Notes on the Genus Chelipodozus Collin (Diptera: Empididae: Hemerodromiinae) with Descriptions of Seven New Species from Chile

Notes on the genus Chelipodozus Collin (Diptera: Empididae: 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 Insects, 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 indicate 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

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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 Chelipoda 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 posteroapically). 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- Phyllodromia 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.

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Chelipodozus araucariana sp. n. 1. Thorax predominantly brown or blackish, Fig. 13 T1 with simple setae above (Fig. 19) . . . . . 2 – Thorax predominantly yellowish, T1 with Type material. Holotype , Chile, Region x, Cau- dorsal fringe of pennate setae (Fig. 18) . . . . 3 tin, 1150 m, Conguillo Nat. Park, Araucaria / 2. C1 almost as long as thorax; postpedicel Nothofagus, 4.ii.1988, L. Masner Chile Exp. (CNC). elongate lanceolate (Fig. 7), much longer Paratypes; 47, 120, as holotype, 4.ii.1099 and than wide, arista 0.3 as long . . . C. cinereus 5.ii.1988 (CNC, EMEC, MNNC, NMWC). – C1 less than 0.7 as long as thorax; postpedicel hardly longer than wide, arista 2.5 Derivatio nominis as long (Fig. 16) ...... C. chelipodiformis From Araucaria, a genus of tree present at the type 3. Postpedicel of antenna elongated, (at least locality. 10 as long as wide) often swollen distally (Figs 9, 11, 13, 14, 15), arista absent or Diagnosis greatly reduced (equal or less than 0.2 as A predominantly yellow species with a strongly long) ...... 4 marked median stripe on scutum continued over – Postpedicel of antenna shorter (about 6.5 scutellum and metanotum. In males, T1 is some- as long as wide), apically pointed with dis- what darkened only anteriorly and dorsally on the tinct arista at least 0.2 as long (Fig. 8) . . . 7 apical third. The postpedicel is distinctively elon- 4. Thorax with distinct dark median dorsal gate, slightly but distinctly narrowed medially stripe continuing over scutellum and metan- (but not obviously constricted as in C. trauco and otum ...... C. araucariana C. mapucheensis ) with a rather bulbous apex bearing – Thorax without median stripe or median a distinct short arista stripe very indistinct, if distinct then not continuing over scutellum and metanotum . . . Description ...... 5 Male. Length: body + head 3.0 mm, wing 3.5 mm. 5. Thorax with median stripe absent or very Similar to C. trauco sp. n. but differing as follows. faint; postpedicel at most weakly narrowed Head with lower postocular ciliation shorter, rather medially, distinct arista 0.2 as long as post- stout. Postpedicel of antenna (Fig. 13) greatly elon- pedicel (Fig. 15) ...... C. luteothorax gated, 12 as long as narrowest, rather flattened, – Thorax with median stripe usually present gradually narrowed medially, broadened gradu- on scutum, not continuing onto scutellum ally distally (not abruptly flattened and widened or metanotum; postpedicel distinctly nar- near tip), arista about 0.2 as long. Thorax with rowed medially, arista vestigial or absent strong brownish black dorsal median stripe as wide (Figs 9, 11) ...... 6 as frons in front of ocelli on front of scutum, some- 6. Lower occipital setae strong; T1 gradually times becoming narrower in prescutellar depression, darkened from base to apex; postpedicel then broader and more diffuse on scutellum and more pointed apically (Fig. 9) . . . . . C. trauco metanotum. Legs yellow with apical tarsal segments – Lower occipital setae weaker; T1 with distal obscurely darkened; T1 darkened dorsally and ante- 0.5 intensely black, contrasting with yellow- riorly on distal 0.3, posteroapically yellowish or only ish basal 0.5; postpedicel more rounded api- slightly darkened; T3 sometimes slightly darkened cally (Fig. 11) ...... C. mapucheensis subapically. F1 with av denticles less numerous, fem- 7. T1 hardly blackish apically with somewhat oral formula 5–6/13–14/11–12/5–7. Male genitalia narrow pennate setae commencing 0.3 from very similar to and doubtfully distinguishable from base positioned rather posterodorsally but C. trauco; apical ‘tooth’ of cercus conspicuous; epan- becoming dorsally placed on distal 0.5, long- drium with longer setae, rather evenly distributed est at 0.6, becoming shorter distally . . . . . near posterior margin...... C. chodlipang Female. Length: body + head 3.0–3.5 mm, wing – T1 distinctly darkened apically, pennate se- 3.5 mm. Very similar to male, differing chiefly as tae positioned distinctly posterodorsally at follows. Antenna shorter, postpedicel lanceolate, commencement of series 0.3 from base, be- narrowly pointed, similar to C. cinereus (Fig. 7) and coming longer and gradually more dorsally C. chodlipang (Fig. 8) but arista almost as long. T1 positioned distally and truly dorsal at apex . not darkened dorsally and anteriorly on distal 0.3 ...... C. australis and with no short hairs dorsally. Cercus very long.

