexual Kimmins, 1954 dimorphism. In most Chauliodinae genera, the female Archichauliodes diversus New Zealand antenna is not strongly modified, either filiform or subser- (Walker, 1853) rate. In Ctenochauliodes and Chauliodes pecticornis, the Chauliodes Latreille, 1802 female antenna is similar to that of the male, also pectinate. Chauliodes pecticornis U.S.A. (eastern), (Linnaeus, 1763) Canada (eastern) Chauliodes rastricornis U.S.A. (eastern), Thorax Rambur, 1842 Canada (eastern) Ctenochauliodes Weele, 1909 Ctenochauliodes friedrichi China (southern and 10. Tarsal claw: (0) simple; (1) with additional subdistal claw. Nava´ s, 1932 south-western) The tarsal claw is of a similar simple shape in most species Ctenochauliodes meridionalis China (southern) of Megaloptera. However, in addition to the distal claw, Yang & Yang, 1986 there is an additional subdistal claw in Neohermes. Neochauliodes Weele, 1909 11. Pigmentation marks of wing: (0) dotted in a disperse Neochauliodes fraternus China manner; (1) united into rather distinct transverse band. (McLachlan, 1869) In Chauliodinae, the pigmentation marks of the wings are Neochauliodes sinensis China (southern) usually small and dotted in a disperse manner, whereas, in (Walker, 1853) Nigronia and most species of Neochauliodes, the marks are Neohermes Banks, 1908 connected into a distinct transverse band in the middle. Neohermes concolor U.S.A. (Davis, 1903) 12. Coloration of wing: (0) hyaline or subhyaline with dark Neohermes filicornis U.S.A., Mexico marks; (1) entirely blackish. (Banks, 1903) In most species of Chauliodinae, the wings are hyaline or Nigronia Banks, 1908 subhyaline with several dark marks, whereas they are Nigronia fasciatus U.S.A. (eastern), Mexico entirely blackish in Sinochauliodes fujianensis. (Walker, 1853) 13. Coloration of veins: (0) predominantly brown; (1) Nigronia serricornis U.S.A. (eastern) alternating dark and pale pattern. (Say, 1824) In most genera of Chauliodinae, the veins are mostly Parachauliodes Weele, 1909 brown throughout. In Anachauliodes and Chauliodes, the Parachauliodes japonicus Japan (northern) veins show a distinct alternating dark and pale pattern (McLachlan, 1867) Parachauliodes neblosus China (Taiwan) (synapomorphy). (Okamoto, 1910) 14. R2: (0) two-branched or more; (1) simple. Protochauliodes Weele, 1909 The simple R2 is considered to be apomorphic, and is Protochauliodes bullocki Chile shared by Protochauliodes, Neohermes and three South Flint, 1973 African genera which are excluded in the present analysis. Protochauliodes cinerascens Chile In Archichauliodes, Chauliodes, Nigronia, Nothochauliodes, (Blanchard, 1851) Dysmicohermes, Orohermes and the five Asian genera, R2 is Sinochauliodes Liu & Yang, of two branches or more. gen.n. 15. R4: (0) simple; (1) two-branched. R4 is simple in most genera of Chauliodinae, but it is two-branched in Protochauliodes and Neohermes.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 ora compilation Journal # 06TeAuthors The 2006 # 06TeRylEtmlgclSociety, Entomological Royal The 2006
Table 2. Character matrix.
11111111112222222222333333333344444444445555555555 12345678901234567890123456789012345678901234567890123456789
Chloroniella peringueyi 00000000000000010000100000000000000000000000000010000000000 Neoneuromus ignobilis 00000000000000000000100000000000000000000000000010000000000 Anachauliodes sinensis 11111031100010011001000100010011100011100010010000000110110 Anachauliodes tonkinicus 11111031100010011001000100010011100011100010010000000110110 Archichauliodes chilensis 11111100000000010001000010000000001010100000100000000110101 Archichauliodes diversus 11111100000000010001000010000000001010100000100000000010101 Chauliodes pecticornis 11111031200010010001000100010011100011100000000000000110110 Chauliodes rastricornis 11111031100010010001010100010011100011100000000011000110110 ytmtcEntomology Systematic Ctenochauliodes friedrichi 11111030200000010001001100100110020010100000100111110010100 Ctenochauliodes meridionalis 11111030200000010001001100100110020010100000100111110010100 Neochauliodes fraternus 11111031101000010010001100111101010011111000010000000101100 Neochauliodes sinensis 11111031101000010010001100111101010011111000010000000101100 Neohermes concolor 11111110010001110101100010000000001011100000000010000001001 Neohermes filicornis 11111110010001110101100010000000001011100000000010000001001 Nigronia fasciatus 11111033101000010011001100111101010011131000010011101101100 Nigronia serricornis 11111020101000010011001100111101010011131001010011101101100 , hlgn ftesbaiyChauliodinae subfamily the of Phylogeny
31 Parachauliodes japonicus 11111020100000010011001100011101010011120001010000000101100
652–670 , Parachauliodes neblosus 11111000100000010011001100011101010011120000010000000????00 Protochauliodes bullocki 11111100000001110101100010000000001011100100000100000001001 Protochauliodes cinerascens 11111130000001110101100010000000001011100101000000000001001 Sinochauliodes fujianensis 11111031100100010011001101011101010011110001011000000101100 http://www.paper.edu.cn Sinochauliodes griseus 11111031100000010011001101011101010011110001010000000101100 Sinochauliodes macuosus 11111031100000010011001101011101010111110001011000000?????? Sinochauliodes squalidus 11111031100000010011001101011101010011110001011000000101100 655 中国科技论文在线 http://www.paper.edu.cn
656 X. Liu and D. Yang
16. R5: (0) branched; (1) simple. is incised, whereas it is truncate in Archichauliodes, Neo- Branched R5 is considered to be plesiomorphic and hermes and Protochauliodes. shared by the outgroup species and two Nearctic fishfly 26. Ninth tergum in dorsal view: (0) wider than long; (1) genera Dysmicohermes and Orohermes, whereas it is simple longer than wide. in all the present ingroup taxa. In most genera of Corydalidae, the ninth tergum is wider 17. 1A of forewing: (0) two-branched; (1) three-branched or than long, but longer than wide in Sinochauliodes. more. 27. Ninth tergum in lateral view: (0) with ventral margin In Anachauliodes, 1A of the forewing is three-branched or obtusely produced; (1) with ventral margin acutely more (autapomorphy). produced. 18. 2A of forewing with anterior branch: (0) not fused to 1A; In Ctenochauliodes, Neochauliodes and Nigronia,the (1) medially fused to 1A. ninth tergum is acutely produced at the ventral margin in The anterior branch of 2A is not fused to 1A in the lateral view (synapomorphy). forewing in many genera of Corydalidae and is medially 28. Ninth sternum: (0) as long as ninth tergum; (1) shorter fused to the footstalk of 1A in a short distance, which is than ninth tergum. synapomorphic for grouping Neohermes, Protochauliodes, In the ‘basal’ genera of Chauliodinae, the male ninth Nothochauliodes and Taeniochauliodes. sternum is broad and as long as the male ninth tergum, 19. Crossvein between 1A and 2A of forewing: (0) placed whereas it is reduced and rather shorter than the ninth between posterior branch of 1A and anterior branch of tergum in Chauliodes, Nigronia, Anachauliodes, Neochauli- 2A; (1) placed between footstalks of 1A and 2A. odes, Parachauliodes and Sinochauliodes. In Corydalinae and many genera of Chauliodinae, the 29. Ninth sternum: (0) without membranous apical process; crossvein between 1A and 2A in the forewing is placed (1) with membranous apical process. between the posterior branch of 1A and the anterior branch The male ninth sternum is connected by a membrane to of 2A, whereas, in Nigronia, Neochauliodes, Parachauliodes the basal portion of the tenth sternum. In Nigronia, Neo- and Sinochauliodes, this crossvein is placed rather basally, chauliodes, Parachauliodes and Sinochauliodes, the mem- connecting the footstalk of 1A and 2A. brane is produced backward, forming a small subtriangular 20. 2A of forewing: (0) not sinuated; (1) strongly sinuated. process at the tip of the male ninth sternum. 2A of forewing is strongly sinuated in most Chauliodinae 30. Tenth tergum: (0) as long as or longer than ninth tergum; genera except Neochauliodes. (1) shorter than ninth tergum.
