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