A revision of the Neotropical genus Chespiritos (Diptera: Sphaeroceridae: Limosininae)

The genus Chespiritos Marshall, 2000 (Diptera: Sphaeroceridae) is revised, including C. sindecimus Marshall, 2000, C. pervadens Roháček and Buck, 2003 and 19 new species. The following new species are described, keyed and placed in a morphological phylogeny: C. anguineus sp. n. from Bolivia, C. attenboroughi sp. n. from Argentina, C. balrogiformis sp. n. from Peru, C. bolanosi sp. n. from Mexico, C. calceus sp. n. from Peru, C. chicobrazos sp. n. from Costa Rica, C. coronatus sp. n. from Bolivia, Brazil, Colombia, Costa Rica, Ecuador and Trinidad, C. dolabratus sp. n. from Costa Rica, C. elephantus sp. n. from Ecuador and Peru, C. extendido sp. n. from Bolivia, Colombia, Costa Rica, Ecuador, Mexico, Panama and Venezuela, C. ganchopico sp. n. from Argentina, Bolivia, Ecuador, Panama and Tobago, C. gladiator sp. n. from Costa Rica, C. hojagrande sp. n. from Costa Rica and Mexico, C. jamaicensis sp. n. from Jamaica, C. lepustergum sp. n. from Costa Rica and Venezuela, C. metroidiformis sp. n. from Costa Rica, C. paraiso sp. n. from Dominican Republic, C. peckorum sp. n. from Ecuador, and C. ventrisetis sp. n. from Brazil, Costa Rica and Venezuela. ZooBank Article Registration: http://zoobank.org/urn:lsid:zoobank.org:pub:80DCB50F-37BD-4164-8D51-80B05FEC7428


INTRODUCTION
Chespiritos Marshall, 2000 is unique among sphaerocerid genera for the combination of a broad head, bulging lunule, three pairs of interfrontal setae (one small and two large), and six marginal scutellar setae (four large setae and two smaller basal setulae). This genus is further characterized by the presence of two pairs of dorsocentral setae, a costa that extends far beyond R 4+5 , and the unusual complete absence of the female epiproct (tergite 10). There are currently two described species of Chespiritos, C. sindecimus Marshall, 2000 from Costa Rica and C. pervadens Roháček & Buck, 2003 from Brazil and the Canary Islands. The genus is mostly neotropical, ranging from southern Mexico to northern Argentina at elevations from sea level to 2100 m although most records are from above 800 m. We here revise the genus with the description of 19 new Neotropical species, of which eight are known only from Central America, six are known only from South America, and fi ve are known from both regions.
Little is known of the natural history of Chespiritos, though species are generally associated with decaying plant material. For example, Chespiritos ganchopico sp. n. has been reared and collected from Heliconia Linnaeus, 1771 (Heliconiaceae) in Ecuador and Panama and specimens of C. sindecimus were originally collected from bromeliads -Field Museum of Natural History, Chicago, Illinois, USA; LACM -Los Angeles County Museum, Los Angeles, California, USA; MIZA -Museo del Instituto de Zoología Agrícola, Maracay, Venezuela; MNCR -Museo Nacional de Costa Rica, San José, Costa Rica; MUSM -Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru; MZSP -Museu de Zoologia, Universidade de São Paulo, Brazil; QCAZ -Museo de Zoología, Pontifi ca Universidad Catolica del Ecuador, Quito, Ecuador; USNM -United States National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia, USA. Marshall (2000) suggested Sclerocoelus Marshall, 1995 as a possible sister group to Chespiritos on the basis of the shared broad lunule and similar small setae in the intra-alar or posterior supra-alar position. Roháček & Buck (2003) disputed the latter character as a possible generic synapomorphy, but genitalia of these genera provide some evidence for a close relationship between Sclerocoelus and Chespiritos. The probable ground plan condition of the surstylus in Chespiritos, as refl ected in the basal Chespiritos species C. jamaicensis sp. n. and C. calceus sp. n., is similar to that found in Sclerocoelus. Both groups have distinctive ventromedially cleft surstyli as illustrated for Sclerocoelus hemorrhoidalis Marshall, 1997and S. brasilensis Marshall, 1997in Marshall (1997. We therefore root our phylogenetic analysis of Chespiritos with Sclerocoelus (represented in the matrix by S. hemorrhoidalis plus S. brasilensis) as the outgroup.

