Neotrop Entomol (2015) 44:47–58 DOI 10.1007/s13744-014-0254-5

SYSTEMATICS, MORPHOLOGY AND PHYSIOLOGY

Review of the longipalpus-Group of Chrysotus Meigen (Diptera: Dolichopodidae), with Description of Four New Species RS CAPELLARI Depto de Biologia, Fac de Filosofia, Ciências e Letras de Ribeirão Preto, Univ de São Paulo, Ribeirão Preto, SP, Brasil

Keywords albipalpus-group, Diaphorinae, philtrum-group, taxonomy Correspondence RS Capellari, Depto de Biologia, Fac de Filosofia, Ciências e Letras de Ribeirão Preto, Univ de São Paulo, bloco 7, sala 5. Avenida Bandeirantes 3900, Ribeirão Preto, SP, 14040-901, Brasil; rscapellari@gmail. com Edited by Roberto A Zucchi – ESALQ/USP Received 9 May 2014 and accepted 6 November 2014 Published online: 15 January 2015 * Sociedade Entomológica do Brasil 2014

Abstract The longipalpus-group of Chrysotus Meigen is reviewed and comprises eight species: Chrysotus coquitos n. sp. (Mexico), Chrysotus crosbyi Van Duzee (Eastern US, Bermuda, Cuba, Puerto Rico and Brazil; introduced in Australasian Region), Chrysotus longipalpus Aldrich (=Chrysotus sagittarius Van Duzee, n. syn.; Eastern US, Saint Vincent, Grenadas and Brazil; introduced in the Afrotropical, Australasian, Oriental and Palaearctic regions), Chrysotus miripalpus Parent (Costa Rica and Brazil), Chrysotus neopedionomus n. sp. (Brazil), Chrysotus pachystoma n. sp. (Belize), Chrysotus xiphostoma Robinson (Dominica and Saint Lucia), and Chrysotus zumbadoi n. sp. (Costa Rica). Lectotype and paralectotypes are designated for Chrysotus pallidipalpus Van Duzee, and a neotype for C. miripalpus. Illustrations of the hypopygium and ovipositor, photos of the male palpus and a key to species of the group are provided.

Introduction Chrysotus Meigen is the largest genus of Diaphorinae (Diptera: Dolichopodidae), comprising over 320 described species worldwide (Yang et al 2006). However, it is also the most poorly defined genus in that subfamily. This is historically indicated by the number of species described under Chrysotus but later transferred to many different genera: Achradocera Becker (Diaphorinae), Acropsilus Mik (Peloropeodinae), Asyndetus Loew (Diaphorinae), Chrysotimus Loew (Peloropeodinae), Diaphorus Meigen (Diaphorinae), Elmoia Evenhuis (Sympycninae), Eurynogaster Van Duzee (Sym pycni nae), Lamprochromus Mik (Rhaphiinae), Lyroneurus Loew (Diaphorinae), Melanostolus Kowarz (Diaphorinae), Nematoproctus Loew (Rhaphiinae), Telmaturgus Mik (Sympycninae), Teuchophorus Loew (Sympycninae), Thinophilus Wahlberg (Hydrophorinae), Thrypticus Gerstaecker (Medeterinae), Sympycnus Loew (Sympycninae), and Xanthina Aldrich (Achalcinae). The New World fauna was particularly challenging for taxonomists during the past century, since recognition of Chrysotus as usually defined in Europe hardly fits many Nearctic and Neotropical species. Early attempts to restrict the genus to a meaningful concept addressed diagnostic characters

between Chrysotus and Diaphorus, traditionally seen as closely related (Loew 1864, Aldrich 1896, 1902, Becker 1922, Van Duzee 1924). The Robinson & Vockeroth (1981) key offered the most useful distinction between the two genera, providing clearer limits to Diaphorus and allowing recombination of several species in Chrysotus from Diaphorus or viceversa (e.g., Bickel 1997, Pollet et al 2004, Capellari & Amorim 2010). Nevertheless, couplet 16 in Robinson & Vockeroth’s (1981) key leads to a dichotomy between Diaphorus and Achradocera plus Chrysotus, in such a manner that characters used to diagnose Diaphorus are not exclusive of Chrysotus. Pollet et al (2004) considered Lyroneurus a junior-synonym of Chrysotus, since the genus also runs to that same couplet as Achradocera and Chrysotus in Robinson & Vockeroth’s (1981) key. More recently, Capellari & Amorim (2012) found that the monotypic Azorean genus Falbouria Dyte shares that same set of characters with Achradocera, Chrysotus, and Lyroneurus, and suggested that they compose a clade within Diaphorinae. As such, although distinction between Chrysotus and Diaphorus is currently well established, this is actually achieved after an improved definition of Diaphorus. Characters in the second half of couplet 16 in Robinson & Vockeroth’s (1981) key fit a group of

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genera—Achradocera, Chrysotus, Falbouria, and Lyroneurus— of which Chrysotus is probably a paraphyletic residue and still persists as a taxonomic imbroglio (Capellari & Amorim 2012). Relatively few authors established species-groups within Chrysotus (e.g., Van Duzee 1924, Negrobov 1980, Bickel & Sinclair 1997, Wei & Zhang 2010), and a number of poorly known species still remain unassigned to any group in the vaguely defined genus. Van Duzee’s (1924) revision of the North American Chrysotus recognized 20 species-groups (Achradocera included). At that time, four species were ascribed to his longipalpus-group: Chrysotus crosbyi Van Duzee, Chrysotus flavus Aldrich (later transferred to Xanthina), Chrysotus longipalpus Aldrich, and Chrysotus sagittarius Van Duzee. In this paper, the longipalpus-group of Chrysotus is reviewed and argued to compose a clade within that genus. Two known species are formally referred to the group, and four are described as new from Mexico, Belize, Costa Rica, and Brazil.