21. M1 þ 2 of hindwing: (0) simple; (1) two-branched. In Ctenochauliodes, Nigronia, Neochauliodes, Parachauli- In most genera of Chauliodinae, M1 þ 2 is simple, odes and Sinochauliodes, the male tenth tergum is much whereas, in Protochauliodes and Neohermes,M1 þ 2 is two- shorter than the male ninth tergum, whereas it is usually as branched. long as or longer than the male ninth tergum in the other 22. Basal r-m crossvein of hindwings: (0) present; (1) absent. ingroup genera and the outgroups. In most genera of Corydalidae, there is a basal r-m 31. Tenth tergum: (0) not narrowing towards apex; (1) crossvein in the forewing, but this crossvein is absent in narrowing towards apex. Platychauliodes, Taeniochauliodes, Nothochauliodes and In Anachauliodes, Chauliodes and Ctenochauliodes, the Chauliodes rastricornis, which may represent parallel loss. tenth tergum is wide at the base and narrows towards the 23. Basal r-m crossvein of hindwings: (0) simple; (1) modified. apex, whereas, in the other genera of Chauliodinae, the tenth In the outgroup species, the basal r-m crossvein of the tergum does not narrow towards the apex and even with hindwing is simple, whereas it reconnects to M by an addi- the apex inflated. tional short branch or a short fusion in Nigronia, Ctenochauli- 32. Tenth tergum in lateral view with base: (0) narrower than odes, Neochauliodes, Parachauliodes and Sinochauliodes. one-half of ninth tergum; (1) wider than two-thirds of ninth tergum. In Anachauliodes, Chauliodes, Neochauliodes, Nigronia, Male genitalia Parachauliodes and Sinochauliodes, the base of the tenth tergum is wider than two-thirds of the ninth tergum in 24. Ninth tergum in dorsal view: (0) posteriorly incised; (1) lateral view, whereas it is narrower than one-half of the posteriorly truncate or arcuately produced. ninth tergum in the other Chauliodinae genera. In the outgroup species, Archichauliodes, Neohermes 33. Tenth tergum: (0) without strongly sclerotized distal and Protochauliodes, the posterior margin of the ninth claws; (1) with a few strongly sclerotized distal claws. tergumisincisedmoreorless,whereasitistruncate In Anachauliodes and Chauliodes, the tenth tergum bears or arcuately produced in Anachauliodes, Chauliodes, Nigro- a few strongly sclerotized distal claws (synapomorphy). nia, Ctenochauliodes, Neochauliodes, Parachauliodes and 34. Tenth tergum with spinous setae: (0) feebly modified; (1) Sinochauliodes. moderately modified; (2) strongly modified. 25. Ninth tergum in dorsal view: (0) anteriorly incised; (1) In Chauliodinae, there are usually several spinous setae anteriorly truncate. on the distal portion of the tenth tergum. In Archichauliodes, In the outgroup species, Chauliodes, Nigronia and the Neohermes and Protochauliodes, the spinous setae are simply Asian fishfly genera, the anterior margin of the ninth tergum slender and dispersed without modified arrangement. In
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 657
Nigronia, Neochauliodes, Parachauliodes and Sinochaulio- 43. Tenth sternum in lateral view: (0) curved dorsally; (1) des, the spinous setae are moderately modified in several curved ventrally. rows. In Ctenochauliodes, the spinous setae are sparse, In most genera of Chauliodinae, the tenth sternum is rather stout and strongly sclerotized, showing the strongly curved dorsally, but, in Anachauliodes, the distal portion of modified style. the tenth sternum is curved ventrally. 35. Tenth tergum in dorsal view with proximal inner portion: 44. Tenth sternum in lateral view: (0) mostly visible; (1) (0) strongly sclerotized; (1) almost membranous. mostly invisible. The tenth tergum is strongly sclerotized throughout in most In Parachauliodes and Sinochauliodes, the tenth sternum Chauliodinae genera, whereas the proximal inner portion of is mostly enveloped by the male ninth tergum and invisible the tenth tergum is almost membranous in dorsal view in in lateral view (synapomorphy). Archichauliodes, Neohermes and Protochauliodes. 45. Tenth sternum with bifurcation: (0) symmetric; (1) 36. Tenth tergum in lateral view with proximal margin: (0) asymmetric. vertical; (1) oblique. In most genera of Chauliodinae, the tenth sternum is In Sinochaulioides fujianensis and S. maculosus, the prox- bifurcated or with its apex slightly incised, forming a pair of imal margin of the tenth tergum is rather oblique (synapo- symmetric lobes. However, in some species of Archichauli- morphy). odes and Ctenochauliodes, the tenth sternum is bifurcated 37. Cercus: (0) free, between ninth gonostylus and tenth with a pair of asymmetric lobes. tergum; (1) fused to base of tenth tergum. 46. Tenth sternum with bifurcation: (0) distinct; (1) indistinct. The cercus in Corydalinae is free between the ninth The distinctly bifurcated tenth sternum is considered to be gonostylus and the tenth tergum, whereas it is fused to the plesiomorphic and shared by many genera of Chauliodinae, base of the tenth tergum in Chauliodinae (apomorphy). whereas, in Anachauliodes, Nigronia, Neochauliodes, Para- 38. Cercus: (0) distinctly prominent; (1) feebly prominent. chauliodes and Sinochauliodes, the bifurcation of the tenth The feebly prominent cercus is considered to be apomor- sternum is indistinct. phic and shared by most genera of Chauliodinae, but in 47. Tenth sternum with apex: (0) incised or feebly produced; Archichauliodes and Ctenochauliodes, the cercus is distinctly (1) strongly produced. prominent. The distally unbifurcated tenth sternum is usually slightly 39. Ninth gonocoxite and gonostylus: (0) entire; (1) reduced. incised or produced feebly, but it is strongly produced in The reduction of the male ninth gonocoxite and gono- three Sinochauliodes species. stylus is a synapomorphic character of Chauliodinae. 48. Tenth sternum with distal half in lateral view: (0) feebly 40. Ninth gonocoxite and gonostylus: (0) absent; (1) present widened; (1) strongly widened. but indistinct; (2) present and roundly produced; (3) In Ctenochauliodes, the distal half of the tenth sternum is present and spinously produced. strongly widened in lateral view, as well as in Protochauli- The morphology of the reduced male ninth gonocoxite odes bullocki, which may be a parallel derived state. and gonostylus varies in Chauliodinae. The absence of the However, it is considered to be autapomorphic to Cteno- male ninth gonocoxite and gonostylus herein is considered chauliodes. to be plesiomorphic; thus, the presence of the structures is 49. Lateral lobes: (0) absent; (1) present. secondarily derived. In Neochauliodes and Sinochauliodes, In many genera of Corydalinae, there is a pair of the reduced ninth gonocoxite is present but indistinct. In lateral lobes on the posterior margin of the tenth sternum. Parachauliodes, the ninth gonocoxite is distinctly produced However, the absence of the lateral lobes is herein with the round tip forming a ‘bilobed’ tenth tergum. In considered to be plesiomorphic. In many genera of Nigronia, the ninth gonocoxite is acutely produced with the Chauliodinae, the lateral lobes are absent, but are present spinous gonostylus. in many species of Ctenochauliodes, Nigronia and Chauli- 41. Tenth sternum with lateral arm: (0) as long as or longer odes rastricornis. The reduced ninth gonostylus in Neo- than one-half of the median plate; (1) shorter than one- hermes, which is figured by Contreras-Ramos (2004), is quarter of the median plate. considered herein to be the lateral lobe of the tenth In Chauliodinae, the tenth sternum generally comprises sternum also. a pair of lateral arms and a median plate. The lateral arm is 50. Lateral lobes: (0) fused to lateral arms of tenth sternum; the structure for connecting the tenth sternum to the ninth (1) separated from tenth sternum. tergum. In most genera of Chauliodinae, the lateral arm is as In Neohermes, the lateral lobes are fused to the lateral long as or longer than one-half of the median plate, but it is arms of the tenth sternum, whereas, in Ctenochauliodes, shorter than one-quarter of the median plate in Nigronia Nigronia and Chauliodes rastricornis, the lateral lobes are and Neochauliodes. separated from the tenth sternum. 