Similar genera and relationships
Pullimosina (Dahlimosina) darwini Richards, 1931 is superfi cially similar to most Chespiritos in having a small pair of basal scutellar setulae, the costa extending well past with this sclerite it is distinctly separated from and oriented perpendicular to sternite 5. We therefore refer to it here as a separate sclerite.
Abbreviations include S -sternite(s); T -tergite(s). Label data are given in a standardized format rather than given verbatim from labels.
Male terminalia were prepared by soaking entire abdomens in room temperature 10% potassium hydroxide for fi ve minutes prior to heating the potassium hydroxide to boiling point for 15-30 min. Cleared abdomens were subsequently neutralized in glacial acetic acid for at least 30 min, then rinsed with deionized water and placed in glycerin for examination. All dissected parts of the specimens were stored in glycerin in genitalia vials pinned below the specimens.
Illustrations were created via free-hand drawings made while viewing specimens through a Leica Wild M3Z stereo microscope or a Nikon Labophot compound microscope. The illustrations were then compiled and edited using Photoshop Elements 9 (Adobe Services, CA, USA).
Measurements were taken from ten specimens of common species or all specimens from rarer species, from a variety of locations when possible. Measurements are approximate because dried specimens are subject to varying degrees of shrinkage and distortion. Body length measurements were taken from the tip of the head (lunule) to the tip of the abdomen.
Acronyms of depositories. Specimens are deposited in the following institutions: BMNH -Natural History Museum (formerly British Museum of Natural History), London, UK; CBFC -Colección Boliviana de Fauna, La Paz, Bolivia; CNCI -Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario, Canada; DEBU -School of Environmental Sciences, University of Guelph, Guelph, Ontario, Canada; FMNH Table 1. Character state matrix for Neotropical species of Chespiritos, including two species of Sclerocoelus as the outgroup. Plesiomorphic states are indicated by (0) and apomorphic state(s) by (1-2). Multistate characters are treated as unordered.

Character state matrix
The plesiomorphic state is indicated by (0) and the apomorphic state(s) by (1-2). Multistate characters are treated as unordered. Two equally parsimonious trees (length 87, consistency index 0.41, retention index 0.66), differ only in the relationships between three small groups: the C. sindecimus group, C. ganchopico and C. extendido, and the C. dolabratus group. The preferred tree (Fig. 27) places the C. dolabratus species group as sister to C. ganchopico and C. extendido based on the unique synapomorphy of a long medial connecting sclerite (character 19) The most strongly supported clade within Chespiritos is the C. ventrisetis species group. This group is characterized by the presence of large anterior lobes and a dense, posterior setal brush on the male fi fth sternite (characters 4 & 5, though C. chicobrazos sp. n. has a weak setal brush), postgonites with tooth-like posterior lobes originating from the basal third (character 14) and a hypandrium with long posteromedial arms (character 20). The C. ventrisetis group is distributed across Central and South America, with the widespread C. ventrisetis sp. n. found as far south as the southern Brazilian state of Santa Catarina. Chespiritos ventrisetis sp. n., C. chicobrazos sp. n. and C. gladiator sp. n. have all been collected from the Monteverde Biological Reserve in Costa Rica.
The C. sindecimus species group is characterized by the presence of posteromedial lobes on the male S5 (character 7) and large, leaf-like surstyli. The C. sindecimus group occurs from Mexico to southern Brazil, with C. coronatus having the largest known range in the group.
The C. dolabratus species group is characterized by a medially short male S5 with long posterolateral lobes (character 6) and distinctively shaped postgonites with the apical quarter constricted. Additionally, females of this group have paired spermathecae with smooth bulbs and a single spermatheca with a grooved bulb (character 27). The C. dolabratus group is known from Mexico, Costa Rica and the Dominican Republic, with each species recorded from only a single country.

Distribution
Most species are known from only one or two countries, but a few are more widespread and some, such as C. ganchopico sp. n., C. extendido sp. n., and C. coronatus sp. n., occur in both Central and South America. Chespiritos extendido sp. n. is the only species currently known from the Galápagos Islands. Chespiritos pervadens, recorded from the Canary Islands as well as Brazil, is the only species of the genus known from outside the Neotropics; Roháček and Buck (2003) suggest that the species was introduced to the islands from South America. Costa Rica seems to have the highest diversity of species of Chespiritos, with 10 species, but the Costa Rican fauna has been more extensively sampled than that of other Neotropical countries. Typically, only one or two species of Chespiritos are collected from any given locality, but some locations in Costa Rica, such as the Monteverde Biological Reserve in Puntarenas and Pandora in Limón, have yielded several species.