Material and Methods The morphological nomenclature follows Cumming & Wood (2009). Measurements of the leg segments are representative ratios given according the formula: trochanter+femur, tibia, tarsomeres 1, 2, 3, 4, and 5. The CuAx ratio represents the length of the dm-cu crossvein/distal section of the anal vein. The abbreviations used in the text include: I, II, and III: pro–, meso–, and metathoracic legs; MSSC, male secondary sexual character(s). While describing the hypopygium, ‘dorsal’ and ‘ventral’ refer to the morphological position prior to genitalic rotation and flexion; as such, the top of the drawings in lateral view is actually ventral on the specimens. Features found in all of the species are listed under the species-group description and not repeated in each species treatment. Exclamation marks “[!]” in “Distribution” sections refer to new records. Photographs of palpi were produced using a Leica DFC camera attached to a Leica M205 C stereomicroscope, and Photoshop CS4. Specimens examined or only cited in this study belong to the following institutions: American Museum of Natural History (AMNH, New York, US), Bernice Pauahi Bishop Museum (BPBM, Honolulu, US), California Academy of Sciences (CAS, San Francisco, US), Cornell University Insect Collection (CUIC, Ithaca, US), Canadian National Collection (CNC, Ottawa, Canada), Hawaii Department of Agriculture (HDOA, Honolulu, US), Instituto Nacional de Biodiversidad (INBio, San José, Costa Rica), Museu de Zoologia da Universidade de São Paulo (MZSP, São Paulo, Brazil), National Museum of Natural History (USNM, Washington, US), Natural History Museum (BMNH, London, UK), Universidad Nacional Autónoma de México (UNAM, Mexico D.F.), Washington State University

(WSU, Pullman, US), and Zoologisches Museum, Universität von Hamburg (ZMUH, Germany, Hamburg). Label data for primary types are cited verbatim in quotation marks (lines separated by “|”), and annotations in square brackets. Genus Chrysotus Meigen Chrysotus Meigen (1824): 40. Type species: Musca nigripes Fabricius, designation by Westwood (1840: 134). See Capellari & Amorim (2012) for further discussion on the type-species of the genus. The longipalpus species-group Diagnosis (based on males) Head. Eyes nearly contiguous below antennae, face as broad as middle ocellus. Antennal post-pedicel sub-triangular. Palpus variously enlarged (Fig 1a–h), usually longer than face (MSSC). Legs. First tarsomere I with ventral swelling (Fig 2b) (MSSC). Tibia III with row of anterior to anterodorsal setae. Hypopygium. Anteroventral margin of epandrium pointed; surstylus with single spine; phallus with small projection at apex; postgonites and hypandrial apodemes short. Description Male. Small-sized diaphorines (less than 2 mm long). Head. Frons about half of head width at the dorsal-most point, converging towards antennae; face as broad as middle ocellus, widening above and below; eyes nearly contiguous below antennae, ventral facets enlarged (MSSC); palpus variously enlarged and ornamented, usually longer than face (MSSC, Fig 1a–h); pair of strong divergent, proclinate ocellar setae; pair of minute post-ocellar setae; pair of strong convergent, proclinate vertical setae; pair of paravertical setae, around a third of the length verticals; postoculars in one row, those on ventral half whitish, the ventral-most longer. Antenna: scape and pedicel short, pedicel with crow of apical setae; postpedicel sub-triangular, pubescent, arista-like stylus inserted near apex, slightly displaced laterad, bi-articulate at base. Thorax. Acrostichals in two irregular rows; six pairs of dorsosentrals, first pair very small; one strong and two smaller post-pronotals; upper part of proepisternum, in front of anterior spiracle, bare, lower part usually with two setae; one pre- and one sutural intra-alar setae; one pre- and two postsutural supra-alar setae; one post-alar; two notopleurals; pair of strong medial scutellars and one smaller pair laterad. Wing (Fig 1i). Membrane hyaline, veins brown. R2+3 reaching C at 3/4 of wing length. R4+5 and M subparallel. CuAx ratio 0.3– 0.5. Legs. Usually mostly yellow, except by the brown coxae II and III (sometimes also I), and tips of femur III and tarsi. Claws and pulvilli small. Setae black, except when noted. I.

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Fig 2 Anterior male tarsus of Chrysotus. a Chrysotus gramineus (Fallén); b Chrysotus zumbadoi n. sp. Arrow indicates the ventral swelling of first tarsomere. Scale bars: 0.1 mm.

n. sp.). Femur I with posteroventral row of brownish setae and row of two to four posterior setae at apex. Tibia II with one strong anterodorsal seta at basal third and three to four apicals. III. Coxa III with 1 long basal seta. Femur III with row of three to five anteroventral setae. Tibia III with row of anterior to antero-dorsal setae (MSSC) and three to four apicals. Abdomen. Cylindrical, tergites brown, sternites lighter; tergites covered by short setation, tergite 6 setose (setae as long as setae on preceding tergites). Sternite 8 only with small setae (without bristles). Hypopygium (Figs 3a–b and 4a, d, g, j–m). Globular, partially concealed by tergite 6; genital foramen left lateral, central. Anteroventral margin of epandrium pointed; epandrial lobe with three to four setae arising from short projections. Hypandrial apodeme short. Surstylus as a single lobe (ventral lobe when compared with other diaphorines), with spine at apex and small seta posteriad. Apex of phallus with ventral projection at apex (cf. Fig 3a–b: arrow). Postgonite short, covered by microtrichia. Cercus short, pale. Female. Similar to males, except by MSSC and as noted. Head. Eyes approximate below antennae, but face broader, 1/4–1/3 of frons width; clypeal suture evident; palpus shorter, usually unmodified. Antennal postpedicel trapezoid. Legs. Tibia III without row of antero-dorsal setae. Abdomen. Segments 6–8 and acanthophorites telescoped into preceding segments. Ovipositor (Figs 3c–d and 4b–c, e–f, h–i). Tergite and sternite 8 divided into two rod-like sclerites, tergites with a delicate dorsal plate each with minute seta; acanthophorite (=tergite 10) with three to five pointed spines.