42. Tenth sternum with median plate: (0) longer than wide, 51. Lateral lobes: (0) separated from each other; (1) con- distally pointed; (1) wider than long, distally widened. nected by a sclerotized plate. In most genera of Chauliodinae, the median plate of the In Ctenochauliodes and Nigronia, the lateral lobes are tenth sternum is longer than wide and pointed distally, but it connected by a sclerotized plate, which may be a remnant of is wider than long and distally widened in Protochauliodes the paramere. (autapomorphy). 52. Plate between lateral lobes: (0) narrow; (1) broad.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
658 X. Liu and D. Yang
In Ctenochauliodes, the sclerotized plate connecting the and the branches with bootstrap values of < 50% collapsed. lateral lobes is broad, but it is rather narrow in Nigronia. Bremer’s decay index was calculated with Autodecay ver- 53. Plate between lateral lobes: (0) feebly sclerotized; (1) sion 4.0 (Eriksson, 1998) and PAUP* version 4.0b10. The strongly sclerotized. unambiguous characters were mapped by MacClade version In Ctenochauliodes, the plate connecting the lateral lobes 4.0 (Maddison & Maddison, 2000). is generally feebly sclerotized, but it is strongly sclerotized in Nigronia. Results Female genitalia The heuristic analysis resulted in three most parsimonious 54. Eighth sternum with posterior margin: (0) nearly truncate; trees [length ¼ 92, consistency index (CI) ¼ 0.71, retention (1) produced. index (RI) ¼ 0.90, rescaled consistency index (RC) ¼ 0.65]. The nearly truncate posterior margin of the female eighth The three most parsimonious trees only differed in the sternum is plesiomorphic and shared by most species of topology of Anachauliodes and Chauliodes, with Chauliodes Corydalinae and many genera of Chauliodinae. In Ana- pecticornis as the sister group of Anachauliodes,with chauliodes, Chauliodes, Nigronia, Neochauliodes, Parachau- Chauliodes rastricornis as the sister group of Anachauliodes, liodes and Sinochauliodes, the female eighth sternum is or with two Chauliodes species to form a monophyletic usually posteriorly produced into a subgenital plate. group. One of the three most parsimonious trees is shown in 55. Tenth tergum: (0) thick; (1) slender. Fig. 1, and the strict consensus for the three most parsimo- The thick female tenth tergum is plesiomorphic and nious trees is given in Fig. 2. In the latter, the ingroup is presented in most genera of Corydalidae, but in Anachauli- divided into two monophyletic clades. Clade 1 contains the odes, Archichauliodes, Chauliodes and Ctenochauliodes, the five Asian Chauliodinae genera and two Nearctic genera female tenth tergum is rather slender, much narrower than (Chauliodes and Nigronia). Within clade 1, Ctenochauliodes the diameter of the cercus. is sister to the remaining clade, which consists of three 56. Cercus: (0) strongly prominent; (1) feebly prominent. monophyletic lineages: Anachauliodes lineage (Anachaulio- The strongly prominent cercus is considered to be plesio- des þ Chauliodes), Neochauliodes lineage (Neochauliodes þ morphic and is presented in a few Chauliodinae genera, such Nigronia)andParachauliodes lineage (Parachauliodes þ as Anachauliodes, Archichauliodes, Chauliodes and Cteno- Sinochauliodes). Clade 2 comprises Archichauliodes, Neo- chauliodes. hermes and Protochauliodes, with the last two genera as 57. Gonostylus on gonocoxite: (0) present; (1) absent. sisters. In Corydalidae, the female gonostylus is placed at the tip of the gonocoxite and plesiomorphically articulated to the gonocoxite. In most genera of Chauliodinae, the female Phylogeny gonostylus is absent, but it remains in Protochauliodes and Neohermes. Clade 1 58. Gonocoxite with apex: (0) pointed; (1) rounded. The general form of the gonocoxite is triangular or foliate According to the present cladogram, the Asian fishflies with a pointed apex; however, a derived form of the were assigned as a corroborated monophyletic group with gonocoxite with a rounded apex is presented in Anachauli- two Nearctic genera, Chauliodes and Nigronia, and the odes and Chauliodes. monophyly was supported by the pectinate male antenna 59. Gonocoxite with lateral sclerotized plate: (0) subtriangu- (character 7:3) and the male ninth tergum with the posterior lar; (1) subquadrate. margin not incised (character 24:1). The modified basal r-m In Corydalidae, there is usually a lateral sclerotized plate crossvein in the hindwing (character 23:1) also partially on the female gonocoxite, which is generally subtriangular. supports the monophyly of clade 1, whereas, in Anachauli- However, in some genera of Chauliodinae (Archichauliodes, odes and Chauliodes, the absence or reduction of the basal Neohermes, Protochauliodes), the lateral sclerotized plate is r-m crossveins may be lost secondarily. large and subquadrate. Ctenochauliodes was well supported to be monophyletic by the spare stout spinous setae on the male tenth tergum Cladistic analysis (character 34:2) and the broad plate connecting the lateral lobes of the male tenth sternum (character 52:1). The The cladistic analysis was performed in PAUP* version strongly developed pectinate antenna in both sexes may 4.0b10 (Swofford, 2002) using heuristic parsimony analysis, also be synapomorphic to the Ctenochauliodes species, and with 1000 random stepwise additions of taxa [tree-bisection– parallel in Chauliodes pecticornis. Compared with the other reconnection (TBR) branch swapping] under ACCTRAN opti- genera within clade 1, Ctenochauliodes retains more plesio- mization, characters unordered and of equal weight and morphic characters, such as the totally absent male ninth MulTrees option in effect. Bootstrap values for clades were gonocoxite and gonostylus, the well-developed male ninth calculated in 1000 replicates using a general heuristic search sternum without distal membranous process and the
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 659
Fig. 1. One of the three most parsimonious trees with Chloroniella peringueyi and Neoneuromus ignobilis as the outgroups (see text for further information). Only unambiguous characters are shown. Filled squares represent apomorphic characters, open squares represent parallelisms and reversals. Character states for multistate characters and reversals are placed above the squares.
