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Revision and Phylogenetic Systematics of the Neotropical Ceratomerinae (Insecta: Diptera: Empidoidea: Brachystomatidae)

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Revision and Phylogenetic Systematics of the Neotropical Ceratomerinae (Insecta: Diptera: Empidoidea: Brachystomatidae) Arthropod Systematics & Phylogeny 197 68 (2) 197 – 228 © Museum für Tierkunde Dresden, eISSN 1864-8312, 22.06.2010 Revision and phylogenetic systematics of the Neotropical Ceratomerinae (Insecta: Diptera: Empidoidea: Brachystomatidae) BRADLEY J. SINCLAIR Canadian National Collection of Insects, Ottawa Plant Laboratory – Entomology, CFIA, K.W. Neatby Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6 [[email protected]] Received 11.i.2010, accepted 13.iv.2010. Published online at www.arthropod-systematics.de on 22.06.2010. > Abstract Thirteen Neotropical species of Ceratomerus, including nine new species (C. apterus, C. argutus, C. comarapa, C. hibbsi, C. irramus, C. longicornis, C. masneri, C. paraconnexus, C. penai) are described and illustrated. One of these species, C. apterus, from Ecuadorian páramo above 4000 m, lacks wings and has reduced halteres. A preliminary phylogeny of the genera and species groups of the Ceratomerinae is presented, with a discussion of the generic concept of Ceratomerus Philippi. A provisional biogeographic hypothesis of this Gondwanan lineage is discussed. Signifi cant relationships include the Southern Gondwana Pattern, the Inverted Southern Pattern and the Neotropical C. paradoxus group evolved within the Australian C. campbelli group, illustrating both an intra-generic and intercontinental pattern. > Key words Empidoidea, Brachystomatidae, Ceratomerus, Neotropics, Chile, Ecuador, Argentina, Bolivia. 1. Introduction The family Brachystomatidae is a recently recognized PHILIPPI (1865), which possesses remarkably modi- worldwide family, classifi ed into three subfamilies: fi ed male legs. COLLIN (1928) fi rst erected the sub- Brachystomatinae, Ceratomerinae and Trichopezinae family in recognition of the distinctive conus, a (SINCLAIR & CUMMING 2006). This family was recog- fi nger-like projection from the pedicel (2nd antennal nized for empidoid taxa with modifi ed female termi- segment) projecting into the postpedicel (3rd anten- nalia: female tergum 7 often with fringe of setae, and nal segment), and absence of an anal cell in the wing female cercus held upright. There are currently 21 (except in the New Zealand genus Glyphidopeza). recognized genera and some 163 described species Two additional Chilean species were described by prior to this study (YANG et al. 2007). As in other em- COLLIN (1933) and one species by PLANT (1995). pidoid families, adults range in size from large and Ceratomerinae is confi ned to the Southern Hemi- slender species of Heterophlebus Philippi, 1865 (10 sphere and in addition to South America, is known mm) to Glyphidopeza Sinclair, 1997 (2 mm). Species from Australia, New Zealand and Norfolk Is. The occur in a variety of habitats including arctic tundra historical classifi cation of the subfamily was more (Heleodromia Haliday, 1833), Chilean rainforests thoroughly reviewed in SINCLAIR (2003a). Currently (Heterophlebus), tropical rainforests (Apalocnemis there are three genera assigned to Ceratomerinae, Philippi, 1865) and cold temperate streams (Cer- Glyphidopeza (2 spp.), Icasma Collin, 1928 (7 spp.) atomerus Philippi, 1865). and Ceratomerus (36 spp.). The former two genera The ceratomerine group was fi rst recognized are endemic to New Zealand, with Glyphidopeza from the Neotropical Region with the description confi ned to small cascading streams and Icasma of the type species, Ceratomerus paradoxus by occurring in humid forests, often near streams or 198 SINCLAIR: Neotropical Ceratomerinae 1 2 3 4 Figs. 1 – 4. Habitus photographs of dried mounted species of Ceratomerus, lateral view. 1: C. masneri, male. 2: C. mediocris, male. 3: C. apterus, male. 4: C. paradoxus, male. (scale bar: 1.0 mm) small pools (SINCLAIR 1997). Ceratomerus appears and New Zealand faunas (SINCLAIR 1997, 2003a). The to occur both in streams and humid forests (SINCLAIR genus Ceratomerus exhibits a classic Gondwanan or 2003a). Trans-Antarctic pattern and this study attempts a fi rst This is the fi rst revision of the Neotropical species analysis of the biogeographic and phylogenetic rela- of Ceratomerinae based on a large number of speci- tionships among species groups of Ceratomerus and mens; it builds on earlier studies of the Australian genera of Ceratomerinae. Arthropod Systematics & Phylogeny 68 (2) 199 2. Material and methods sion 4.0b10 (SWOFFORD 2002). A heuristic search with stepwise addition was implemented to fi nd the most parsimonious trees using random addition sequence of 2.1. Material taxa, tree-bisection-reconnection (TBR) branch swap- ping and 1000 random replications. A posteriori char- This study is based on some 