Comments

Fig 1 Male palpus and wing of Chrysotus. a Chrysotus coquitos n. sp.; b Chrysotus crosbyi; c Chrysotus longipalpus; d Chrysotus miripalpus; e Chrysotus neopedionomus n. sp.; f Chrysotus pachystoma n. sp.; g Chrysotus xiphostoma; h Chrysotus zumbadoi n. sp.; i Chrysotus crosbyi. Scale bars: 0.5 mm.

Anterior surface of coxa I covered by pale to whitish setae (brownish in Chrysotus pachystoma n. sp.). First tarsomere I with ventral swelling (Fig 2b) (MSSC). II. Setae on anterior surface of coxa II pale to whitish (brownish in C. pachystoma

The hypothesis of monophyly for the longipalpus-group can be based on apomorphic characters of males, such as the ventral swelling on first tarsomere I (Fig 2b; compare with the unmodified condition in Fig 2a), anterodorsal setation on tibia III, and phallic ventral projection (cf. Fig 3a–b: arrow). Although modifications of the male palpus are also apomorphic (Fig 1a–h), it is difficult to characterize common shared features, given the uniqueness of shape and color for each species. Indeed, species of this group differ mostly by characters of the male palpus and, to a lesser, degree by the

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Key to males of the longipalpus-group of Chrysotus

Fig 3 Chrysotus crosbyi. a–b hypopygium: left lateral (a) and ventral (b); c–d ovipositor: dorsal (c) and right lateral (d). Arrows indicate the ventral phallic projection. Abbreviations: ac acanthophorite, ce cercus, el epandrial lobe, pg postgonite, sur surstylus, st sternite, tg tergite. Scale bar: 0.1 mm.

color of the antenna and coxae. Even hypopygial characters show very few differences among the species (see Figs 3a–b and 4a, d, g, j–m), and none of them are regarded as diagnostic, as is commonly found for other Insect groups (a more detailed illustration is presented only for C. crosbyi). MSSC of the palpus in the group certainly play a role during courtship recognition, in such a way isolation of the species is probably due to pre-copulatory mechanisms related to the palpus, rather than copulatory ones related to hypopygial morphology. Most of the species of the longipalpus-group are known from Central and northern South America, and the group can be considered as primarily Neotropical in distribution. Nevertheless, both C. crosbyi and C. longipalpus are widespread and found in other zoogeographical regions due to secondary introductions (see further details in species sections).

1. Palpus clearly divided into a basal enlarged part and an apical strap-like projection after abrupt tapering .......…2 1’. Palpus not clearly divided into two parts................…4 2. Palpus brownish, thick (Fig 1f); coxa I light brown, yellow at apex, with brownish setae on anterior surface ….......................................................C. pachystoma n. sp. 2’. Palpus mostly yellowish, thin; coxa I entirely yellow, with white setae on anterior surface ..........................…3 3. Basal part of palpus densely covered with brownish setae; distal part slightly tapering towards apex, without apical lamella (Fig 1a) …...........................C. coquitos n. sp. 3’. Basal part of palpus covered with few brownish setae; distal part strip-like, with apical lamella (Fig 1d) …....................................................... C. miripalpus Parent 4. Palpus oblong, and rounded distally, entirely bright silvery (Fig 1b) …................................C. crosbyi Van Duzee 4’. Palpus either uniformly narrow or only gradually tapering toward apex, never entirely bright silvery, but eventually bright yellowish ..........................................…5 5. Palpus uniformly narrow and straight …......................6 5’. Palpus tapering towards apex ….................................7 6. Antenna brown; palpus brown with silvery tip (Fig 1h); coxa I brown, femur I yellow ….............. C. zumbadoi n. sp. 6’. Antenna yellow; palpus yellow, bright silvery (Fig 1g); coxa and femur I whitish yellow.......................................... ...................................................... C. xiphostoma Robinson 7. Palpus sub-triangular with rounded apex, entirely yellow, bright silvery (Fig 1c) …........C. longipalpus Aldrich 7’. Palpus pointed, basal half yellow, apical half brownish, silvery pruinose (Fig 1e) …..................................................... ......................................... Chrysotus neopedionomus n. sp.

Chrysotus coquitos n. sp. (Figs 1a and 4j) Type material Holotype male, “Mexico: Chiapas: Biosphere Reserve | La Encrucijada; 15°10′N 93°50′W; | 20–29.v.2012; sticky trap; mangrove. | M. Solórzano-Kraemer leg.”, “Chrysotus | coquitos | Capellari HOLOTYPE [red label]” (UNAM). Paratypes: six males, one female, same data as the holotype (two males in UNAM, two males in CNC, two males, and one female in MZSP). Re-description Male. Body length, 1.1–1.4 mm. Wing length, 1.0–1.1 mm, width, 0.4–0.5 mm. Head. Frons greenish, with coppery reflections; antenna yellow, postpedicel lighter; face light

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Fig 4 Hypopygia and ovipositors of Chrysotus. a–c Chrysotus longipalpus; d–f Chrysotus miripalpus; g–i Chrysotus pachystoma n. sp.; j Chrysotus coquitos n. sp.; k Chrysotus neopedionomus n. sp.; l Chrysotus xiphostoma; m Chrysotus zumbadoi n. sp. Hypopygium, left lateral (a, d, g, j–m); ovipositor, dorsal (b, e, h); ovipositor, right lateral (c, f, i). Scale bar: 0.1 mm.

brown; proboscis yellow with whitish setae; palpus (Fig 1a) with basal part enlarged, oblong, mostly yellow and covered by brownish setae; distal part tapering, around 2/3 as long as basal part, mostly brownish, but silvery tip, without conspicuous setae, except one to two near base (ratio tibia I: palpus,