C. extendido sp. n. and C. coronatus sp. n. are both found at an extremely wide range of elevations and in a variety of habitats, with C. extendido sp. n. having been collected at altitudes of 250 m to 2100 m above sea level, in habi-tats ranging from rainforests to cloud forests. Specimens have been taken in a variety of collection devices including dung traps, carrion traps, compost pan traps and Malaisefl ight intercept traps. Chespiritos coronatus sp. n. has been collected at elevations ranging from 160 m to 2000 m. Unlike C. extendido sp. n. and C. coronatus sp. n., most Chespiritos species are known from a single country, and often a single locality. Narrowly distributed species in the genus include C. jamaicensis sp. n. from Jamaica, C. balrogiformis sp. n. from Peru and C. attenboroughi sp. n. from Argentina. Head. Very broad, frontal width ~ 3.0 × interfrontal height. 2 pairs of large interfrontal setae preceded by 1 pair of smaller setae; 2 large lateroclinate orbital setae; ocellar setae large and diverging; postvertical and postocellar setae small; inner occipital seta large; vibrissa very large with two distinct subvibrissal setae. Clypeus large, dark and often exposed, lunule rounded and bulging. Palpus clavate, ventrally setose with a distinct preapical ventral seta. Orbital plate, interfrontal stripe and ocellar triangle with silver pollinosity. First fl agellomere compressed; arista pubescent, dorsobasal, ~ 2.0 × height of head. Compound eye with shallow notch along anterior edge, posteroventral edge straight; eye height ~ 2.0-2.5 × genal height.
Legs. Fore femur with 1 row of long ventral setae. Basal half of mid tibia with 6-8 dorsal setae including 2-3 small and 1 large anterodorsal setae, and 3-4 small posterodorsal setae above 1 at least slightly stouter posterodorsal seta. Distal half of mid tibia with 7 dorsal setae (2-3 anterodorsal, 2 dorsal and 2-3 posterodorsal). Mid tibia with large apicoventral and anteromedial setae, distal third of male mid tibia with two rows of stout setae ventrally; proximal quarter of mid femur with two rows of 3-5 corresponding ventral setae. Mid basitarsus with a prominent apicoventral seta. Mid coxa with three distinct lateral setae.
Wing. Elongate, ~ 2.5 × long as wide; third costal sector slightly shorter than second costal sector; costa basal to subcostal break with strong setae and a large costagial seta; R 4+5 upcurved, meeting costa well before wing tip; costa extending far beyond R 4+5 (~ 7 × costal width); distance between crossveins dm-cu and r-m ~ 2.0-3.0 × the length of crossvein dm-cu; both M and CuA 1 extending slightly beyond dm cell; alula narrow.
Male abdomen. S3-4 unmodifi ed and well sclerotized. S6 and S7 expanded and fused dorsolaterally to form a genital pouch; left side of S6+7 wrapped underneath S5 and connected via the medial connecting sclerite, typically bearing two stout setae (Fig. 8); sclerite may not be visible in some species. Cerci typically rounded, pad-shaped and medially fused, with 1 or 2 larger setae and numerous smaller setulae. Hypandrium typically elongate and sinuate; cleft posteriorly with a set of forked arms: lateral arms extending to and continuous with ventrolateral corners of epandrium and posteromedial arms small, extending posteriorly to surround the anterior half of the distiphallus. Ejaculatory apodeme simple, S-shaped and largely desclerotized. Basiphallus stout and wedge-shaped, epiphallus often expanded and hooked giving the basiphallus an axeblade shape. Distiphallus in two distinct sections, a sclerotized and tubular proximal half, and a membranous distal half with some supporting sclerites. Phallapodeme long, dark and slightly sinuate with a pair of posterior lobes and a small ventral lobe just prior to the paired lobes; phallapodeme articulating with the basiphallus around these three lobes.
