Notes on the genus Chelipodozus Collin (Diptera: : Hemerodromiinae) with descriptions of seven new species from Chile

Adrian R. Plant

The genus Chelipodozus Collin (Diptera: Empididae) is reviewed and Chelipodozus cinereus Collin is designated type species of the genus. Chelipodozus trauco sp. n., C. mapucheensis sp. n., C. luteothorax sp. n., C. chodlipang sp. n., C. chelipodiformis sp. n., C. australis sp. n. & C. araucariana sp. n. are described from Chile and the descriptions of C. cinereus Collin and C. ochraceus Collin are augmented. Definitions of the Chelipodini are discussed and although Chelipodozus is regarded as belonging to this tribe, the laterotergite is bare and interpreted as a reversal of the synapomorphic condition of Chelipodini in which setae are present. Pennate setae are present as secondary sexual characters on the legs of males of the new species. This morphology has not been noted previously in Hemerodromiinae and is interpreted as having an epigamic display function. It is argued that speciation has been driven primarily by mate recognition and sexual selection traits of the antenna and front leg rather than differentiation of genitalic characters. The genus has Andean biogeographic affinities with loci of endemism in the main Cordillera, Valdivian ranges and Magellanic sector. Adrian R. Plant, Department of Biodiversity & Systematic Biology, National Museum of Wales, Cardiff, CF10 3NP, United Kingdom. [email protected]

Introduction CNC Canadian National Collection of , The genus Chelipodozus Collin, 1933 (Diptera: Ottawa, Ontario, Canada Empididae: Hemerodromiinae) was erected by EMEC Essig Museum of Entomology, University Collin (1933) for two species both described from of California, Berkeley, USA single specimens collected in Región de Los Lagos NMWC National Museum Wales, Cardiff, Great in central southern Chile; C. cinereus Collin from Britain a male, and C. ochraceus Collin from a female. The MNNC Museo Nacional de Historia Natural, present work supplements the descriptions of these Santiago, Chile two species and describes seven more, all collected in UMO Oxford University Museum of Natural the cool temperate rainforests of southern Chile. History, Oxford, Great Britain Collection locality abbreviations are coded to indi- cate the Chilean administrative regions, numbered Materials and methods from North to South using Roman numerals I to The following abbreviations refer to institutions XII: Regions specifically mentioned are: housing specimens: Region viii Región del Biobio BMNH Natural History Museum, London, United Region ix Región de La Araucania Kingdom Region x Región de Los Lagos

Tijdschrift voor Entomologie 151: 103–114, Figs 1–24. [ISSN 0040–7496]. http://www.nev.nl/tve © 2008 Nederlandse Entomologische Vereniging. Published 1 June 2008. 104 Tijdschrift voor Entomologie, volume 151, 2008

Region xi Región de Aisén del General Carlos twice the length of the postpedicel. However, the va- Ibáñez del Campo lidity of this character is doubtful as the present work Region xii Región de Magallanes y Antártica and studies of the closely related genus in Chilena. the Australasian region (Plant 2007) have revealed a Morphological terms are essentially those of wide range of aristal morphology in the Chelipodini, McAlpine (1981) and Stuckenberg (1999). Inter- the arista varying from very long to completely ab- pretation of genitalic homology follows Cumming sent. Reduction in length of arista is often associated et al. (1995) and Sinclair (2000). Thoracic setae are with a concomitant trend toward pronounced elon- abbreviated as – dc, dorsocentral; h, humeral (post- gation of the postpedicel. In Chelipodozus, the later- pronotal); lnp, lower notopleural; ocl, ocellar; ph, otergite is bare and is here interpreted as a reversal posthumeral; sa, supraalar; sct, scutellar; unp, upper of the otherwise synapomorphic condition of the notopleural; vt, vertical. Orientation is denoted by Chelipodini in which the laterotergite bears distinct – av, anteroventral; pd, posterodorsal, pv, posterov- setae. It is clear from the foregoing that tribal defini- entral. C1, C2 and C3 refer to the front, mid and tions of Hemerodromiinae are imperfect and will not hind coxae respectively; F1–F3 and T1–T3 to the be resolved without wider study and phylogenetic corresponding femora and tibiae. The front femora analysis of the world fauna of Hemerodromiinae. bear two rows of long setae ventrally between which Chelipodozus can readily be recognized amongst is a double row of much shorter peg-like setae. This other Hemerodromiinae with a short, arched thorax study employs the term spine to describe setae of bearing strong setae on the scutum by the following the outer rows and denticle denotes shorter setae combination of characters – laterotergite bare; wing between these rows. The femoral formula (Plant (Fig. 23) with vein R4+5 forked; cell cup broad, 2007) records the number of spines or denticles in almost as long as cell bm, apically rounded with vein each series starting from the most anterior and work- CuA2 only slightly recurrent basally and becoming ing posteriorly. Thus a femoral formula of 5/10/12/6 distinctly recurrent and rounded distally where the indicates that there are 5 av spines, 10 av denticles, outer angle between it and vein A1 is distinctly acute 12 pv denticles and 6 pv spines. (clearly less than 90º). Confusion is most likely with Chelipoda which nor- mally has vein R4+5 unforked, the laterotergite bears Systematic account distinct setae and cell cup is narrower, rather more Genus Chelipodozus Collin quadrate apically (and often slightly produced pos- teroapically). In local species of Chelipoda vein CuA2 Chelipodozus Collin, 1933: 279. Collin (1933) did not is not recurrent basally (e.g. in Chelipoda remissa designate a type species and the subsequent citation Collin, 1933, Fig. 24) and is more evenly curved or by Smith (1967) – ‘Type species, cinereus Collin (orig. sublinear throughout its length and the outer angle des.)’ appears to be in error. Chelipodozus cinereus Collin is here designated type species. between it and vein A1 is more obtuse (usually 90º or more). In Chelipodozus the female eighth abdomi- The tribe Chelipodini, includes Afrodromia Smith, nal segment is tubular and telescopic and the cerci 1969; Chelipoda Macquart, 1823; Drymodro- are exceptionally long although in C. chelipodiformis mia Becker, 1914; Monodromia Collin, 1928; sp. n., both are shorter and more resemble local spe- Zetterstedt, 1837; Ptilophyllodromia, cies of Chelipoda. Bezzi, 1904 and Chelipodozus Collin, 1933 (Sinclair Chelipodozus may be recognized using the key to & Cumming 2006). Cephalodromia pictipennis (Bezzi, Chilean Hemerodromiinae in Collin (1933: 10) 1912) from Taiwan is undoubtedly a Chelipoda-like with the following minor modification of couplets taxon, properly included in the Chelipodini (Plant four and five: 2007) but which appears to have been incorrectly 4(5) Anal cell as long as second basal cell . . . . . assigned to Cephalodromia Becker, 1914 (Mythico- ...... Chelipodozus myiidae) (Sinclair & Cumming 2006). Chelipodini 5(4) Anal cell shorter than second basal cell. are distinguished from other Hemerodromiinae by the presence of a relatively short and distinctly Key to the Chilean species of Chelipodozus arched thorax with strong setae on the scutum and The following key includes only males of Chilean with male terminalia partially reflexed anteriorly over Chelipodozus. Females are not included as females the abdomen (MacDonald 1988, Plant 1993, 2005, of C. mapucheensis sp. n., C. luteothorax sp. n. and Sinclair & Cumming 2006). These workers have of- C. chodlipang sp. n. are as yet unknown and few de- ten separated Chelipodini from Hemerodromiini us- finitive characters have been identified to confidently ing the additional character of a long arista more than separate females of most of the yellow species.