22:21) (MSSC). Thorax. Mesonotum and scutellum metallic green with coppery reflections; mesopleuron brown with some bluish reflections and little pruinosity. Wing. Halter light yellow, calypter brownish. Legs. I: 24, 22, 11, 4, 3, 2, 2. II: 32, 30, 14, 6, 4, 3, 2. III: 33, 31, 8, 7, 4, 3, 2. Coxa II and basal

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half of III light brown; apical 1/3–1/4 of femur III and apex of tarsi brown; legs otherwise yellow. Setae on coxae I and II white. Hypopygium. As in Fig 4j. Female. Body length, 1.4 mm. Wing length, 1.2 mm, width, 0.4 mm. Similar to males, except by MSSC and as noted. Head. Palpus a little enlarged, as long and as broad as face, light brown, covered by black setae, 2 longer at apex. Scape and pedicel dark yellow, inner surface of postpedicel brownish. Legs. I: 23, 22, 11, 4, 3, 2, 2. II: 30, 30, 13, 6, 4, 3, 2. III: 33, 31, 5, 5, 4, 3, 3. Ovipositor (not dissected). Similar to those illustrated for other species in the group. Etymology Alluding to the type-locality (Biosphere Reserve La Encrucijada), referred to by the residents as “Coquitos.” The name is treated as a noun in apposition. Comments Both C. coquitos n. sp. and C. miripalpus have similar shape and color of male palpus, but they can be distinguished by the apical lamella in C. miripalpus. Distribution Mexico. Chrysotus crosbyi Van Duzee (Figs 1b, i and 3) Chrysotus crosbyi Van Duzee (1924): 43. Holotype in the CUIC. Chrysotus magnipalpus Van Duzee (1927): 2 (synonymized by Woodley 1996: 202). Holotype in the AMNH. Material examined United States, Florida, Highlands Hammock State Park, three males, 17.iv.1989, J.R. Vockeroth leg. (CNC); Bermuda, Paget, Paget Marsh, one male, one female, 3.vi.1991, N. Woodley leg. (CNC); French Guiana, Patawa, malaise, 180 m, 13 males, two females, viii–x.2008, J. Cerda, O. Morvan leg. (CNC). Brazil, Rondônia, Campo Novo de Rondônia (10°40′06″S, 63°29′00″), malaise, 248 m, 44 males, 14 females, 3– 15.xii.2011, D.S. Amorim, D.C. Ament & P.R. Riccardi leg. (MZSP); three males, one female, same data, but 5.2 km from Porto Velho (08°43′40″S 63°57′44″W) (MZSP); 12 males, one female, same data, but Monte Negro (10°16′35″S 63°20′ 40″W), sweeping (MZSP); three males, Pernambuco, Recife, Parque dos Dois Irmãos, malaise, trilha ponto 5, 17– 20.vii.2002, S.T.P. Amarante et al leg. (MZSP); one male, same data, but trilha ponto 1 (MZSP); two males, same data,

but 20–23.vii.2002 (MZSP); eight males, same data, but bosque, ponto 3 (MZSP); two males, Bahia, Porto Seguro, Estação Ecológica Pau Brasil (16°23′17″S 39°10′55″W), malaise, 107 m, 17.v.2002, C.O. Azevedo et al leg. (MZSP); two males, one female, same data, but Mata de São João, Reserva de Sapiranga (11°22′39″S 37°25′04″W), malaise, ponto T1, 22– 25.vii.2001, M.T. Tavares et al leg. (MZSP); two males, same data, but ponto T6 (12°33′38″S 38°02′57″W) (MZSP); four males, Espírito Santo, Linhares, Reserva Biológica de Sooretama (18°58′03″S 40°07′56″W), 21–27.iii.2002, malaise, ponto 2, C.O. Azevedo et al leg. (MZSP); one male, same data, but ponto 5 (18°58′02″S 40°08′06″W) (MZSP); one male, one female, same data, but ponto 1 (18°58′02″S 40°07′53″W) (MZSP); one male, same data, but ponto 4 (18°58′03″S 40°08′03″W) (MZSP). Reported material (Dan Bickel, pers. comm.) Commonwealth of the Northern Mariana Islands, Rota, one male, four females, farm plots, stream area, 14.v.2004, 14°07.111′ N 145°10.708′ E. Guam, Yigo Village, six males, University of Guam, Yigo Expertiment Station, 13°31.924′ N 144°52.2984′ E, 6.v.2004; two males, Inarajan Village, Pauliluc Farm & Pauliluc River, 5.v.2004, 13°15.948′ N 144°45.166′ N; one male, one female, Mangilao Village, University of Guam campus, 6.v.2004, 13°25.714′ N 144°47.913′ N; one male, one female, Merizo Village, stream at Santa Maria Kamatlin Park, 8.v.2004, 13°15.948′ N 144°40.125′ E (all WSU, collected by R.S. Zack). French Polynesia, Austral Islands, Rimatara I., Oromana Plateau, one male, 20°39.05′ S 152°48.29′W, 30.x.2004, E. Claridge leg. (BPBM). Re-description Male. Body length, 1.2–1.5 mm. Wing length, 1.0–1.1 mm, width, 0.4–0.5 mm. Head. Frons brownish, with bluish to violet reflections; antenna brown; face brown; proboscis brownish with pale setae; palpus (Fig 1b) thin, oblong, entirely bright silvery, bare or only with an inconspicuous whitish seta near apex (ratio tibia I: palpus, 11:11) (MSSC). Thorax. Mesonotum metallic green with coppery reflections, scutellum lighter; mesopleuron brown with some bluish reflections and little pruinosity. Wing. (Fig 1i) Halter dark yellow, calypter brownish. Legs. I: 26, 25, 11, 6, 4, 3, 2. II: 32, 32, 16, 9, 5, 4, 3. III: 35, 36, 9, 10, 5, 4, 3. All coxae light brown; femur I dorsally to entirely brownish; apical 1/3–1/4 of femur III infuscated; apex of tarsi brown; legs otherwise yellow. Setae on coxae I and II pale. Hypopygium. As in Fig 3a–b. Female. Body length, 1.2–1.4 mm. Wing length, 1.1–1.2 mm, width, 0.5–0.6 mm. Similar to males, except by MSSC and as noted. Head. Palpus a little enlarged, as long and as broad as

Review of the longipalpus-Group of Chrysotus

face, dark brown, covered by black setae. Legs. I: 27, 25, 15, 5, 4, 3, 2. II: 35, 33, 15, 7, 5, 4, 3. III: 37, 35, 10, 10, 4, 3, 2. Ovipositor. As in Fig 3c–d.