630 adult specimens of acter weighting was implemented using successive Ceratomerinae borrowed from the following institu- approximations according to the rescaled consistency tions: Biosystematics Laboratory, Kyushu University, index (RC). Character evolution, character state distri- Fukuoka, Japan (BLKU); The Natural History Mu- butions and alternative tree topologies were examined seum, London, England (BMNH); Bernice P. Bishop using the program MacClade 4 (MADDISON & MADDI- Museum, Honolulu, USA (BPBM); Canadian Na- SON 2003). The dataset may be downloaded from the tional Collection of Insects, Ottawa, Canada (CNC); TreeBASE database (URL: http://purl.org/phylo/tree- University of Guelph, Guelph, Canada (DEBG); Ki- base/phylows/study/TB2:S10399). takyushu Museum of Natural History, Kitakyushu, Ja- pan (KMNH); Museum National d’Histoire Naturelle, Paris, France (MNHN); National Museum of Wales, 2.3. Morphological abbreviations Cardiff, Wales, U.K. (NMWC); United States Na- tional Museum of Natural History, Washington, USA The following abbreviations are used in the species (USNM); Zoological Museum, University of Copen- descriptions, material examined sections, and fi gures: hagen, Denmark (ZMUC). Abbreviations given here are used throughout the text to indicate deposition of ad anterodoral (position of setae/bristles) specimens. alc alcohol All dissections were made in glycerin and tissues av anteroventral (position of setae/bristles) cleared using hot 85% lactic acid. Terms used for adult cerc cercus CALPINE (1981), except structures primarily follow M d discal cell for the antenna and the wing venation, where the terms dc dorsocentral bristles/setulae of STUCKENBERG (1999) and SAIGUSA (2006) are used, respectively. Homologies of the male terminalia fol- ej apod ejaculatory apodeme epand epandrium low those of SINCLAIR & CUMMING (2006). Lists of scutal setae in descriptions refer to one-side only, ex- FIT fl ight intercept trap cept scutellum. for. forest Label data for primary types are exactly as they ap- gcx apod gonocoxal apodeme pear. Labels are listed from the top downward, with hypd hypandrium data from each label enclosed in single quotation hypd proc hypandrial process marks; lines are delimited by a ‘pipe’ (|). Additional hyprt hypoproct information is included in [square] brackets. M1,2,4 medial vein MT Malaise trap 2.2. Cladistic analysis NP national park npl notopleural bristles Characters were scored for all known genera and re- pal postalar bristle presentative species groups of the Ceratomerinae and pd posterodorsal (position of setae/bristles) three outgroup genera, two from the Trichopezinae pgt postgonite (Brachystomatidae) and Oreogeton Schiner, 1860 ph phallus (Empidoidea, incertae sedis). A complete list of exem- pprn postpronotal bristle plars is provided in Tab. 1. Forty-six characters were presut spal presutural supra-alar bristles analyzed in the cladistic analysis, including four mul- psut spal postsutural supra-alar bristles ti-state characters (Tab. 2). All characters were treated PT pan traps as unordered with multi-state characters considered as pv posteroventral (position of setae/bristles) non-additive, and all characters were equally weight- R radial vein ed. Character polarity was determined by rooting the 1,2 + 3,4,5 tree with the three empidoid outgroups, which togeth- Sc subcostal vein er were constrained to be paraphyletic in relation to sctl scutellar bristles the ingroup. sur surstylus Parsimony analysis of the character state matrix T8,10 tergum (Tab. 3) was performed using the program PAUP* ver- ypans yellow pan traps 200 SINCLAIR: Neotropical Ceratomerinae 3. Taxonomy wide; apex of stylus concolorous with postpedicel. Base of labrum lacking dorsal process; palpus short, pale and slender, 0.2 × length of labrum, with several 3.1. Genus Ceratomerus Philippi long, dark setae. Due to wingless condition, thorax shortened. Me- Ceratomerus Philippi, 1865: 765. Type-species Ceratomerus sonotum, postnotum and pleura dark. Acrostichals paradoxus Philippi (monotypy). lacking; 2 dc, with several setulae interspersed; 1 pprn; Tomia Paramonov, 1961: 100 nec MARTYNOV (1936: 1262). 1 – 2 short presut spal; 1 npl; 0 psut spal; 1 pal; 2 sctl; Type-species Tomia campbelli Paramonov (original de - lacking setulae scattered on scutum. Antepronotum signation). with pair of short setulae; laterotergite bare. Wing reduced to mere vestige. Halter reduced to Diagnosis. Ceratomerus is distinguished by the fol- short, pale, knob-like process, bearing long apical seta. lowing combination of characters: posterior ocelli usu- Fore coxa much paler than pleura, remaining coxae ally widely separated with ocellar bristles inserted an- slightly paler than pleura; fore and mid femora with teriorly, postpedicel lengthened and tapered apically, light anterior surface and dark brown posterodorsal acrostichal setulae present, 2 – 5 dorsocentral
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