bifurcated male tenth sternum. Therefore, the origin of (character 29:1), the rows of the spinous setae (character 34:1), Ctenochauliodes may be earlier than some ‘younger’ taxa the presence of the reduced male ninth gonocoxite and within clade 1, such as the Neochauliodes lineage and the gonostylus (character 40:1) and the feebly prominent male Parachauliodes lineage. cercus (character 56:1). Kimmins (1954) mentioned that the The monophyly of the Anachauliodes lineage þ Neochauli- female genital characters cannot be distinguished from each odes lineage þ Parachauliodes lineage was well supported other between Neochauliodes and Parachauliodes, and he by the shortened male ninth sternum (character 28:1), the proposed that these two genera should be considered as male tenth tergum with the base wider than two-thirds of synonyms. However, in the present study, the rather different the width of the ninth tergum in lateral view (character 32:1), features of the male sternum justify generic status. the feebly prominent cercus (character 38:1) and the distinctly The Neochauliodes lineage was well supported by the produced female eighth sternum (character 54:1). peculiar wing patterns (character 11:1), the male ninth The grouping of the Nearctic genus Chauliodes with the sternum with the ventral margin acutely produced (charac- Oriental genus Anachauliodes was well supported by the ter 27:1) and the shortened lateral arms of the male tenth peculiar coloration of the veins (character 13:1), the pres- sternum (character 41:1). The non-sinuate anal veins are ence of the distal claws of the male tenth tergum (character reversed autapomorphic characters of Neochauliodes. Nigronia 33:1) and the roundly produced female gonocoxite (charac- appears to be a rather derived genus from the common ter 58:1). However, the grouping of the two Chauliodes ancestor with Neochauliodes in having many highly modi- species was not convincing, which may be a result of more fied autapomorphic characters. plesiomorphic characters retained in Chauliodes. The Ana- The lineage Parachauliodes was well supported by the chauliodes lineage is considered to be more primitive than male tenth sternum enveloped in the male ninth tergum the group of the Neochauliodes lineage and Parachauliodes (character 44:1). Van der Weele (1910) mentioned that lineage by having some distinguished plesiomorphic char- Parachauliodes may have close affinities with Protochauli- acters, such as the bifurcated male tenth sternum and the odes, which is distributed in the New World and the strongly prominent female cercus. Australian Realm, and he also noted that Parachauliodes The monophyletic group of the Neochauliodes lineage and may be the precursor of Neochauliodes.Bycontrastwith Parachauliodes lineage was well supported by the position of van der Weele‘s assumption, the current phylogenetic the crossvein between 1A and 2A (character 19:1), the presence hypothesis is that Parachauliodes belongs to an indepen- of the membranous distal plate of the male ninth sternum dent lineage of Neochauliodes, and has a rather distant
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
660 X. Liu and D. Yang
Fig. 2. The strict consensus tree of the three most parsimonious trees with boot- strap value (above branches) and Bremer’s decay index (below branches).
affinity with Protochauliodes. Compared with Sinochauli- As a result of a lack of the three Afrotropical genera odes, the subserrate male antenna in Parachauliodes implies (Platychauliodes, Madachauliodes and Taeniochauliodes) its primitiveness. The monophyly of the new genus Sino- and three New World genera (Dysmicohermes, Orohermes chauliodes was supported by the elongation of the male and Nothochauliodes), the current hypothesis concerning ninth tergum (character 26:1). Although the general fea- clade 2 is preliminary. However, some aspects on the generic ture of the genitalia shows the great similarity in Para- phylogeny can be inferred on the basis of the current results chauliodes and Sinochauliodes, the generic status of and previous studies (Barnard, 1931; Paulian, 1951; Chandler, Sinochauliodes is corroborated by the more derived male 1954; Flint, 1983; New & Theischinger, 1993; Penny, 1999). antenna, the strongly produced male tenth sternum and the Firstly, the nine genera (Archichauliodes, Platychauliodes, elongated male ninth tergum. Madachauliodes, Dysmicohermes, Orohermes, Neohermes, Within Sinochauliodes, S. squalidus, S. fujianensis and Protochauliodes, Nothochauliodes and Taeniochauliodes) S. maculosus form a monophyletic group supported by the probably are grouped into a monophyletic clade based on strongly produced male tenth sternum (character 47:1). The the long filiform antenna and the anteriorly truncate male sister group relationships between S. fujianensis and ninth tergum. Thus, the clade may consist of more primitive S. maculosus are supported by the male tenth tergum with genera than clade 1 by having many endemic genera, which the rather oblique basal margin in lateral view (character retain many plesiomorphic characters (such as the filiform 36:1). Thus, Sinochauliodes griseus may be sister to the rest antenna and the presence of the female gonostylus) and are of of the genus. rather disjunct and limited distributions. Secondly, within the nine genera, Neohermes, Protochauliodes, Nothochauliodes and Taeniochauliodes may form a monophyletic subclade, Clade 2 which is supported by the first branch of 2A medially fused with 1A in the forewing (character 18:1). Furthermore, the The monophyly of Archichauliodes þ Neohermes þ Pro- sister relationships between Neohermes and Protochauliodes tochauliodes was well supported by the long male antenna are confirmed by the branched R4 in the forewing. (character 6:1), the anteriorly truncate male ninth tergum The distinguished lineage of Dysmicohermes þ Orohermes (character 25:1), the male tenth tergum with the inner seems to be highly derived and closely related to the proximal portion membranous (character 35:1) and the Protochauliodes lineage by the branched R4 in the forewing. female gonocoxite with the lateral sclerite strongly devel- The branched R5 and M1 þ 2 in the hindwing are considered oped (character 59:1). herein to be plesiomorphic, although they may also be
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 661
secondarily derived characters for grouping the Dysmico- antenna in P. buschi originally was described as pectinate, hermes lineage and the Protochauliodes lineage. Thus, the and 1A in the forewings of P. laboissierei was illustrated
polarization of the states of R5 and M1 þ 2 requires further originally as four-branched, which is the autapomorphic evaluation by adding more outgroups. character of Anachauliodes. Thus, the distribution pattern of Penny (1999) proposed close relationships between Parachauliodes is confirmed herein, probably along the Archichauliodes, Platychauliodes and Madachauliodes based West Pacific Island Arc. Interestingly, the genus Sinochauli- on the unbranched R5, 1A not fused to 2A in the forewing odes, which is the sister to Parachauliodes, occurs only on and the filiform antenna. However, all three characters are the Chinese mainland and is restricted mostly to south- probably plesiomorphic. Therefore, any hypothesis on the eastern China, which is close to the West Pacific Island Arc. relationships between the three genera should be postponed pending future comprehensive studies. Distribution tracks of the Asian fishflies
According to the distribution patterns and the present Biogeography cladogram, the Asian fishflies reveal north-eastward and south-eastward distribution tracks. The north-eastward Distribution patterns of the Asian fishflies distribution track can be recognized from south-western China and north-eastern India, through central China to Compared with the distributions of the other Asian south-eastern and northern China. The relatively young Chauliodinae genera, the distribution of Neochauliodes is genera of Asian fishflies (Parachauliodes and Sinochaulio- widest, from south-eastern to far eastern Asia, and most des) are restricted to south-eastern China, even extending ‘primitive’ Neochauliodes species are restricted to the northwards to Palaearctic Japan. The distribution patterns Oriental Realm, especially concentrated in south-western of the Neochauliodes species also fit this track, with many China and north-eastern India (Liu & Yang, 2005a). The ‘younger’ species, such as Neochauliodes sinensis complex, Ctenochauliodes species are restricted to the Oriental Realm dominantly occupying northern China (Liu & Yang, and are richest in China (van der Weele, 1910; Nava´ s, 1932; 2005b). According to the fossil record of the Chauliodinae Kimmins, 1954; Yang C. K. & Yang D., 1986, 1990, 1992, from the Early Cretaceous of Baissa in Siberia (Cretochaulus 1999). The distribution of Anachauliodes is extremely nar- lacustris; Ponomarenko, 1976), the north-eastward distribution row, and this genus can be found only in a few mountainous track may be at least 140 Myr old. The combination of the regions of south-western China and northern Vietnam two Nearctic genera, Chauliodes and Nigronia, with the (Kimmins, 1954; Yang D. & Yang C. K., 1992). Undoubt- Asian genera and the Baltic Amber Chauliodes (Pictet, 1854; edly, four Parachauliodes species are distributed in Japan Wichard, 2003) also implies that Chauliodinae may have and Taiwan (Hayashi, 1990b), showing peculiar adaptation been widely distributed in Laurasia with some recent genera to the island habitats. According to our unpublished studies, originating during the Late Cretaceous or early Eocene. The records of the other two species of Parachauliodes from the distribution areas of Chauliodinae probably decreased and Chinese mainland and Vietnam (Nava´ s, 1913, 1924) are then increased several times during glacial and postglacial inaccurate, perhaps due to misidentifications. The male periods in the Cenozoic until the recent fauna was formed.