Comments Robinson (1975) suggested that this species (as C. magnipalpus) is a possible relative of Chrysotus mediocaudatus Robinson. Nevertheless, Robinson’s (1975) description of C. mediocaudatus does not fit diagnostic characters for the longipalpus-group, and the species should be placed elsewhere. Woodley (1996) offered a detailed description of C. crosbyi and stressed that color features can be unreliable to identify widespread species, such as this one. In fact, the distribution of C. crosbyi proved to be much wider than previously known, and the species is also present in Pacific islands (Dan Bickel, pers. comm., see below). A species of Chrysotus “similar to C. crosbyi” reported by Robinson (1977: 310) from Bahia (Brazil) is probably that species.

Distribution Eastern US (Nearctic), Bermuda, Cuba, French Guyana [!], Puerto Rico, Brazil [!] (Neotropical), Guam [!], French Polynesia [!] and Commonwealth of the Northern Mariana Islands [!] (Australasian) (Pollet et al 2004, Bickel, pers. comm.).

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Additional material United States, Hawaii, Honolulu, Oahu, 26 males, 11 females, 8–24.ix.1966, J.R. Vockeroth leg. (CNC); BRAZIL, Amazonas, Biological Reserve ZF2, near Base Camp (02°38′16″S 60°09′ 22″W), four males, one female, 02–06.xii.2013, malaise, 109 m, J.A. Rafael & D.S. Amorim leg. (MZSP). Reported type material (Neal Evenhuis, pers. comm.) Chrysotus pallidipalpus Van Duzee, male lectotype, here designated: “Honolulu, T.H. [black printing] | 1-21-30 [black handwritten]”, “O.H. Swezey | Collector [black printing]”, “Chrysotus | pallidipalpus [handwritten] | Holotype. Van Duzee [red printing on a red-ringed square label]”; one female paralectotype, same data as the lectotype [both glued on their left side to the same point, the lectotype closer to the pin] (BPBM Type No. 4069). Remaining paralectotypes: one female, same data as the lectotype; one female: Waialua, Oahu, 16 Jan 1930, F.X. Williams (both in HDOA). Additional reported material (Dan Bickel, pers. comm.) Kirabati Line Islands: Palmyra Atoll, Eastern Islands, two males, four females, 11–16.vi.2003, Malaise trap, Haines et al leg. (BPBM). Re-description

Chrysotus longipalpus Aldrich (Figs 1c and 4a-c) Chrysotus longipalpus Aldrich (1896): 329. Syntypes in the BMNH and USNM. Chrysotus sagittarius Van Duzee (1924): 42, new synonym. Holotype in the CAS. Chrysotus pallidipalpus Van Duzee (1933): 313 (synonymized by Evenhuis 1996: 28). Lectotype in the BPBM. Chrysotus vulgaris Van Duzee (1933): 313, preoccupied by Van Duzee (1924: 15) (synonymized with C. pallidipalpus by Hardy & Kohn (1964): 239). Holotype in the BPBM. Chrysotus elegans Parent 1937: 68 (synonymized with C. pallidipalpus by Parent 1939: 248). Syntype in the BMNH.

Male. Body length, 1.3–1.6 mm. Wing length, 1.1–1.3 mm, width, 0.4–0.5 mm. Head. Frons dark green, with coppery and violet reflections; antenna yellow, postpedicel lighter; face light brown; proboscis brownish with pale setae; palpus (Fig 1c) sub-triangular with rounded apex, yellow, bright silvery, lateral margin darker, basal 2/3 covered by sparse brownish setae (ratio tibia I: palpus, 20:15) (MSSC). Thorax. Mesonotum metallic green with coppery reflections, scutellum lighter with some bluish reflections; mesopleuron brown with some bluish reflections and little pruinosity. Wing. Halter dark yellow, calypter brownish. Legs. I: 22, 20, 11, 5, 4, 3, 2. II: 30, 30, 15, 6, 4, 2, 2. III: 32, 33, 9, 9, 5, 4, 3. Coxa II and extreme base of coxa III light brown; apical 1/3–1/4 of femur III and apex of tarsi brown; legs otherwise yellow. Setae on coxae I and II white. Hypopygium. As in Fig 4a.

Type material examined Chrysotus sagittarius Van Duzee, male holotype, “Buffalo, N.Y. | 4-6-09 | M.C.V. Coll.”, “Chrysotus | sagittarius [sic, handwritten] | Holotype Van Duzee [red printing on a redringed square label]”, “California Academy | of Sciences | Type n°. 3555” (CAS).

Female. Body length, 1.2–1.5 mm. Wing length, 1.2–1.4 mm, width, 0.4–0.5 mm. Similar to males, except by MSSC and as noted. Head. Palpus a little enlarged, as long and as broad as face, dark brown, covered by black setae. Antenna brown, scape lighter. Legs. I: 25, 22, 11, 5, 4, 3, 2. II: 30, 29, 14, 5, 4, 3, 2. III: 30, 32, 8, 8, 5, 4, 2. Ovipositor. As in Fig 4b–c.