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1 2 Figs 1–5. Male genitalia of Chelipodozus. – 1. C. mapucheensis terminalia in lateral view; 2. C. trauco, cercus; 3. C. australis, cercus; cer 4. C. chodlipang cercus; 5. C. mapucheensis terminalia epan in dorsal view. Legend: cer, cercus; epan, epandrium; hyp, hypandrium. 3

hyp

epan

hyp 5 4

cer

Chelipodozus australis sp. n. Description Figs 3, 10 Male. Length: body + head 3.0 mm, wing 3.5 mm. Type material. Holotype , Chile, Region xii, Head similar to C. trauco but face brownish white Magellanes, Isla Navarinos, Seno Grandi, 55°S, and posterior margin of mouth opening blackish; 6.ii.1959 (BMNH). Paratypes; 1, 1, Isla setae of lower occiput weaker; antenna similar to Navarinos, Nothofagus forest, i.1996, A. R. Plant C. chodlipang (Fig. 8) with postpedicel elongate lan- (NMWC); 1, Isla Bertrand 55°S, 650 feet, ceolate and arista 0.3 as long. Thorax yellow with Nothofagus forest, 4.ii.1959 (BMNH). narrow median blackish brown stripe on scutum, scutellum and metanotum similar to C. chodlipang. Derivatio nominis Legs similar to C. trauco, yellow becoming darker From the Latin, australis, meaning southern. distally with apical and preapical tarsal segments on all legs blackish; pennate setae on T1 blackish, posi- Diagnosis tioned distinctly posterodorsally at commencement A predominantly yellow species with a complete of series 0.3 from base, becoming longer and gradu- median thoracic stripe dorsally, yellowish legs with ally more dorsally positioned distally and truly dorsal distinctly darkened apical tarsi and in males a lan- at apex. Preapical segment of front tarsi not much ceolate postpedicel closely resembling C. chodlipang. longer than wide and slightly shorter than apical seg- Males are distinguished from C. chodlipang primarily ment. Femoral formula 6/12/17/6. Abdomen and by having the distal process of the cercus very strongly male terminalia similar to C. chodlipang but upper curved inward and the series of pennate setae basally margin of cercus (Fig. 3) rather less concave and distal on T1 being distinctly posterodorsal, only gradually process very strongly curved inwards, almost hidden becoming truly dorsal apically. (in contrast, the pen- in lateral view, bearing several minute setae dorsally nate setae on T1 of C. chodlipang are more or less along its length and distinct apical ‘tooth’. Wing with dorsal throughout and only slightly posterodorsal at fork in R4+5 rather long with second submarginal the extreme base of the series). cell (r4) elongate, narrowed basally and somewhat