Female abdomen. Tergites uniformly setose with long posterolateral setae and a sparse posterior setal brush. T7 with a pale posteromedial notch and a small anterolateral lobe. T8 slightly desclerotized dorsomedially, with small anterolateral and posterolateral lobes; posterolateral lobe often slightly separated from main body of T8. Epiproct absent. Cercus short and tapered, with 3 small medial setae, 1 large apical seta and a smaller curved preapical dorsal seta. S3-7 uniformly setose with sparse posterior setal brush. S7 large and shield-like; S8 reduced to a pair of rounded plates normally mostly hidden by S7 in ventral view, each with a pair of setae. Vaginal sclerite semicircular with a distinct posteromedial process; hypoproct reduced to a pair of indistinct plates located just under the cerci. Spermathecae (2+1) mushroom-shaped: apical section ovoid or spherical, basal section forming a conical stem connecting to the sclerotized distal parts of the spermathecal ducts; apical section of single spermatheca sometimes different in shape or texture from that of paired spermathecae. Pleural membrane uniformly setose.

Key to species
This key is based almost entirely on features of the male terminalia as most species are otherwise indistinguishable morphologically. distinctly fl attened apical seta and dense lateral setae but without a dense patch of microtrichia apically (   smaller setae in addition to large medial seta; surstylus with a broad anterobasal lobe and a triangular preapical lobe, resembling a sock ( Fig. 17.2); postgonite broadly emarginate apically (

Species descriptions
The following species descriptions are almost entirely restricted to the male and female terminalia. Other external characters were carefully examined and, with the exception of a few features included in the species descriptions, did not differ signifi cantly between species. Species descriptions are arranged in the same order the species appear on the phylogenetic tree (Fig. 27). Male abdomen. S5 with scattered setae, a pair of triangular, long-setose posterolateral lobes each terminating in thin, curved extensions; posterolateral lobes connected to main body of S5 via a membranous sheet posteriorly edged by a thin sclerite with a pair of reclinate setae; S5 medially short, length ~ 0.20 × width. Epandrium uniformly longsetose. Cercus pad-like. Surstylus short, long-setose, shoeshaped with a rectangular lobe extending posteriorly. Postgonite dark, fl attened, curved forward along entire length, apex broadened and truncate. Basiphallus stout, rectangular; epiphallus small, rounded and directed ventrally. Distiphallus with basal third sclerotized and tubular, apical two-thirds membranous with a broad, scoop-like (when viewed dorsolaterally) dorsal lobe and a ventral lobe distinctly curved downwards, dorsoapical quarter reticulated.
Female unknown. Distribution. This species is known only from its type locality in Peru. Male abdomen. S5 uniformly setose with 1 pair of incurved posterolateral lobes, each with numerous stout setae; lobes connected by a membranous sheet with 2 me-dial setae; S5 medially long, length ~ 1.0 × width. Epandrium uniformly long-setose. Cercus pad-like with triangular ventral margin. Surstylus distally bilobed, anterior lobe shorter than posterior lobe, both with patches of setae. Postgonite dark, decurved, medially expanded and apically bifurcated. Basiphallus large, rectangular; epiphallus triangular, rounded apically and directed posteriorly. Distiphallus with basal half heavily sclerotized and tubular, apical half membranous with a large, scoop-like ventral lobe and a triangular dorsal lobe.
Female unknown. Etymology. This name refl ects the similarities between the fi fth sternite of Chespiritos metroidiformis and the bulbous body and large mandibles of the alien species "Metroid" from Nintendo's video game series of the same name.
Distribution. This species is known only from its type locality in Costa Rica. Male abdomen. S5 large, broad, with a pair of short, bare posteromedial lobes that curl upwards slightly; S5 medially long, length ~ 0.50 × width. Medial connecting sclerite stout, posteriorly emarginated with a pair of stout setae. Epandrium uniformly long-setose with a large ventrolateral seta. Cercus pad-like. Surstylus complex, with two distinct ventral lobes: long-setose basal lobe extending to a rounded point and bare apicoventral lobe extending to a sharp point. Postgonite dark, tapered and decurved, anterior edge slightly sinuate, basal half with a broad lateral lobe, apex with rounded lateral lobe. Basiphallus stout, rectangular; epiphallus large, curved and directed posteroventrally. Distiphallus with basal third heavily sclerotized and tubular with sclerotization extending dorsally to apex, apical two-thirds highly membranous with a large, bilobed, scoop-like lobe.