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Comments

Additional material

Van Duzee (1924) described C. sagittarius and redescribed C. longipalpus based on the type material in the same work. I compared the male holotype of C. sagittarius with Hawaiian and Brazilian specimens of C. longipalpus, and differences in form of the palpus and color of antennae and coxae listed by Van Duzee (1924) were regarded as minor variations. As a consequence, I treated both species as synonyms. Chrysotus longipalpus is a tramp species, known from all zoogeographical regions and found overseas in sympatry with C. crosbyi in Guam (see below). The observed association of C. longipalpus with greenhouses (Parmenter 1942) can be related to the worldwide distribution of this species, and the same could be conjectured for C. crosbyi. Neal Evenhuis (pers. comm.) noted that Van Duzee’s (1933) description of C. pallidipalpus failed to designate a holotype from the type series, which originally consisted of nine specimens (three males, six females). A specimen labeled as holotype is here designated as the male lectotype to fix the proper interpretation of the species name. Additionally, one female glued in the same pin as the lectotype and two females in the HDOA are regarded as paralectotypes (...), as well as a couple (one male and one female) in CAS. The remaining specimens of the type series could not be found and are treated here as lost.

One male, same data as the neotype; one male, same but Puntarenas, PN Corcovado, one male, 16.iii–6.iv.2003, malaise trap, 130 m, K. Caballero, M. Moraga, A. Azofeita leg; one male, two females, same data, but 80 m, 22.iii–6.iv.2003; one male, same data, but 100–300 m, 19.vi–8.vii.2003; one male, same data, but Golfito, 50–100 m, 22–29.iv.2004, W. Porras, B. Gamboa, D. Briceño, M. Moraga leg; one male, same data, but Guanacaste, Bagaces, PN Palo Verde, 5.i– 7.ii.2000, 50 m, I. Jiménez leg; one male, same data, but Liberia, PN Santa Rosa, 6–7.ix.2002, sweeping, 300 m, D. Briceño leg.; one male, same data, Alajuela, San Carlos, PN Arenal, 1–26.x.1999, malaise trap, 600 m, G. Garballo leg. (all INBio).

Distribution Eastern US [!] (Nearctic), Saint Vincent, Grenadas, Brazil [!] (Neotropical), Hawaii, Guam, French Polynesia, Kirabati [!] (Australasian), Mauritius (Afrotropical), Diego Garcia (Oriental), England, France, The Netherlands, and Finland (Palaearctic) (Parmenter 1942, Bickel 2005, and pers. comm., Grichanov 2011). Chrysotus miripalpus Parent (Figs 1d and 4d–f) Chrysotus miripalpus Parent 1928: 165. Holotype in the ZMUH (destroyed; neotype in the INBio).

Re-description Male. Body length, 1.2–1.4 mm. Wing length, 1.2–1.3 mm, width, 0.5–0.6 mm. Head. Frons brownish, with coppery and violet reflections; antenna yellow, postpedicel lighter; face light brown; proboscis brownish with pale setae; palpus (Fig 1d) with basal part enlarged, oblong, distal part strip-like, as broad as face and as long as basal part; basal part mostly yellow (inner margin darker) covered by sparse long, brownish setae; distal part brown, short pubescent, ending in an apical lamella with silvery tip (ratio tibia I: palpus, 27:31) (MSSC). Thorax. Mesonotum metallic green with coppery reflections, scutellum lighter with some bluish reflections; mesopleuron brown with some bluish reflections and little pruinosity. Wing. Halter light yellow, calypter brownish. Legs. I: 29, 27, 13, 5, 4, 3, 2. II: 32, 34, 15, 10, 6, 3, 2. III: 37, 36, 10, 10, 5, 3, 2. Coxa II and basal half of III light brown; apical 1/3–1/4 of femur III and apex of tarsi brown; legs otherwise yellow. Setae on coxae I and II white. Hypopygium. As in Fig 4d. Female. Body length, 1.4 mm. Wing length, 1.4 mm, width, 0.6 mm. Similar to males, except by MSSC and as noted. Head. Palpus a little enlarged, as long and as broad as face, dark brown, covered by black setae, at least two to three setae at apex longer. Scape and pedicel dark yellow, postpedicel brown. Legs. I: 27, 25, 12, 5, 3, 2, 2. II: 31, 30, 15, 7, 4, 3, 2. III: 35, 36, 9, 10, 5, 3, 2. Ovipositor. As in Fig 4e–f.

Type material examined Comments Male neotype, here designated, “Costa Rica. Prov. Limón, P.I. L.A., Valle del | Silencio, Send. Circular, 2480 m, 20–26 SEP | 2003, D. Rubi, R. González, R. Delgado, M. Alfaro, Amarilla, L_S_340258_577465 | #75068”, “INB0003995283 | INBIOCRI COSTA RICA” [barcode label], “Chrysotus | miripalpus | Parent NEOTYPE [red label]”.

The type of C. miripalpus (ZMUH) was destroyed during the Second World War, but Parent’s (1928) description allows identification of the species, and thus, a specimen from Costa Rica is here designated as the male neotype to ensure proper interpretation of the species name.

Review of the longipalpus-Group of Chrysotus

55

Distribution

Chrysotus pachystoma n. sp. (Figs 1f and 4g–i)

Costa Rica, Panama, and Brazil (Parent 1930, Robinson 1970).

Type material

Chrysotus neopedionomus n. sp. (Figs 1e and 4k) Type material Holotype male, “SISBIOTA- | CNPQ/FAPESP [vertical line] | BRASIL, RO [Rondônia State], Campo Novo de Rondônia | Amorim Farm 10°40′06″ S 63°29′00″W | 03–15.xii.2011, malaise trap, 248 m | Amorim, Ament & Riccardi leg.”, “Chrysotus | neopedionomus | Capellari HOLOTYPE [red label]” (MZSP). Paratypes: two males, same data as holotype (MZSP). Additional material examined Brazil, Tocantins, Brejinho do Nazaré, Crixás River, one male, 11.iv.1998, malaise, S.H. Tozoni leg. (MZSP). Description Male. Body length, 1.2–1.3 mm. Wing length, 1.0–1.1 mm, width, 0.4–0.5 mm. Head. Frons metallic green, more bluish above, with little pruinosity below; face dark brown; antenna yellow, postpedicel lighter; proboscis yellow with whitish setae; palpus (Fig 1e) pointed, basal half yellow, covered with few brownish setae (smaller and pale on inner margin), distal half brown, silvery pruinose, without setae (ratio tibia I: palpus, 19:21). Thorax. Mesonotum metallic green with coppery reflections, scutellum concolourous with mesonotum; mesopleuron brown with some bluish reflections and little pruinosity. Wing. Halter dark yellow, calypter brownish. Legs. I: 21, 19, 10, 4, 3, 2, 2; II: 23, 24, 11, 5, 3, 2, 2; III: 26, 27, 6, 7, 4, 3, 2. Coxa II, basal 1/2–2/3 of coxa III, apical 1/2–1/3 of femur III, basal 1/2–2/3 of tibia I, and apex of tarsi brown; legs otherwise yellow. Setae on coxae I white, on coxae II pale to brownish. Hypopygium. As in Fig 4k.