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ciliation longer, 4–5 serial forming patch of distinct pubescence on lower occiput. Basal antennal segments short, postpedicel not much longer than wide, clothed in short but distinct pale pile; arista 2.5 as long. Proboscis short, about as long as frons; palpus small with pale hairs. Fig. 6. Female terminalia of Chelipodozus trauco. Thorax dark brown thickly covered in greyish white dusting; humerus and lower pleura yellowish; cha- etotaxy similar to C. cinereus and C. trauco, all setae yellowish to brownish. Laterotergite bare. Legs yel- bell-shaped. In the paratype male a spurious cross- low, C1 0.65–0.7 as long as thorax, 3–4 as long vein is present on one wing running between R4 and as wide, series of three or four distinct yellow setae R5 across the base of radial fork. on distal 0.5 becoming shorter on basal 0.5. C2 and Female. Similar to male but antenna (Fig. 10) with C3 with anteroapical yellow setae. F1 1.3–1.4 as scape and pedicel yellow, postpedicel brown, shorter long as C1, 4–5 as long as wide, widest 0.1–0.2 than in male, elongate conical with equally long from base, gradually narrowed distally; ventral spines arista. T1 lacking pennate fringe. Femoral formula and denticles black, small and fine compared with 6/17/19/6 (Isla Navarinos specimen) 6/12/13/6 other members of genus, spines hardly as long as (Isla Bertrand specimen). Abdomen whitish brown, F1 is deep; each series of denticles virtually congru- tergites only weakly pigmented. Cercus very long. ent with and hardly separated from adjacent series of spines; femoral formula 3/14–17/11–16/3–4, Chelipodozus chelipodiformis sp. n. no distinct basal ventral spine. T1 with only short decumbent hairs dorsally. F2, F3, T2 and T3 with Figs 12, 16 short hairs, F2 without pd setae distally. Mid and Type material. Holotype , Chile, Region x, hind legs with metatarsal segment less than 0.6 Cautin, 1150 m, Conguillo Nat. Park, Araucaria / as long as corresponding tibia. Abdomen brown Nothofagus, 4.ii.1988, L. Masner Chile Exp. (CNC). with pale hairs and stronger setae on posterior mar- Paratypes; 101, 45, as holotype, 4.ii.1988 and gin of tergite five. Terminalia (Fig. 12) rather com- 5.ii.1988 (CNC, EMEC, MNNC, NMWC). pact, reflexed anteriorly (Chelipoda-like), brown; Additional material. 2, Osorno, 1250 m, Hypandrium somewhat keel-shaped with short Puyehue N. P., Argentine border, Nothofagus tree setae posteriorly; epandrium broadly triangular, line, 13.ii.1988, L. Masner, Chile Exp. (CNC). weakly sclerotized posteriorly with strong brown Derivatio nominis: from forma (Latin) meaning setae dorsally and posteriorly; cerci not fused with shape, and Chelipoda Macquart, in reference to the epandrium, pale, erect, constricted medially, apical superficial similarity between that genus and the new lobe with a few minute hairs but no apical ‘tooth’. species. Phallus broad, strongly curved anteriorly, blunt- ended; subepandrial process very long, white basally, Diagnosis black distally, strongly twisted and curved anteriorly. A predominantly brownish species which needs com- Wing veins dark, vein A1 not reaching margin, radial parison only with C. cinereus from which it may be fork broad. Haltere white. distinguished by having C1 distinctly shorter than Female. Length: body + head 2.5 mm, wing 3.0 the thorax and a very short postpedicel bearing a mm. Similar to male but postpedicel (Fig. 16) often much longer arista. Like, C. cinereus, males also lack brown and mouthparts brownish yellow. C1 slightly pennate setae on T1. narrower than in male. F1 with ventral spines longer, as long as F1 is deep but very fine. Abdominal cha- Description etotaxy weaker, no strong setae on tergite five; eighth Male. Length: body + head 2.2–2.5 mm, wing 3.0 abdominal segment almost quadrate, not greatly mm. Head black with scape, pedicel and mouthparts elongated; cercus short with longish dark bristles. yellow; postpedicel dusky yellow; face 0.5 as wide as pedicel above, widening and slightly produced in Comment profile below; frons wide, minute pair of frontal setae Chelipodozus chelipodiformis superficially resembles behind antennal sockets; ocellar tubercle prominent, local species of Chelipoda in that the female eighth ocl setae strong, brownish black to yellowish, pro- abdominal segment and cerci are both shorter than clinate; 4–5 pairs brownish vt setae. Upper posto- in other species of Chelipodozus, the thorax is rather cular ciliation uniserial, yellowish; lower postocular shorter and more arched, and the front coxa is of

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7 Figs 7–11. Antenna of Chelipodozus spp. – 7. C. cinereus ; 8. C. chodlipang ; 9. C. trauco ; 10. C. australis ; 8 11. C. mapucheensis .