Female unknown. Male abdomen. S5 broad with 2 small incurved longsetose posterolateral lobes; posterior edge with a dense brush-like band of setae; S5 medially short, length ~ 0.20 × width with a single large, anteromedially emarginate anterior lobe. Medial connecting sclerite thin, inverted V-shaped and bearing a pair of setae; hidden under posterior edge of S5. Epandrium uniformly long-setose with 1-2 large ventrolateral setae. Cercus pad-like. Surstylus elongate, long-setose and tapered with apical third slightly angled posteriorly. Postgonite dark, slightly decurved, blade-like with anterior margin slightly sinuate and a small, distally projecting lobe originating from the posterior edge of the basal third. Hypandrium unique: anterior lobe long, posterior fork of arms distinctly long, about as long as anterior fork of arms. Basiphallus stout, square; epiphallus extremely large, curved and directed posteriorly. Distiphallus with basal half heavily sclerotized and tubular, apical half membranous, somewhat tubular and angled downwards with a slightly bifurcated apex.
Female abdomen. T7 with 4 pairs of smaller setae and 2 pairs of larger posterolateral setae; width ~ 1.50 × width of T3; lacks posteromedial notch but has rounded anteromedial notch, extending posteriorly ~ 0.17 × length of T7, anterior width ~ 0.33 × width of T7; anterolateral lobe absent. T8 with 7 pairs of smaller setae; both antero-and posterolateral lobes triangular and connected from T8, posterolateral lobe elongate. Apical bulb of spermathecae stout, spherical, grooved with apical invagination, ~ equal to length of smooth, stout basal bulb. Distribution. This species is widespread but has an apparently disjunct distribution, with no specimens having been collected between the southern end of Brazil and Venezuela.
Chespiritos chicobrazos sp. n. Male abdomen. S5 with a pair of broad, long-setose posterolateral lobes projecting downwards and angled inwards; S5 medially short, length ~ 0.25 × width with a single large, anteromedially emarginate anterior lobe. Epandrium uniformly long-setose with a ventrolateral patch of large setae. Cercus pad-like. Surstylus rectangular, long-setose with a long, thin apical projection angled posterolaterally. Postgonite dark, narrowed in apical third with a small, distally-projecting lobe originating from the posterior edge of the basal third. Basiphallus large, ventromedially emarginate resembling a kidney bean; epiphallus small, rounded and directed posteriorly. Distiphallus with basal two-thirds heavily sclerotized and tubular, apical third highly membranous with a large, laterally folded, scoop-like lobe.
Distribution. This species is known only from its type locality in Costa Rica.
Male abdomen. S5 with a pair of stout, incurved longsetose posterolateral lobes projecting downwards; S5 medially short, length ~ 0.33 × width with a single large, anterior lobe; posteromedial edge of S5 with a dense patch of setae. Medial connecting sclerite stout, posteriorly emarginated and bearing a pair of stout setae; hidden under posterior edge of S5. Epandrium uniformly longsetose with a large ventrolateral seta. Cercus with a slight ventromedial indentation. Surstylus elongate, long-setose and tapered with apex angled inwards; anterior edge with a row of long setae; apical half with a sparse patch of setae. Postgonite dark, slightly decurved, and blade-like with a small, distally projecting lobe originating from the posterior edge of the basal third. Hypandrium unique: anterior lobe with spatulate apex; anterior fork of arms broad and recurved apically, posterior fork of arms distinctly long, about half as long as anterior arms. Basiphallus stout, triangular; epiphallus small, triangular and directed posteriorly. Distiphallus with basal two-thirds heavily sclerotized and tubular, apical third membranous with a pair of smaller dorsolateral lobes and a large ventral lobe.
Etymology. This name refers to the fi fth sternites, which look like the galea-style helmet worn by Roman gladiators (Latin 'gladiator': swordsman). The posterolateral lobes resemble the plates of the helmet that cover the sides of the face.
Distribution. This species is known only from Costa Rica. Male abdomen. S5 complex, with a pair of large, rectangular, long-setose posterolateral lobes angled inwards and bare posteromedial lobe wrapping under posterolateral lobes; posteromedially with 2 stout setae; S5 medially short, length ~ 0.20 × width. Epandrium uniformly long-setose. Cercus pad-like. Surstylus elongate, triangular with a small triangular anteromedial lobe; anterior surface with numerous long setae, laterally with some small setae. Postgonite tapered, with anterior margin sinuate and apical third angled posteriorly. Basiphallus rectangular; epiphallus small, rounded and directed ventrally. Distiphallus with basal half heavily sclerotized and tubular, apical half semimembranous with 2 elongated dorsal and ventral sclerites supporting 3 scoop-like, apicolaterally reticulated lobes which form a tube.