Holotype male, “Belize: Belize District. | Caye Caulker Is. | 6.xii.2001, L. Masner | Yellow Pan Trap | mangrove”, “Chrysotus | pachystoma | Capellari HOLOTYPE [red label]” (CNC). Paratypes: 18 males, six females, same data as the holotype (all CNC, but five males and five females in MZSP). Description Male. Body length, 1.5–1.7 mm. Wing length, 1.2–1.3 mm, width, 0.4–0.5 mm. Head. Frons brownish, with coppery reflections; antenna orange to dark yellow, postpedicel lighter; face dark yellow; proboscis brownish with pale setae; palpus (Fig 1f) large and thick, basal part grayish and covered by black setae, distal part tapering, shorter than basal part, dark brown, without setae and with silvery tip (ratio tibia I: palpus, 27:27) (MSSC). Thorax. Mesonotum metallic green with coppery reflections, scutellum lighter; mesopleuron brown with some bluish reflections and little pruinosity. Wing. Halter yellow, calypter brownish. Legs. I: 29, 27, 12, 5, 4, 3, 3. II: 32, 30, 15, 7, 4, 3, 2. III: 35, 35, 9, 9, 4, 3, 3. All coxae light brown, coxae I and III yellowish at apex; apex of tarsi brown; apical 2/4 of femur III infuscated, darkening towards apex; legs otherwise yellow. Setae on coxae I and II brownish. Hypopygium. As in Fig 4g. Female. Body length, 1.5–1.6 mm. Wing length, 1.3–1.4 mm, width, 0.5–0.6 mm. Similar to males, except by MSSC and as noted. Head. Palpus a little enlarged, as long and as broad as face, dark brown, covered by black setae, at least two to three setae at apex longer. Antennal postpedicel brown. Legs. I: 27, 25, 12, 5, 4, 3, 3. II: 32, 30, 15, 7, 4, 3, 2. III: 36, 37, 7, 7, 4, 2, 2. Ovipositor. As in Fig 4h–i. Etymology From Greek pachys (thick) and stoma (mouth), referring to the thick palpus of males.

Female. Unknown. Comments Etymology From Greek, neos (new) and pedionomos (inhabitant of the plain), referring to the type-locality, Campo Novo (“new plain”) de Rondônia.

C. pachystoma n. sp., C. miripalpus, and C. coquitos n. sp. have male palpus basally enlarged and followed by a distal narrow projection (couplet 2 in the key above). Moreover, C. pachystoma and C. coquitos are the only members of the longipalpus-group so far reported from mangroves.

Distribution

Distribution

Brazil.

Belize.

Capellari

56

Chrysotus xiphostoma Robinson (Figs 1g and 4l) Chrysotus xiphostoma Robinson 1975: 84. Holotype in the USNM. Material examined Saint Lucia, Praslin, seven males, iv.2000, S. Lesmond leg. (CNC). Re-description Male. Body length, 1.3–1.4 mm. Wing length, 1.1–1.2 mm, width, 0.4–0.5 mm. Head. Frons metallic green, with bluish reflections and little pruinosity below; antenna yellow, postpedicel lighter; face light brown; proboscis yellow with whitish setae; palpus (Fig 1g) light yellow, bright silvery, long and narrow, strip-like, slightly curved, some brownish setae near inner margin (ratio tibia I: palpus, 22:21) (MSSC). Thorax. Mesonotum metallic green with coppery reflections, scutellum lighter with some bluish reflections; mesopleuron brown with some bluish reflections and little pruinosity. Wing. Halter light yellow, calypter brownish. Legs. I: 27, 22, 11, 4, 3, 2, 2. II: 35, 33, 13, 6, 4, 2, 2. III: 37, 37, 9, 9, 4, 3, 2. Coxa and femur I whitish yellow; coxa II, extreme base of coxa III and basal 1/2– 2/3 of tibia I light brown; apical 1/2–1/3 of femur III and apex of tarsi brown; legs otherwise yellow. Setae on coxae I and II white. Hypopygium. As in Fig 4l.

INB0003825918 and INB0003826237, and two males in MZSP, bar codes INB0003802861 and INB0003775303). Description Male. Body length, 1.4–1.6 mm. Wing length, 1.2–1.3 mm, width, 0.4–0.5 mm. Head. Frons brownish, with bluish reflections and silvery pruinose; face dark brown; antenna brown; proboscis brownish with pale setae, palpus (Fig 1h) long, straight and narrow, mostly light brown, covered with sparse setae, tip silvery pruinose, without setae (ratio tibia I: palpus, 26:27). Thorax. Mesonotum metallic green with coppery reflections, scutellum lighter; mesopleuron brown with some bluish reflections and little pruinosity. Wing. Halter light yellow, calypter brownish. Legs. I: 30, 27, 12, 6, 4, 3, 2. II: 32, 32, 15, 9, 7, 3, 2. III: 36, 35, 10, 11, 5, 3, 3. All coxae dark brown, coxa I lighter, apex of tarsi brown; apical 1/4 of femur III infuscated; legs otherwise yellow. Setae on coxae I and II pale to brownish. Hypopygium. As in Fig 4m. Female. Unknown. Etymology Named after the Costa Rican dipterist and colleague Manuel Zumbado (INBio). Distribution