Fig. 3. Distribution map of the genus Sinochauliodes and its sister genus Para- chauliodes, implying the division of the sister genera caused by sea invasion between the Chinese mainland and the Western Pacific Island Arc (the full line demarcates the distribution areas of Sino- chauliodes and the broken line demarcates the distribution areas of Parachauliodes).
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
662 X. Liu and D. Yang
The south-eastward distribution track is probably from the Asian dobsonflies proposed by Penny (1993). As he south-western China and north-eastern India, through mentioned, the Asian dobsonfly genera could have been southern China, Burma, Thailand and Vietnam, to Malay- present either in south Asia from a previous vicariant event sia and Indonesia. Part of the Neochauliodes species may (perhaps the sundering of Pangea) or may have been in- have dispersed along this track. troduced as single or multiple ancestors on the Gondwanan- derived Indian subcontinent as it drifted northwards. In Origin of the Asian fishflies addition, he pointed out that the biogeographical pattern of Chauliodinae may also fit his theory of Gondwanan So far, the biogeography of the Chauliodinae is still distribution. According to the above discussion on the poorly known, with a working hypothesis for the origin of distribution track of the Asian fishflies, the Chauliodinae
Fig. 4. Habitus photographs of the Sinochauliodes species. A, S. fujianensis, holotype male; B, S. fujianensis, allotype female; C, S. griseus, paratype male; D, S. griseus, paratype female; E, S. maculosus, holotype male; F, S. squalidus, holotype male; G, S. squalidus, paratype female.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 663
may have originated before the Early Cretaceous, at the with V-shaped basal incision; ninth sternum less sclero- time of separation of Laurasia and Gondwana. Thus, the tized and much shorter than ninth tergum, with one hypothesis that the Corydalidae was introduced firstly to membranous subtriangular apical process; tenth tergum Laurasia by the Indian subcontinent is rejected. Considering simple and laterally flattened with round apical margin, the current phylogenetic hypothesis, the two main clades of apical portion somewhat inflated with rows of black Chauliodinae may have formed by the splitting of Laurasia spinous setae, ventral portion basally with shallow incision from Gondwana. near inner margin; tenth sternum strongly sclerotized and On the basis of the distribution patterns of Asian fishflies, subtriangular, much shorter than ninth tergum, mostly by harbouring many relatively primitive genera and species, enveloped by ninth tergum, curved upwards. Female south-western China and north-eastern India are considered eighth sternum subtrapezoidal, usually produced apically to be the origin centre of the recent fauna of Chauliodinae as a subgenital plate; ninth tergum deep and pointed in Asia. The speciation may have been prompted in the downwards; tenth tergum shortly claviform with upper diversified mountainous habitats formed by the collision of apical corner bluntly produced; gonocoxite subtrapezoidal the Indian subcontinent and Laurasia. with pointed tips.
Remarks. The genus Sinochauliodes comprises four spe- Biogeography of Parachauliodes and Sinochauliodes cies, with two species described as new to science. The new genus appears to be closely related to the genus Para- The Western Pacific Island Arc mainly comprises Taiwan, chauliodes in having similar venation and genitalic struc- Ryukyus and Japan, which may have been formed after the tures, but can be easily separated from Parachauliodes by Cenozoic. Taiwan has been connected to the Chinese the sexual dimorphism in the antenna, the elongated male mainland several times since the mid-Pleistocene, offering ninth tergum and the strongly produced tenth sternum in some direct pathways to introduce species of the mainland the male. In Parachauliodes, the distinct sexual dimorphism into this newly elevated land (Zhang, 1999). Judging from in the antennae is absent, the male ninth tergum is not the distribution patterns of Parachauliodes and Sinochauli- elongated and the male tenth sternum is obtuse at the tip. odes, the precursor of Parachauliodes probably was first Furthermore, the reduced male ninth gonocoxite is roundly introduced to Taiwan by way of south-eastern China, then produced in Parachauliodes, whereas, in Sinochauliodes, this extending northwards along the Western Pacific Island Arc feature is indistinct. to Japan. The division of the two sister genera may have taken place as a result of the vicariance between Taiwan and the mainland of Asia during the frequent transgression of Key to the adults of Sinochauliodes the Pacific after the mid-Pleistocene (Fig. 3). 1. Body and wings almost blackish brown throughout ...... S. fujianensis (Yang & Yang), comb.n. Taxonomy Body and wings much paler, not blackish brown throughout...... 2 Genus Sinochauliodes Liu & Yang, gen.n. 2. Wings without any distinct brownish marks; tenth sternum with tip truncate ...... Etymology. The new genus is named after the unique ...... S. griseus (Yang & Yang), comb.n. distribution in China. Wings with distinct brownish marks; tenth sternum with tip strongly produced...... 3 Type species. Sinochauliodes squalidus sp.n. 3. Wings with dense small brownish spots; male tenth tergum with rather oblique proximal margin in lateral General characters. Fishflies of median size (20–45 mm), view; tenth sternum narrowing at apical portion with tip with brown to black body. Wings hyaline with dark marks slightly narrowing again ...... S. maculosus sp.n. or entirely blackish brown. Head subtriangular and robust Wings with several brownish cloudy marks, but with distinctly prominent compound eyes. Antennae pec- with small spots indistinct; male tenth tergum with tinate in male and subserrate in female. Labrum suboval. vertical proximal margin in lateral view; tenth Prothorax subcylindrical, not longer than wide. Meso- and sternum narrowing at apical portion but with tip not metathorax distinctly robust. Legs densely setose, with narrowing again ...... S. squalidus sp.n. some long setae. Wings somewhat narrow and elongated, about 4.0 longer than wide, with round or slightly curved Sinochauliodes fujianensis (Yang & Yang, 1999), comb.n. tips; Rs four-branched, usually with R2 two-branched, R3 (Figs 4A, B; 5) and R4 usually with Z-bend near apices in male; three crossveins between R1 and Rs; M two-branched; 1A two- Neochauliodes fujianensis Yang & Yang, 1999: 174. branched, with posterior branch strongly sinuated; 2A two-branched, with both branches strongly sinuated. Male Diagnosis. Body and wings almost blackish brown; male ninth tergum subquadrate, much longer than wide, usually ninth tergum with oblique apical margin.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
664 X. Liu and D. Yang
Fig. 5. Sinochauliodes fujianensis.A, wings; B, male genitalia, lateral view; C, tenth sternum, ventral view; D, male geni- talia, dorsal view; E, male genitalia, ventral view; F, female genitalia, lateral view. Scale bars: A, 5 mm; B–F, 1 mm.