reduced length. However, the shape of cell cup, Description absence of setae on the laterotergite and fork in vein Male. Length: body + head 3.0 mm, wing 3.5 mm. R4+5, confirm that it is more properly placed in Similar to C. trauco but differing as follows. Face Chelipodozus. more distinctly produced in profile below, vt almost as strong as ocl, setae of lower occiput shorter. Chelipodozus chodlipang sp. n. Postpedicel (Fig. 8) elongate lanceolate but much shorter than C. trauco and with arista 0.5 as long; Figs 4, 8 dorsal seta on scape longer, reaching to pedicel. Type material. Holotype , Chile, Region ix, Volcan Thorax yellow, narrow well defined blackish brown Villarica, 39°23'13.9''S, 71°57'46.7''W, (1346m) median stripe on scutum, scutellum and metanotum. stunted Nothofagus forest with sparse understorey, Legs yellowish becoming darker distally and rather 9.i.2006, A. R. Plant (MNNC). abruptly almost black on apical and preapical tarsal segments of all legs. Faint indications of darkening Derivatio nominis dorsally on F2 and F3 0.8 from base. T1 not black- The specific epithet derives from the Mapuche, chod ish apically, series of black rather narrowly pennate (yellow) & lipang (limb) in reference to the yellow setae commencing 0.3 from base positioned slightly legs of this species. posterodorsally but becoming dorsally placed on distal 0.5, longest at 0.6, becoming shorter distally. Diagnosis Femoral formula 6/18/16/6. Male terminalia very Male only. A predominantly yellow species with a similar to C. mapucheensis but cercus (Fig. 4) with well marked dark median strip across the entire tho- upper margin concave, slightly produced antero- racic dorsum. T1 is yellowish throughout its length dorsally; distal process strongly curved inwards but but the apical tarsal segments on all legs are dark, hardly hidden in lateral view, bearing strong apical distinctly contrasting with basal segments. The ‘tooth’ and two strong setae on outer face of curve. postpedicel, while elongate, is clearly shorter than Wing rather elongate, radial fork rather long such in C. trauco, C. mapucheensis, C. araucariana or that second submarginal cell (r4) distinctly elongate, C. luteothorax and is distinctively narrowly lanceolate narrowed basally and somewhat bell-shaped. with an apical arista half its length. Female. Unknown.

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occiput with uniserial to biserial dark setae becoming 3–4 serial and paler on lower occiput; postpedicel (Fig. 7) elongate lanceolate, arista 0.3 as long; palpus small with pale hairs. Thorax black, paler on lower pleura and almost yellowish on postpronotal lobe and proepisternum; all setae black; three pairs of strong presutural dc, one pair minute prescutellar dc; sa, and unp rather shorter, h and ph distinct but weaker still; one pair of strong sct; laterotergite bare. Legs yellowish becoming brownish distally; mid and hind legs slightly darker with trochanters blackened ventrally; C1 0.9 as long as thorax bearing two to three erect black setae anteriorly in distal 0.5 and one fine posteroapical seta; F1 1.1 as long as C1, 4 as long as wide, considerably inflated, widest 0.4 from base; femoral formula of holotype 6–7/14– 20/13–18/5–6, with strong ventral spine almost at base; T1 0.75 as long as F1; front tarsi with first segment 0.4 as long as T1, segments two and three shorter and equally sized. Mid and posterior legs more slender, F2 lacking pd bristles; posterior meta-  Fig. 12. – Chelipodozus chelipodiformis terminalia in tarsus 0.6 as long as T3, equal length as remaining lateral view. segments. Abdomen blackish brown, slightly paler ventrally; tergites with short hind marginal bristles Chelipodozus cinereus Collin much stronger on tergite 5. The terminalia have been illustrated previously (Sinclair & Cumming Fig. 7, 19 2006: Fig. 287) and are structurally similar to the Chelipodozus cinereus Collin, 1933: 279–280. Holotype species described here but with greater development , Chile, Region x, labelled : Type  / Chelipodozus of hypandrium and postgonites and broader phallus.  cinereus, Coll. Type / Casa Pangue, 4–10.xii.1926 / Wing veins dark and membrane slightly darkened; S. Chile Llanquihue Prov. F. & M. Edwards. B.M. 1927–63 / Holotype  / BMNH(E) # 248009. radial fork rather short, not distinctly narrowed (BMNH) [examined] basally; halteres grey. Female. Similar to male but slightly larger (body + Additional material. Chile: 1, 1, Region x, Oncol, head 3.5mm, wing 4.0mm). Mid and posterior legs 39°41'59''S, 73°19'11''W, (490m), 13.i.2006, A. R. Plant; tending to be darker, particularly outer face of C3. Region X, Reserva Costera Valdiviana, SW of Chaihuin, C1 as long as thorax. Ventral denticles of F1 slightly 40°05'18''S, 73°33'02''W, Fitzroya cupressoides forest at more numerous, femoral formula 6–7/17–24/15– 700m, 15.i.2006, A. R. Plant (NMWC); 5, 10, Re- gion xi, Conquillo, Valdivia, Chiloé Is (Region X), Puerto 18/6–7. Abdomen with only short bristles. Cisnes, 1988 & 1957, L. Masner & L. E. Penn (CNC). Comment A single female from middle Chile, collected by Diagnosis S. Schönemann [UMO], has R4+5 simple (not A predominantly brownish species with C1 almost as forked) in both wings, superficially resembling long as the thorax and an elongate, narrowly lanceo- Chelipoda. However, apart from a rather broad face late postpedicel bearing a distinct arista. Males lack it is otherwise similar to Chelipodozus cinereus and pennate setae on T1. is probably an aberrant species of Chelipodozus. In the absence of an associated male this species is not Re-description described here. Male. Length: body + head 2.5 mm, wing 3.0 mm. Head black, paler on face, scape dirty yellow, pedi- Chelipodozus luteothorax sp. n. cel darker, postpedicel black, mouthparts brown- Fig. 15 ish yellow; face narrow, not as wide as scape is deep, only slightly produced in profile below; ocl Type material. Holotype , Chile, Region x, strong, divergent; two pairs vt almost as strong as Malleco, 14 km E. Malacahuello, 3–31.xii.1982, ocl with smaller seta between and behind; upper 1570 m, A. Newton & M. Thayer (CNC).