Comments Costa Rica. Females of C. xiphostoma were not available for study, but Robinson (1975) described females from Dominica with palpus similar to that of other females in this group and coxa and femur I not as whitish yellow as in males. Distribution Dominica and Saint Lucia [!]. Chrysotus zumbadoi n. sp. (Figs 1h and 4m) Type material Holotype male, “Costa Rica. Prov. Puntarenas, PN | Corcovado, Cerro Puma, 100–300 m, | malaise trap, 19 Jun–8 Jul 2003, M. Moraga, | A. Azofeita, K. Caballero, Malaise #1, | L_S_267700_518900 #74506”, [bar code label], “Chrysotus | zumbadoi | Capellari HOLOTYPE [red label]” (INBio). Paratypes: three males, same data as holotype; two males, same data, but 10.i–9.iii.2003;five males, same data, but 200 m, 14.iii– 5.iv.2003; one male, same data, but 26.vi.2003, M. Moraga leg. (all INBio, but two males in CNC, bar codes

Discussion The monophyly of the longipalpus-group as here defined seems to be reasonably founded based on apomorphic characters of males (see Comments under the species-group section). As stated above, species of the longipalpus-group are recognized mostly by their distinctive male palpus, while remaining body parts show little variation. This situation is critical to assess phylogenetic relationships based on morphology, since the few characters that vary more consistently among the species also vary so much that it is difficult to recognize shared apomorphies. Nevertheless, the palpus divided into two parts (i.e., basally enlargement followed by a strap-like projection after abrupt tapering) can be synapomorphic for C. coquitos n. sp., C. miripalpus, and C. pachystoma n. sp. Molecular data can help to clarify the phylogenetic relationships within the longipalpus-group, but comments on the placement of the group within Chrysotus can still be made.

Review of the longipalpus-Group of Chrysotus

Grichanov (2012) described Chrysotus chukotkensis based on a single male specimen from Russia (Chukotka Peninsula), referring it to the albipalpus-group (sensu Van Duzee 1924), with species known both from Nearctic and Neotropical regions. Grichanov (2012) also noted that the albipalpusgroup and the longipalpus-group should be united based on the enlargement of the male palpus in both groups, providing an identification key for all species of these groups. I examined specimens of C. argentatus Van Duzee and C. halteralis Van Duzee (CNC) and checked descriptions of the remainder of the species of the albipalpus-group, and they do not share diagnostic features with the species of the longipalpus-group, such as the ventral swelling on male tarsus I, antero-dorsal setation on tibia III, and general color pattern. Actually, even the enlargement of the palpus is not so pronounced (shorter than face height in studied species) as it is in males of the longipalpus-group. Although the male palpus of C. chukotkensis is indeed enlarged (see Grichanov 2012: Fig 2), a comparative study of the species in the albipalpus-group is needed to check if that Russian species belong there. Thus, a close relationship between the two groups seems unlikely, since enlargement of the palpus probably arose independently in these lineages, as it did in several other species of Chrysotus. Males with an enlarged palpus also occur in the philtrumgroup (sensu Van Duzee 1924), which includes C. philtrum Melander, C. simulans Van Duzee, and C. tarsalis Van Duzee (all Nearctic). Van Duzee’s (1924) diagnosis for the group listed “[e]yes contiguous; femora and tibiae yellow; one pair of tarsi greatly compressed and ornamented”. Nevertheless, C. philtrum has a modified tarsus II (Melander 1903: Fig 1, Van Duzee 1924: Fig 44), while tarsus I is modified in C. tarsalis and C. simulans (Van Duzee 1924: Fig 2), which are obviously non-homologous conditions. I examined specimens of C. philtrum, C. tarsalis (CNC) and an undescribed species from Costa Rica close to C. tarsalis (INBio), and the philtrumgroup, as proposed by Van Duzee (1924), probably includes two unrelated clades: one with C. philtrum and other with C. simulans, C. tarsalis and the undescribed Costa Rican species. Males of this later group are characterized by the yellow palpus compressed and ovoid (assumed for C. simulans, described from a decapitated male) and tarsomeres 1–3 of leg I strongly flattened and enlarged. Although distinct, these characters could be homologous to those in the longipalpus-group, suggesting an exclusive ancestor for both groups. However, males of C. tarsalis and the undescribed Costa Rican species lack anterodorsal setae on tibia III and a ventral phallic projection, weakening the hypothesis that characters of the palpus and tarsus are synapomorphic for this group plus the longipalpus-group. At this time, it is better to keep C. simulans and C. tarsalis separated from the longipalpus-group and recognize the latter as a rather isolated clade within Chrysotus.

57 Acknowledgments Carlos Lamas (MZSP) (funded by CNPq-SISBIOTA FAPESP 2010/52314-0), Jeffrey Cumming (CNC), Manuel Zumbado (INBio), and Norman Penny (CAS) kindly facilitated loan of specimens. Harold Robinson, Norman Woodley (Smithsonian Institution), and Dan Bickel (Australian Museum) offered useful suggestions to the manuscript. Dan Bickel is also thanked for making available material of C. coquitos and kindly providing information on new records of C. crosbyi and C. longipalpus. Mónica Solórzano Kraemer (Senckenberg Forschungsinstitut und Naturmuseum) collected the specimens of C. coquitos, funded by the German Research Foundation (DFG, project SO894/3-1). Duncan Sivell (BMNH) provided useful information on type material, and Neal Evenhuis (BPBM) shared his data on the type series of C. pallidipalpus, allowing lectotype designation. Danilo Ament, Diego Porto, and Eduardo Almeida (FFCLRP) helped with the photos. This study benefited from FAPESP grants 2008/58224-3 and 2013/01392-0.

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