Female unknown. Male abdomen. S5 with 2 large, long-setose posterolateral lobes, each tipped with a distinctly larger, fl attened seta; S5 medially short, length ~ 0.10 × width. Medial connecting sclerite very broad, triangular and bearing a pair of stout setae. Epandrium uniformly long-setose with 2 large ventrolateral setae. Cercus with large, ovular, long-setose ventral lobe. Surstylus elongate, thin, anteriorly long-setose with apical third bent posteriorly 90°and tipped with stout seta. Postgonite dark, slightly sinuate along anterior Etymology. This name refers to the very large setae on the posterolateral lobes of the fi fth sternite, which resemble the fangs of a snake (Latin 'anguineus': having the appearance of a snake).
General. Eye height ~ 2.5 × genal height. Mid femur with ventral row of 5 setae on proximal quarter. Distance between crossveins dm-cu and r-m ~ 2.5 × length of dm-cu.
Female abdomen. T7 with 6 pairs of smaller setae and 1 pair of larger posterolateral setae; width ~ 1.0 × width of T3; posteromedial notch large, squared, extending anteriorly ~ 0.40 × length of T7, posterior width ~ 0.16 × width of T7; anterolateral lobe absent. T8 with 3 pairs of smaller setae and narrow, desclerotized medial band; anterolateral lobe elongate, triangular and connected to T8, posterolateral lobe large, triangular and separated from T8. Apical bulb of spermathecae stout, spherical and weakly grooved, apically invaginated, slightly shorter than smooth, elongate basal bulbs. Distribution. This species is known from Brazil as well as the Canary Islands.
Female abdomen. T7 with 4 pairs of smaller setae and 2 pairs of larger posterolateral and medial setae; width ~ 1.25 × width of T3; posteromedial notch very small, extending anteriorly ~ 0.13 × length of T7, posterior width ~ 0.33 × width of T7; anterolateral lobe small and rounded. T8 medially desclerotized with 6 pairs of small lateral setae and wide, desclerotized medial band; anterolateral lobe small, triangular and connected to T8, posterolateral lobe large, triangular and separated from T8. Apical bulb of spermathecae elongate and grooved, ~ equal to length of smooth, elongate basal bulb. Type locality. Argentina, Salta Province, El Rey National Park.
Etymology. This name is in honour of Sir David Attenborough, whose programs about, and advocacy of, wildlife conservation helped inspire the fi rst author to pursue a career in the biological sciences.
Distribution. This species is known only from its type locality in Argentina.
Female unknown.
Etymology. This name is in honour of Dr. Stewart B. Peck and Dr. Jarmila Kukalová-Peck who provided us with insect trap samples that include nine Chespiritos species, including this one.
Distribution. This species is known only from its type locality in Ecuador.
General. Eye height ~ 2.5 × genal height. Mid femur with ventral row of 4 setae on proximal quarter. Distance between crossveins dm-cu and r-m ~ 2.5 × length of dm-cu.
Male abdomen. S5 complex, main body bare with a large medial cleft bordered by a pair of small, bare postero-medial lobes and a pair of elongate, long-setose posterolateral lobes; S5 medially short, length ~ 0.25 × width. Medial connecting sclerite stout, posteriorly bifurcated with a pair of strong setae; may be hidden under posterior edge of S5. Epandrium large, uniformly long-setose with a long ventrolateral seta. Cercus pad-like. Surstylus large, leaf-like, anterior edge with a row of stout setae along entire length and a lateral patch of fi ner setulae in basal half. Postgonite dark, decurved, apex broadly rounded and serrated with a small tooth-like projection on apicoventral edge. Basiphallus elongate, rectangular; epiphallus large, rounded and directed ventrally. Distiphallus very large, basal two-thirds heavily sclerotized and tubular, apical third semi-membranous with 2 large sclerites supporting 3 lobes which form a tube, ventral lobes ventrolaterally reticulated.