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13 Figs 13–16. Antenna of Chelipodozus spp. – 13. C. araucariana ; 14. Probable C. ochraceus  (see text); 15. C. luteothorax ; 16. C. chelipodiformis . 14

Derivatio nominis Chelipodozus mapucheensis sp. n. Figs 1, 5, 11, 21 The specific epithet derives from the Latin, luteus, yellow and refers to the yellow thorax of this Type material. Holotype , Chile, Region x, species. Oncol, 39°41'59''S, 73°19'11''W, (490m), 13.i.2006, A.R. Plant, (MNNC). Paratype 1, as Diagnosis holotype (NMWC). Male only. A predominantly yellow species without a dark median stripe on the thoracic dorsum and Derivatio nominis with T1 somewhat darkened anterodorsally on distal Named in honour of the Mapuche people whose ter- 0.7. The postpedicel is elongate and hardly narrowed ritory coincides with the type locality of this species. medially, conically pointed (not bulbous) apically and bears a short arista Diagnosis Male only. A predominantly yellow species in which Description males are recognized by having T1 abruptly dark- Male. Length: body + head 3.5 mm. Similar to ened on apical half, weak lower occipital setae and C. trauco but differing as follows. Postpedicel a distinctive elongate postpedicel which is narrowed (Fig. 15) rather wide but elongated, superficially medially, abruptly widened distally with a rounded resembling C. araucariana (Fig. 13) but only very apex and no arista. slightly narrowed medially and with distinct arista 0.2 as long as postpedicel. Scutum clear yellow Description with vague impression of narrow darker median Male. Length: body + head 3.0 mm, wing 3.5 mm. stripe when viewed in some lights. Legs yellow with Similar to C. trauco differing as follows. Antenna apical tarsal segments somewhat darkened. T1 yel- (Fig. 11) with scape and pedicel darker brownish low, only anterodorsal face darkened on distal 0.7. yellow, postpedicel rather longer, its flattened broad- Femoral formula 6/16/13/6 and one strong basal ened tip more rounded apically. Front of frons dis- ventral spine. Cercus with distal process bearing 2–3 tinctly yellowish and face whitish yellow. Setae of distinct setae on outer face but otherwise very similar lower occiput much weaker and shorter, less erect. to C. trauco Wing with radial fork rather long and Scutum with faint indications of narrower median narrowed basally, somewhat bell-shaped. stripe. Legs similar to C. trauco but intensely black Female. Unknown. distal 0.5 of T1 (Fig. 21) contrasting abruptly with yellowish basal 0.5; pennate setae rather broader than

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17 18 19

20 21 22

23 24 cup

cup

Figs 17–24. Chelipodozus spp.: habitus (17–19), and details. – 17, C. trauco ; 18, C. trauco ; 19, C. cinereus ; 20, C. trauco , front leg; 21, C. mapucheensis , front leg; 22, C. ochraceus, female head and antennae; 23, C. trauco,  wing; 24, Chelipoda remissa Collin,  base of wing. Legend: cup, cell cup.

C. trauco, intensely black, confined to distal 0.5 and gin more or less linear, not produced; distal process not becoming shorter proximally. Only apical seg- strongly curved inwards and upwards, almost hidden ments of front metatarsi darkened, basal segments in lateral view, 2–3 minute setae distally and minute yellow. Femoral formula 5/18–20/15/5. Abdo- black apical ‘tooth’ dorsally at tip. Postgonites small men paler than C. trauco, tergites dirty yellow to and inconspicuous. Phallus very long and strongly brownish. curved forward; proximal loop emerging below Male terminalia (Figs 1, 5) open, yellow with black epandrium and covered with transparent membrane setae; hypandrium narrow bearing strong black then curving anteriorly behind epandrium with its apical seta; epandrium short oblong quadrate with apex directed forwardly in front of cercus; phallic distinct setae about posterior and dorsal margins. sheath divided and divergent distally. Cercus with some strong setae laterally, upper mar- Female. Unknown.