Description. Male: body length 23–31 mm; forewing four-branched, R2 distally three-branched, R3 and R4 dis- length 28–30 mm; hindwing length 26–27 mm. tinctly with Z-bend near apices; three crossveins between R1 Head yellowish brown, with frons, vertex and posterolat- and Rs; M two-branched; 1A two-branched, with posterior eral margin entirely black. Compound eyes brown, ocelli branch strongly sinuated; 2A two-branched, with both yellowish brown with black inner margin. Antenna black. branches strongly sinuated. Mouthparts blackish brown. Abdomen blackish brown. Ninth tergum (Fig. 5B, D, E) in Thorax entirely blackish brown. Legs brown with dense, lateral view subquadrate, dorsal posterior corner strongly and yellowish setae. Wings narrowly elongated, entirely blackish acutely produced, ventral posterior corner bluntly produced. brown with tips somewhat paler. Veins blackish brown. Rs Ninth sternum semicircular, distally with a small membranous
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 665
Fig. 6. Sinochauliodes griseus. A, wings; B, male genitalia, lateral view; C, tenth ster- num, ventral view; D, male genitalia, dor- sal view; E, male genitalia, ventral view; F, female genitalia, lateral view. Scale bars: A, 5 mm; B–F, 1 mm.
triangular process. Tenth tergum laterally slightly flattened, in Material examined. Holotype #: CHINA, Fujian, Jianyang, lateral view distal half indistinctly narrowing with round distal Huangkeng, 278349N, 1178399E, 10.v.1944 (Xiufu Zhao)(CAU). margin; in ventral view middle portion with a shallow Allotype $: CHINA, Fujian, Jianyang, Dazhulan, 278349N, transverse incision on inner margin, and distal half feebly 1178399E, 15.v.1945 (Xiufu Zhao) (CAU). Paratypes: CHINA, inflated. Tenth sternum (Fig. 5B, C, E) strongly sclerotized, Fujian, 1#: Jianyang, Huangkeng, 278349N, 1178399E, 9.v.1944 curved dorsad, as long as two-thirds of ninth tergum; (Xiufu Zhao)(CAU);1#: Jianyang, Dazhulan, 278349N, proximal margin widely and deeply incised arched; distal half 1178399E, 10.vi.1948 (Xiufu Zhao) (CAU). Other material: slightly narrowing with tip strongly produced. CHINA, 1#, Fujian, Jianyang, Huangkeng, 278349N, Female: body length 42 mm; forewing length 37 mm; 1178399E, 30.iv.2004 (Xingyue Liu)(CAU). hindwing length 31 mm. Colour similar to male. Eighth sternum broad, posterior margin strongly pro- Distribution. China (Fujian). duced. Tenth tergum (Fig. 5F) shortly claviform, slightly narrowing and feebly incised at tip. Gonocoxite (Fig. 5F) Remarks. This species is the most distinctive species of subtrapezoidal with pointed tip, posteriorly produced. Sinochauliodes in having a blackish body and wings. All the
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
666 X. Liu and D. Yang
adults were collected only on Mt. Wuyishan of Fujian vertically straight; tenth sternum slightly widened and Province, which is located in south-eastern China. During truncate at tip. our survey to Mt. Wuyishan in 2004, we found that the adults were very active on sunny days, normally gliding high Description. Male: body length 28–38 mm; forewing above the bamboo forest and sometimes nosing down length 37–48 mm; hindwing length 33–42 mm. towards an open space within the forest. Head yellowish brown, with frons, vertex and posterolat- eral margins entirely blackish brown. Compound eyes Sinochauliodes griseus (Yang & Yang, 1999), comb.n. brown, ocelli yellow with black inner margin. Antenna (Figs 4C, D; 6) blackish brown. Mouthparts pale yellowish brown. Thorax pale brown. Legs yellowish brown with dense, pale Neochauliodes griseus Yang & Yang, 1999: 173. yellow setae; tibiae and tarsi blackish brown, tarsal claws reddish brown. Wings narrowly elongated, pale greyish brown Diagnosis. Wings almost pale greyish brown without without distinct marks. Veins brown. Rs four-branched, R2 distinct marks; male ninth tergum with apical margin nearly distally three-branched, R3 and R4 distinctly with Z-bend near
Fig. 7. Sinochauliodes maculosus.A, wings; B, male genitalia, lateral view; C, tenth sternum, ventral view; D, male gen- italia, dorsal view; E, male genitalia, ven- tral view. Scale bars: A, 5 mm; B–E, 1 mm.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 667
Fig. 8. Sinochauliodes squalidus. A, wings; B, male genitalia, lateral view; C, tenth sternum, ventral view; D, male genitalia, dorsal view; E, male genitalia, ventral view; F, female genitalia, lateral view. Scale bars: A, 5 mm; B–F, 1 mm.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
668 X. Liu and D. Yang
apices; three crossveins between R1 and Rs; M two-branched; Etymology. The specific epithet ‘maculosus’ refers to the 1A two-branched, with posterior branch strongly sinuated; forewings with a large number of small brownish spots. 2A two-branched, with both branches strongly sinuated. Abdomen brown. Ninth tergum (Fig. 6B, D, E) in lateral Diagnosis. Forewings with a large number of small view subquadrate, posterior margin nearly vertical, with brownish spots; male tenth tergum with rather oblique round ventral posterior corner. Ninth sternum semicircular proximal margin in lateral view; distal half of tenth sternum distally with small membranous triangular process. Tenth strongly narrowing with blunt narrowed tip. tergum laterally flattened, in lateral view distal half dis- tinctly narrowing with round posterior margin; in ventral Description. Male: body length 40–45 mm; forewing view middle portion with shallow longitudinal incision on length 40–43 mm; hindwing length 36–39 mm. inner margin, and distal half inflated, globose. Tenth Head yellowish brown, with blackish marks on ocellar sternum (Fig. 6B, C, E) strongly sclerotized, curved dorsad, triangle area and radiating towards vertex. Compound eyes as long as two-thirds of ninth tergum; in lateral view its tip brown, ocelli pale yellow with black inner margin. Antenna acutely pointed; in ventral view its proximal margin widely blackish brown. Mouthparts yellowish brown, with distal incised trapezoidal; distal half slightly narrowing, but tip half of mandibles, maxillary palpi and labial palpi brown. somewhat widened and truncate. Prothorax greyish brown, middle portion dorsally with Female: body length 38–43 mm; forewing length 41–50 a pair of pale yellow marks near lateral margin; meso- and mm; hindwing length 38–45 mm. Colour similar to male. metathorax yellowish brown, dorsally with brownish marks on Eighth sternum subtrapezoidal with posterior margin dis- lateral sides. Legs yellow with dense, brownish setae; tibiae and tinctly produced. Tenth tergum (Fig. 6F) short claviform, tarsi brown, tarsal claws reddish brown. Wings hyaline with ventrally with tip slightly incised. Gonocoxite (Fig. 6F) many small brownish spots; pterostigma indistinct. Forewing broad with tip distinctly narrowing and curved inwards. tinged with brown at basal costal area, and with 7–8 small blackish brown spots before pterostigma; small brownish spots Material examined. Holotype #: CHINA, Zhejiang, Xi- densely dispersed throughout along longitudinal veins, medi- tianmushan, 308269N, 1198349E, 4.v.1980 (Chikun Yang) ally with one subtriangular hyaline area near Rs; anal area also (CAU). Allotype $: CHINA, same data as holotype (CAU). nearly without any marks. Hindwing mostly hyaline, with Paratypes: CHINA, Zheijang: 1#: same data as for holotype sparse small brownish spots at tip. Veins blackish brown, with (CAU); 2#: Xitianmushan, 308269N, 1198349E, 500–1000 m, most veins on hindwings pale brown. Rs four-branched, R2 4.v.1980 (Fasheng Li)(CAU);1#: Hangzhou, Lingyin, distally two-branched, R3 and R4 distinctly with Z-bend near 308169N, 1208109E, 23.iv.1980 (Chikun Yang) (CAU); 1$: apices; three crossveins between R1 and Rs; M two-branched; Hangzhou, 308169N, 1208109E, v.1983. Other materials: 1A two-branched, with posterior branch strongly sinuated; CHINA, Zheijang: 1#: Anji, Longwangshan, 308239N, 2A two-branched, with both branches strongly sinuated. 