Female abdomen. T7 with 3 pairs of smaller setae and 1 pair of larger posterolateral and 1 pair of medial setae; width ~ 1.25 × width of T3; posteromedial notch triangular, extending anteriorly ~ 0.25 × length of T7, posterior width ~ 0.20 × width of T7; anterolateral lobe large and rounded, angled downwards. T8 with 5 pairs of smaller setae and large, desclerotized posteromedial triangle; anterolateral lobe highly reduced and connected to T8, posterolateral lobe large, triangular and separated from T8. Apical bulb of spermathecae stout, spherical and grooved, ~ equal to length of smooth, stout basal bulb. Male abdomen. S5 complex, main body uniformly long-setose with a large medial cleft bordered by a pair of small, bare posteromedial lobes fl anking and pair of elongate, long-setose posterolateral lobes, posterolateral lobes connected via a membranous sheet with dense microtrichia; S5 medially short, length ~ 0.20 × width. Medial connecting sclerite stout, medially constricted with a pair of apicolateral projections, each terminating in a strong seta. Epandrium uniformly long-setose with a ventrolateral pair of large setae. Cercus pad-like. Surstylus very large, leaflike with a row of 5 or 6 stout setae along anterior edge and fi ner setae laterally; apex with a small lateral lobe. Postgonite dark, decurved, apically broad and slightly emarginated. Basiphallus elongate, rectangular; epiphallus broad, rounded and directed posteriorly. Distiphallus with basal half heavily sclerotized and tubular, apical half semi-membranous with a sclerotized dorsal lobe and 2 membranous, laterally reticulated ventrolateral lobes supported ventrally by elongate, tapered sclerites which curl upwards apically.
Female unknown.  medially short, length ~ 0.16 × width. Medial connecting sclerite small, elongate with a pair of short mediolateral projections, each terminating in a stout seta; hidden under posterior edge of S5. Epandrium uniformly long-setose with larger posterolateral seta. Cercus pad-like. Surstylus very large, leaf-like, anterior edge with a shallow emargination across middle half; anterior edge with a row of long setae along anterior edge and fi ner setae laterally. Postgonite dark, elongate, sinuate along anterior margin with apex broad, ventrally fl attened and angled anteriorly. Basiphallus elongate, triangular; epiphallus small, square and directed posteroventrally. Distiphallus with basal two-third heavily sclerotized and tubular, apical third semi-membranous with sclerotized dorsal lobe and 2 membranous, laterally reticulated ventrolateral lobes supported ventrally by elongate, tapered sclerites that curl upwards apically.
Female abdomen. T7 with 4 pairs of smaller setae and 1 pair of larger posterolateral setae; width ~ 1.25 × width of T3; posteromedial notch semicircular, extending anteriorly ~ 0.25 × length of T7, posterior width ~ 0.16 × width of T7; anterolateral lobe small, angled anteriorly. T8 with 5 pairs of smaller setae and small, desclerotized posteromedial triangle; anterolateral lobe triangular and connected to T8, posterolateral lobe rectangular and separated from T8. Apical bulb of spermathecae stout, spherical and grooved, ~ equal to length of smooth, triangular basal bulb. Distribution. This widespread species is known from southern Brazil to Central America. Description Body length. Males 1.9-2.7 mm, females 2.1-2.7. General. Eye height ~ 2.5 × genal height. Mid femur with ventral row of 4 setae on proximal quarter. Distance between crossveins dm-cu and r-m ~ 2.5 × length of dm-cu.
Male abdomen. S5 with scattered setae and a small medial cleft bordered by a pair of very small, bare posteromedial lobes and a pair of long-setose posterolateral lobes; S5 medially short, length ~ 0.25 × width. Medial connecting sclerite short, posterior half gradually expanded and slightly emarginate with a pair of short, lateral projections, each terminating in a strong seta. Epandrium uniformly longsetose. Cercus pad-like. Surstylus very large, leaf-like, anterior edge with a shallow emargination across middle half; anterior edge with long setae along entire length and smaller setae laterally. Postgonite dark, elongate, sinuate along anterior margin with apex broad, ventrally fl attened with two ridges extending laterally, forming a rubber-stamp-like shape. Basiphallus elongate, rectangular; epiphallus small, triangular and directed posteroventrally. Distiphallus with basal two-thirds large, heavily sclerotized and tubular, apical third semi-membranous with a sclerotized dorsal lobe and 2 membranous, laterally reticulated ventrolateral lobes supported by elongate, tapered sclerites which curl upwards apically. Female abdomen. T7 with 3 pairs of smaller setae and 1 pair of larger posterolateral setae; width ~ 1.25 × width of T3; posteromedial notch semicircular, extending anteriorly ~ 0.16 × length of T7, posterior width ~ 0.33 × width of T7; anterolateral lobe rounded, angled anteriorly. T8 with 7 pairs of smaller setae and small, desclerotized posteromedial triangle; anterolateral lobe broadly triangular and connected to T8, posterolateral lobe large, rectangular and separated from T8. Apical bulb of spermathecae stout, spherical and grooved, ~ equal to length of smooth, stout basal bulb. Etymology. This name refers to the very long surstyli, which cause the posterior view of the terminalia to resemble the head of a rabbit (Latin 'lepus': rabbit, hare; Latin 'tergum': back, rear).