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Chelipodozus ochraceus Collin forest, 16.i.2006 (EMEC); 1, Reserva Costera Fig. 14, 22 Valdiviana, 39°59'29''S, 73°34'16''W, 15/i/2006 Chelipodozus ochraceus Collin, 1933: 280. Holotype , (MNNC); 1, Chiloé Is., Valdivian forest 0.6km W Chile, Region x, labelled: Chelipodozus ochraceus of Ruta 5 on turn to Miraflores (80m), 40°05'19''S,  Collin type / Puerto Varas 16.xii.1926 / S. Chile: 73°05'19''W (NMWC) all collected A.R. Plant. Llanquihue Prov. F. & M. Edwards. B.M. 1927– 63 / Holotype / BMNH(E) # 248010. (BMNH) [examined). Derivatio nominis The specific epithet refers to the Trauco, a mythi- cal monster from Chiloé Island in reference to the Diagnosis fierce raptorial legs of this species and is a noun in Female only. A predominantly yellow species with- apposition. out a dark dorsal thoracic stripe and with postpedicel elongate conical, narrowly pointed apically with Diagnosis arista equally long. A predominantly yellow species with a narrow dark The holotype female has yellow thorax with scutum median stripe on the scutum. Males are recognized and metanotum obscurely darker but no median by having T1 gradually darkened from base to apex, dark stripe. The head is black with frons distinctly strong lower occipital setae and a distinctive elongate yellowish in front, face whitish and as wide as pedi- postpedicel which is narrowed medially, abruptly cel and produced in profile below. The setae of the widened distally and sharply pointed apically with lower occiput are moderately long. Antennae dark arista absent. with yellowish basal segments (Fig. 22), postpedicel elongate conical, its narrow apex no wider than base Description of arista which is as long as postpedicel. The legs are Male. Length: body + head 4.0 mm, wing 4.2 mm. very similar to C. trauco sp. n. but the anterior bris- Head black with scape, pedicel, face, posterior mar- tles on C1 have smaller hairs between them and the gin of mouth opening and mouthparts yellow; sub- apical three tarsal segments of all legs, F3 & T3 are spherical, slightly dorsoventrally flattened; face as darker. wide as pedicel above, widened slightly and distinctly produced in profile below; frons wide, minute pair Comment of frontal bristles behind antennal sockets; ocellar This species is known with certainty only from the tubercle prominent, ocl setae black, strong, procli- holotype female. However, there are seven females nate, divergent; two pairs yellowish vt weaker than in CNC from Nahuelbuta (Region VIII) Llanqui- ocl, and smaller seta between and slightly behind; hue (Region X) and Aysen (Region XI) which are postocular ciliation yellow, short and biserial on also probably this species. Two males associated with upper occiput becoming much longer and four the Nahuelbuta females are possibly males of C. – serial on lower occiput. Antenna (Fig. 9) with scape ochraceus; they most closely resemble C. mapucheen- 1.5 as long as wide, bearing two short black setae; sis with almost no median stripe on the scutum and pedicel globular with apical circlet of minute setae; T1 intensely black on distal 0.5 contrasting abruptly postpedicel minutely pilose, 2 as long as head, with yellowish basal 0.5. The postpedicel of the swollen basally, narrower medially, abruptly widened Nahuelbuta males is distinctive (Fig. 14), being elon- and flattened near tip; arista absent but minute api- gate lanceolate, narrowing gradually toward tip with cal sensory bristle present. Proboscis sharply pointed, apical arista less than 0.1 as long and the male rather forwardly directed; palpus with fine hairs in genitalia are very similar to and doubtfully distin- front and two stronger (but still fine) apical setae. guishable from C. trauco. Thorax clear yellow, blackish median stripe on scutum not continued onto prescutellar depression. All Chelipodozus trauco sp. n. setae black; three pairs of strong presutural dc and one minute prescutellar dc; h rather short and stout, Figs 2, 6, 9, 17, 18, 20, 23 ph similarly sized; unp and sa strong; lnp minute; Type material. Holotype , Chile: Region x, Cima pair of strong sct. Laterotergite bare. Legs yellow, Oncol, 39°42'09''S, 73°18'32''W. (490–650m), blackish on apical 0.5 of T1 and fore tarsi; T2 and 10–11.i.2006, A. R. Plant (NMWC): Paratypes: T3 apically and tarsal segments of mid and hind 1, as holotype (NMWC); 1, Cima Oncol, legs slightly darkened but apical and preapical tar- 39°41'38''S, 73°18'51''W, (750m), (MNNC); 1, sal segments blackish. C1 1.2 as long as thorax, Reserva Costera Valdiviana, mouth of Rio Colun, 7.5 as long as wide, slightly narrowed apically, five 40°03'31.1''S, 73°39'22.9''W, Aextoxicon punctatum evenly spaced erect setae anteriorly, one yellowish