1198249E, 490 m, 12/14.vi.1996 (Chikun Yang) (CAU); 2$: Abdomen greyish brown. Ninth tergum (Fig. 7B, D, E) in Anji, Longwangshan, 308239N, 1198249E, 12.v.1996 (Hong lateral view subquadrate, with straight dorsal margin and Wu) (CAU); 1$: Moganshan, 308379N, 1198519E, 7.v.1936 arched ventral margin. Ninth sternum semicircular, distally (IZCAS); 1$: Zhejiang, 31.iv.1929 (IZCAS). with a small membranous triangular process. Tenth tergum laterally flattened, in lateral view with proximal margin rather Distribution. China (Zhejiang). oblique, its distal portion slightly narrowing with round distal margin; in ventral view proximal portion with subsinuate Remarks. This species seems to be closely related to the incision on inner margin, and distal portion slightly inflated. following new species, S. squalidus, in having the male tenth Tenth sternum (Fig. 7B, C, E) strongly sclerotized, curved tergum with a vertical proximal margin in lateral view, but it dorsad, as long as two-thirds of ninth tergum; proximal is easily separated from S. squalidus by the wings lacking margin widely and deeply incised trapezoidal; distal half distinct marks and the tenth sternum with the tip somewhat strongly narrowed, with tip bluntly narrowed again. widened and truncate. In S. squalidus, the wings are tinged Female: unknown. with several brownish cloudy marks and the tenth sternum is strongly tapered distally. Distribution. China (Guangxi, Guizhou). Remarks. The new species is distinguished from other Sinochauliodes maculosus Liu & Yang, sp.n. Sinochauliodes species by the wing pattern of dense brown- (Figs 4D; 7) ish spots. The unique feature of the male genitalia is that the distal portion of the tenth sternum is distinctly narrowed Type specimens. Holotype #: CHINA, Guangxi, Maoer- with the tip slightly narrowed again. shan, Hongjunting, 258539N, 1108259E, 1600 m, 28.vi.2003 (Xingyue Liu) (CAU). Paratypes: CHINA, 2#: same data as Sinochauliodes squalidus Liu & Yang, sp.n. for holotype (CAU); 3#: Guangxi, Maoershan, 258539N, (Figs 4F, G; 8) 1108259E, 2100 m, 5.vii.2003 (Xingyue Liu) (CAU); 1#: Guizhou, Jiangkou, Fanjingshan, 278559N, 1088419E, 800 Type specimens. Holotype #: CHINA, Guangdong, m, 31.v.1988 (Longlong Yang) (IZCAS). Ruyuan, Nanling, 238209N, 1158239E, 7.v.2004 (Mengqing
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
Phylogeny of the subfamily Chauliodinae 669
Wang) (CAU). Paratypes: CHINA, 1# 2$, Guangdong, Distribution. China (Guangdong). Ruyuan, Nanling, 238209N, 1158239E, 950 m, 13/15.v.2001 (Mingyi Tian) (CAU). Remarks. See the remarks on S. griseus.
Etymology. The specific epithet ‘squalidus’ refers to the Acknowledgements cloudy brownish marks on the forewings. We are greatly indebted to Dr Oliver S. Flint, Jr. (Smith- Diagnosis. Forewings with several cloudy brownish sonian Institution, Washington DC, U.S.A.), Dr Fumio marks; male tenth tergum with proximal margin nearly Hayashi (Tokyo Metropolitan University, Tokyo, Japan), vertical; tenth sternum with proximal margin widely and Dr Kazuhiro Masunaga (Lake Biwa Museum, Shiga, Japan) deeply incised V-shaped, and with distal half slightly and Mr Jian Yao (Institute of Zoology, China Academy of narrowing towards tip. Science, Beijing, China) for providing many specimens from their collections. Our thanks also go to Dr Norm D. Penny Description. Male: body length 30–38 mm; forewing (California Academy of Sciences, San Francisco, California, length 36–43 mm; hindwing length 31–39 mm. U.S.A.) for offering many references. We are very grateful Head yellowish brown, with blackish brown marks on to Dr Bradley J. Sinclair (Zoologisches Forschungsinstitut ocellar triangle area and radiating towards vertex. Com- und Museum Alexander Koenig, Bonn, Germany) for pound eyes brown, ocelli pale yellow with black inner evaluating the manuscript before submission. Michael F. margin. Antenna black. Clypeal area brown. Mouthparts Whiting (Brigham Young University, Provo, Utah, U.S.A.) blackish brown. and an anonymous neuropterologist are thanked for pro- Thorax pale brown with meso- and metanotum blackish viding many useful comments on the manuscript. Finally, brown on lateral sides. Legs pale brown with dense, we are much indebted to Professor Chikun Yang and Mr brownish setae; tibiae and tarsi blackish brown, tarsal claws Fasheng Li (Beijing, China) for their kind help in many pale red. Wings hyaline with pale brown cloudy marks; ways. This research was supported by the National Natural pterostigma indistinct. Forewing tinged with brown on Science Foundation of China (30370174, 30225009, basal costal area, and with a brown stripe before pteros- 30200025, 30430100). tigma; small brownish spots densely dispersed along longi- tudinal veins; several brownish cloudy marks located behind R1. Hindwing mostly hyaline, only with a blackish stripe before pterostigma and with sparse small brownish spots on References apical half. Veins pale yellow except blackish brown cross- veins on costal area and brownish marks. Rs four-branched, Asahina, S. (1987) A new corydalid fly from Okinawa Island. Gekkan-Mushi, 201, 17–19 (in Japanese). R distally three-branched, R and R indistinctly with Z- 2 3 4 Asahina, S. (1988) Descriptions of three species of Neochauliodes bend near apices; three crossveins between R1 and Rs; M bowringi group from Hong Kong, the Ryukyus and Thailand two-branched; 1A two-branched, with posterior branch (Megaloptera). Gekkan-Mushi, 207, 6–11 (in Japanese). strongly sinuated; 2A two-branched, with both branches Barnard, K.H. (1931) The Cape alderflies (Neuroptera, Megalop- strongly sinuated. tera). Transactions of the Royal Society of South Africa, 19, Abdomen brown. Ninth tergum (Fig. 8B, D, E) in lateral 169–184. view subquadrate, apical margin nearly vertically straight, Chandler, H.P. (1954) Four new species of dobsonflies from with ventral posterior corner roundly produced. Ninth California (Megaloptera: Corydalidae). Pan-Pacific Entomolo- sternum semicircular, distally with a small membranous gist, 30, 105–111. triangular process. Tenth tergum laterally flattened, in Contreras-Ramos, A. (1995) A remarkable range extension for the fishfly genus Dysmicohermes (Megaloptera: Corydalidae). Ento- lateral view proximal margin nearly vertical, its distal half mological News, 106, 123–126. distinctly narrowing with round distal margin; in ventral Contreras-Ramos, A. (1998) Systematics of the Dobsonfly Genus view middle portion with shallow transverse incision on Corydalus Latreille (Megaloptera: Corydalidae). Thomas Say inner margin, and distal half inflated, globose. Tenth Monographs, Entomological Society of America, Lanham, sternum (Fig. 8B, C, E) strongly sclerotized, curved dorsad, Maryland. as long as two-thirds of ninth tergum; proximal margin Contreras-Ramos, A. (2004) Is the family Corydalidae (Neuro- widely and deeply incised V-shaped; distal half slightly pterida, Megaloptera) a monophylum? Denisia, 13, 135–140. narrowing with round tip. Davis, K.C. (1903) Sialididae of North America. Bulletin of the New Female: body length 40–45 mm; forewing length 36–43 York State Museum, 68, 442–487. mm; hindwing length 31–39 mm. Colour similar to male, but Eriksson, T. (1998) Autodecay, Version 4.0 (program distributed by the author). Bergius Foundation, Royal Swedish Academy of ventral surface of head tinged with blackish brown on Sciences, Stockholm. posterolateral margin. Esben-Petersen, P. (1924) South African Megaloptera. Annals of the Eighth sternum broad, posterior margin slightly pro- South African Museum, 19, 151–158. duced. Tenth tergum (Fig. 8F) short claviform with pointed Evans, E.D. (1984) A new genus and a new species of the dobsonfly tip. Gonocoxite (Fig. 8F) subtrapezoidal with pointed tip, from the far western United States (Megaloptera: Corydalidae). posteriorly produced. Pan-Pacific Entomologist, 60, 1–3.