Distribution. This species is known from Costa Rica and Venezuela. Male abdomen. S5 setose with a pair of broad, triangular, long-setose posterolateral lobes, lobes connected via a membranous sheet with dense microtrichia; S5 medially short, length ~ 0.20 × width. Medial connecting sclerite elongate, apically bifurcated with a pair of setae. Epandrium uniformly long-setose. Cercus pad-like. Surstylus triangular with a row of 5 or 6 stout setae along anterior edge and a large discal patch of fi ner setae. Postgonite dark, tapered, sinuate, apex with slightly emarginate. Basiphallus elongate, rectangular; epiphallus large, decurved and directed posteroventrally. Distiphallus with basal twothirds heavily sclerotized and tubular, apical third semimembranous with 3 scoop-like, apicolaterally reticulated lobes which form a tube.
Male abdomen. S5 with large central cleft bordered by a pair of small, bare posteromedial lobes and pair of large, long-setose posterolateral lobes, lobes connected via a membranous sheet with dense microtrichia; S5 short, length ~ 0.20 × width. Medial connecting sclerite elongate, posterior quarter bulbous with two small apical lobes, 'bulb' with a pair of lateral projections, each terminating with a seta. Epandrium uniformly long-setose. Cercus padlike. Surstylus leaf-like with a row of 5 stout setae along anterior edge and a patch of fi ner setae laterally; distal half with a large triangular lateral lobe. Postgonite dark, elongate, sinuate along anterior margin with a small preapical posterior notch. Basiphallus elongate, rectangular; epiphallus small, rounded and directed posteroventrally. Distiphallus with basal half heavily sclerotized and tubular, apical half semi-membranous with 2 large sclerites supporting 3 scoop-like, apicolaterally reticulated lobes which form a tube.
Male abdomen. S5 with a pair of large, apically incurved long-setose posterolateral lobes, lobes connected via a membranous sheet with dense microtrichia; S5 medially short, length ~ 0.14 × width. Medial connecting sclerite elongate, posterior third expanded with a pair of apicolateral projections, each terminating in a strong seta. Epandrium uniformly long-setose. Cercus with a slight ventral indentation. Surstylus rectangular with a row of stout setae along anterior edge and a small patch of fi ner discal setae near the base; basal two-thirds of posterior edge straight, apical third gently sinuate; apex with a small, blade-like lobe. Postgonite dark, broad, sinuate along anterior margin with a large anterobasal notch, dramatically narrowed about apical quarter. Basiphallus semicircular; epiphallus reduced to a rounded lobe. Distiphallus large, with basal half heavily sclerotized and tubular, apical half semi-membranous with 2 elongate sclerites supporting 3 scoop-like, apicolaterally reticulated lobes which form a tube.
Female abdomen. T7 with 4 pairs of smaller setae and 1 pair of larger posterolateral setae; width ~ 1.25 × that of T3; posteromedial notch small, semicircular, extending anteriorly ~ 0.16 × length of T7, posterior width ~ 0.33 × width of T7; anterolateral lobe large and rectangular, angled anteriorly. T8 with 7 pairs of smaller setae and a small, desclerotized posteromedial cleft; anterolateral lobe small, rounded and connected to T8, posterolateral lobe small, rectangular and separated from T8. Apical bulb of single spermatheca elongate, conical and grooved, slightly longer than length of smooth, tapered basal bulb; apical bulbs of paired spermathecae stout, spherical and smooth, slightly less than length of smooth, elongate basal bulbs. Type locality. Dominican Republic, Barahona Province, 7 km NW Paraiso.
Etymology. This species name is a noun in apposition based on the name of the type locality of the species, the town of Paraíso in the Dominican Republic.
Distribution. This species is only known from the Dominican Republic.   Table 1. The cladogram was rooted by the two species of Sclerocoelus. Solid circles indicate unique and unreversed synapomorphies, empty circles indicate homoplasies and crossed circles indicate reversals. Ones and twos below the circles indicate multistate characters (unordered), zeroes below the circles indicate plesiomorphic characters.