Downloaded from Brill.com09/26/2021 07:13:43PM via free access Plant: Chelipodozus of Chile 113 anteroapical and one posteroapical seta. F1 as long as impossible to distinguish many yellow group C1, strongly inflated, 4 as long as wide, widest 0.4 females, especially as the females of C. chodlipang, from base; femoral formula 6/17/12/6 with a strong C. luteothorax and C. mapucheensis have yet to be basal ventral spine, av and pv black spines strong, recognized. almost as long as F1 is deep. T1 0.7 as long as Only small numbers of specimens of some taxa were F1, dorsal fringe of black pennate setae (Fig. 20) on available making assessment of the range of varia- distal 0.7 longer than T1 is deep distally, becoming tion within these taxa and taxonomic decisions as to shorter proximally. Anterior metatarsus 0.5 as long their identity as true species problematical. However, as T1, distal segments successively shorter, preapical the range of variation recorded, is in my opinion, segment almost 2 as long as wide and about as consistent with that demonstrated in some other long as apical segment. Mid and posterior legs more Hemerodromiinae and to have omitted low sample slender; C3 with strong ‘brush’ of yellow anteroapi- size taxa from this study would have been to waste cal setae; F2 bearing series of four to five short black information on a very poorly understood genus. pd setae distally. T3 with short black seta dorsally Male genitalia are remarkably homogeneous (at least at base. Mid and hind legs with metatarsal segment on external morphology) amongst the yellow species very long, 0.7 as long as corresponding tibia, seg- and taxonomic decisions as to ‘what is a species’ are ments four and five very short, almost quadrate. informed more by characters of the antennae and Abdomen brownish dorsally, paler ventrally, tergites legs than by the genitalia. The modified, enlarged, bearing distinct but small bristles on posterior mar- black antennae and distinctive black pennate setae gin. Terminalia very similar to C. mapucheensis but on T1 of males contrast strongly with an otherwise cercus (Fig. 2) with distal process less strongly curved yellow body and may have behavioural significance inwards, not hidden in lateral view, one strong bris- in Chelipodozus. Pennate setae occur on the legs of tle on outer face of curve; apical ‘tooth’ larger, black. many female Empidinae where they have epigamic or Wing with veins dark, membrane faintly darkened. other display function. The presence of pennate setae Radial fork rather evenly expanded such that second as a secondary sexual character on the fore tibiae of submarginal cell (r4) not noticeably elongate or nar- males of the yellow species of Chelipodozus appears to rowed basally. Halteres yellowish white. be the first time such morphology has been reported Female. Length: body + head 3.5–4.0 mm, wing in the Hemerodromiinae and may indicate epigamic 4.0 mm. Very similar to male, differing as follows. behaviour at variance with other members of the Face less produced in profile below. Postpedicel elon- subfamily. Rapid and divergent evolution of male gate conical, sharply pointed apically; arista present, genitalia giving rise to species-specific genital mor- as long as postpedicel and with single basal article. phology is a widespread but not universal paradigm F1 with spines and denticles of ventral armature in animals with internal fertilization. Other sexually perhaps more numerous; femoral formula 5–6/15– dimorphic traits may often evolve at high rates due 19/12–15/5–7. T1 without dorsal pennate fringe. to their involvement in mate choice and selection Eighth abdominal segment elongate, about 3 as (Huber 2003). It seems likely that this is the case in long as wide, telescoping into previous segment; cer- yellow Chelipodozus species where reproductive isola- cus very long (Fig. 6). tion and speciation may have been driven primarily by mate recognition and sexual selection traits of the antenna and front leg rather than differentiation of General discussion genitalic characters. The yellow species (C. trauco, C. mapucheensis Chelipodozus is restricted to temperate rainforests C. araucariana, C. luteothorax, C. chodlipang, between 35° and 56° S in southern Chile and the C. australis and C. ochraceus) form an homogeneous genus thus has clear Andean biogeographic affinities grouping of predominantly yellow species in males of (sensu Morrone 2006). Chelipodozus araucariana, which the antennal postpedicel is usually lengthened C. luteothorax, C. chodlipang, C. chelipodiformis and the anterior tibia has a dorsal series of highly and possibly C. ochraceus are found in the main modified pennate setae. In contrast, C. cinereus and Cordillera. Chelipodozus trauco and C. mapucheensis C. chelipodiformis are brown species in which both appear to be a Valdivian element from an impor- sexes have similarly proportioned antennae and nor- tant locus of endemism centered on the coastal mal setae on the anterior tibia. While recognition of Valdivian range (Smith-Ramírez 2004). Chelipodo- the brown species and males of some of the yellow zus cinereus is probably more generally distributed species is straightforward, differential characters are while C. australis is likely a Magellanic element subtle in some males of the yellow group (e.g. in restricted to the far south. Little is known of the C. chodlipang and C. australis) and it is virtually biology of Chelipodozus. Larval habitats are poorly