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670 中国科技论文在线 http://www.paper.edu.cn
670 X. Liu and D. Yang
Flint, O.S. Jr (1983) Nothochauliodes penai, a new genus and species Pictet, F.J. (1854) Traite´ de Pale´ontologie ou Histoire Naturelle des of Megaloptera from Chile (Neuroptera: Corydalidae). Entomo- Animaux Fossiles Considere´s dans Leurs Rapports Zoologiques logical News, 94, 15–17. et Ge´ologiques. Baillie` re, Paris. Glorioso, M.J. (1981) Systematics of the dobsonfly subfamily Ponomarenko, A.G. (1976) Corydalidae (Megaloptera) from the Corydalinae (Megaloptera: Corydalidae). Systematic Entomol- Cretaceous deposits of North Asia. Entomologicheskoe Obozre- ogy, 6, 253–290. nie, 55, 425–433. Hayashi, F. (1989) Neochauliodes sinensis (Walker) in Taiwan. Swofford, D.L. (2002) PAUP* – Phylogenetic Analysis Using Parsi- Nature and Insects, 24, 28–31 (in Japanese). mony (*and Other Methods), Version 4.0b10. Sinauer Associates, Hayashi, F. (1990a) Notes on Neochauliodes sinensis (Walker) Sunderland, Massachusetts. (Megaloptera, Corydalidae) on Amami-Oshima Island, Kagosh- Theischinger, G. (1983) The adults of the Australian Megaloptera. ima Prefecture, Japan. Gekkan-Mushi, 228, 14–15 (in Japanese). Aquatic Insects, 5, 77–98. Hayashi, F. (1990b) Distribution of Parachauliodes yanbaru Asahina. Tillyard, R.J. (1918) Australian Megaloptera or alderflies, descrip- Collecting and Breeding, 52, 488–489 (in Japanese). tions of the new genera and species. Proceedings of the Linnean Hayashi, F. (1996) Insemination through an externally attached Society of New South Wales, 43, 819–825. spermatophore: bundled sperm and post-copulatory mate guard- van der Weele, H.W. (1907) Notizen u¨ ber Sialiden und Beschrei- ing by male fishflies (Megaloptera: Corydalidae). Journal of bung einiger neuen Arten. Notes from the Leyden Museum, 28, Insect Physiology, 42, 859–866. 227–264. Hayashi, F. (1999) Ejaculate production schedule and the degree of van der Weele, H.W. (1909) New genera and species of Megaloptera protandry in fishflies (Megaloptera: Corydalidae). Functional Latr. Notes from the Leyden Museum, 30, 249–264. Ecology, 13, 178–189. van der Weele, H.W. (1910) Megaloptera Monographic Revision. Kimmins, D.E. (1954) A new genus and some new species of the Collections Zoologiques du Baron Edm. de Selys Longchamps, 5, Chauliodini (Megaloptera). Bulletin of the British Museum (Nat- 1–93. ural History), Entomology, 3, 415–444. Wichard, W. (2003) Chauliodes, ein Großflu¨ gler im Baltischen Lestage, J.A. (1927) La fauna entomologique indochinoise, 2: les Bernstein (Megaloptera, Corydalidae). Mitteilungen aus dem Megalopteres. Bulletin et Annales de la Socie´te´ Royale d’Entomo- Geologisch-Pala¨ontologischen Institut der Universita¨t Hamburg, logie de Belgique, 67, 71–90, 93–119. 87, 147–158. Liu, X.Y. & Yang, D. (2005a) Notes on the genus Neochauliodes Yang, C.K. & Yang, D. (1986) New fishflies from Guangxi, China from Guangxi, China (Megaloptera: Corydalidae). Zootaxa, (Megaloptera: Corydalidae). Entomotaxonomia, 8, 85–95 (in 1045, 1–24. Chinese). Liu, X.Y. & Yang, D. (2005b) Notes on the genus Neochauliodes Yang, C.K. & Yang, D. (1990) The fishflies from Hainan Island Weele, 1909 (Megaloptera: Corydalidae) from Henan, China. (Megaloptera: Corydalidae). Acta Zootaxonomica Sinica, 15, Entomological Science, 8, 293–300. 98–100 (in Chinese). Maddison, D.R. & Maddision, W.P. (2000) MACCLADE, Version Yang, C.K. & Yang, D. (1992) Megaloptera of Mount Mogan with 4.0. Sinauer Associates, Sunderland, Massachusetts. one new species. Journal of Zhejiang Forestry College, 9, 414–417 Nava´ s, L. (1913) Espe` ces nouvelles de Ne` vropte` res exotiques. (in Chinese). Extrait des Annales l’Association des Naturalistes de Levallois- Yang, C.K. & Yang, D. (1999) Megaloptera, Corydalidae. Fauna of Perret, 19, 10–13. Insects in Fujian Province of China, Vol. 3 (ed. by B.K. Huang), Nava´ s, L. (1924) Insecta Orientalia. III. Series. Memorie della pp. 168–176. Fujian Science Press, Fuzhou (in Chinese). Accademia Pontifica delle Scienze Nuovi Lincei, Rome, 7, Yang, D. & Yang, C.K. (1992) Megaloptera: Corydalidae. Insects 217–228. of Hengduan Mountains Region, Vol. 1 (ed. by S.X. Chen), pp. Nava´ s, L. (1932) Decadas de insectos nuevos. Brote´ria, 1, 145–155. 435–437. Science Press, Beijing (in Chinese). New, T.R. & Theischinger, G. (1993) Megaloptera (alderflies, Yang, D. & Yang, C.K. (1995) Megaloptera and aquatic Neuro- dobsonflies). Handbuch der Zoologie, IV (33), 1–102. ptera. Aquatic Insects of China Useful for Monitoring Water Paulian, R. (1951) Faune des eaux douces de Madagascar. Plecop- Quality (ed. by L.F. Yang, L.X. Tian & J.C. Morse), pp. te` res et Megalopte` res. Me´moires de l’Institut Scientifique de 250–259. Hohai University Press, Nanjing. Madagascar, Series A, 6, 53–61. Yang, D. & Yang, C.K. (1997) Two new species of the fishflies from Penny, D.E. (1993) The phylogenetic position of Chloroniella south China (Megaloptera: Corydalidae). Journal of China peringueyi (Megaloptera: Corydalidae) and its zoogeographic Agricultural University, 2, 31–32 (in Chinese). significance. Entomological News, 104, 17–30. Zhang, Y.Z. (1999) Zoogeography of China. Science Press, Beijing Penny, D.E. (1999) Madachauliodes ranomafana, a new Madagas- (in Chinese). can species of Chauliodinae (Megaloptera: Corydalidae), with a key to the world genera of Chauliodinae. Entomological News, Accepted 2 March 2006 110, 193–220. First published online 30 August 2006
# 2006 The Authors Journal compilation # 2006 The Royal Entomological Society, Systematic Entomology, 31, 652–670