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known in the Hemerodromiinae but, so far as is cur- MacDonald, J.F., 1988. New synonyms pertaining to rently understood, the aquatic habitat is confined to Chelifera and generic key for North American Hemero- the Hemerodromiini and Chelipodini are probably dromiinae (Diptera: Empididae). – Proceedings of the essentially terrestrial (Plant 2007). Adult Chelipodo- Entomological Society of Washington 90: 98–100. Morrone, J.J., 2006. Biogeographic areas and transition zus have been collected in a variety of moist tem- zones of Latin America and the Caribbean Islands perate forest types and there is no evidence of any based on panbiogeographic and cladistic analyses of association with water bodies. the entomofauna. – Annual Review of Entomology 51: 467–494. Plant, A.R., 1993. Sexual dimorphism in the genus Acknowledgements Monodromia Collin (Diptera: Empididae: Hemero- I would like to thank Elizabeth Arias & Kipling dromiinae). – New Zealand Journal of Zoology 20: Will for allowing me to join them in fieldwork in 207–210. Chile and also the many Chilean people who pro- Plant, A.R., 2005. The Hemerodromiinae (Diptera, Em- vided unrivalled hospitality and assistance in the pididae) of New Zealand I. Phyllodromia Zetterstedt. field. The Corporation Nacional Forestal (CONAF) – Studia dipterologica 12: 119–138. kindly granted permission to work in protected Plant, A.R., 2007. The Hemerodromiinae (Diptera: Em- areas of Chile. The support of collection curators, pididae) of New Zealand II. Chelipoda Macquart. Darren Mann (UMO), Nigel Wyatt (BMNH), Jeff – Zootaxa 1537: 1–88. Sinclair, B.J., 2000. 1.2. Morphology and terminology Cumming (CNC), Elizabeth Arias (EMEC) and of Diptera male terminalia. – In: Papp, L. & Darvas, Mario Elgueta (MNNC) is gratefully acknowledged. B. (Eds), Contributions to a Manual of Palaearctic Bradley Sinclair kindly commented on an earlier Diptera, Volume 1. General and Applied Dipterology. draft and usefully suggested the incorporation of Science herald, Budapest, pp. 53–74. additional material from CNC. Sinclair, B.J. & J.M. Cumming, 2006. The morphology, higher-level phylogeny and classification of the Empi- doidea (Diptera). – Zootaxa 1180: 1–172. References Smith, K.G.V., 1967. Family Empididae (Empididae, Collin, J.E., 1933. Diptera of Patagonia and South Hybotidae). – In: A Catalogue of the Diptera of the Chile. – British Museum (Natural History), London, Americas south of the United States: 39.1–39.67. De- 333 pp. partamento de Zoologia, São Paulo. Cumming, J.M., B.J Sinclair, & D.M. Wood, 1995. Stuckenberg, B.R., 1999. Antennal evolution in the Brach- Homology and phylogenetic implications of male ycera (Diptera), with a reassessment of terminology genitalia in Diptera-Eremoneura. – Entomologica relating to the flagellum. – Studia dipterologica 6: Scandinavica 26: 121–151. 33–48. Huber, B.A., 2003. Rapid evolution and species-specificity Smith-Ramírez, C., 2004. The Chilean coastal range: a of arthropod genitalia: fact or artifact? – Organisms, vanishing center of biodiversity and endemism in Diversity & Evolution 3: 63–71. South American temperate rainforests. – Biodiversity McAlpine, J.F., 1981. Morphology and terminology and Conservation 13: 373–393. – Adults. Chapter 2. – In: J.F. McAlpine, B.V. Pe- terson, G.E. Shewell, H.J. Teskey, J.R. Vockeroth, & D.M. Wood (Coords.). Manual of Nearctic Dip- tera, Volume 1. Agriculture Canada Monograph, 27: Received: 7 March 2007 9–63. Accepted: 10 July 2007

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