New genus and species of flea beetles (Coleoptera, Chrysomelidae, Galerucinae, Alticini) from Puerto Rico, with comments on flea beetle diversity in the West Indies and a key to the West Indian Monoplatini genera

Abstract A new genus (Borinken) and five new species (Borinken elyunque, Distigmoptera chamorrae, Kiskeya elyunque, Ulrica eltoro, and Ulrica iviei) from Puerto Rico are described and illustrated. A keyto all West Indian Monoplatini genera is provided, as are keys to all speciesof Kiskeya and to the speciesof Ulrica from Puerto Rico. A list of the flea beetle genera, along with the number of species and some of the faunal features is presented and discussed for the West Indies.


Introduction
The West Indies Islands are one of the World's biodiversity hotspots (Myers et al. 2000). A great variety of ecosystems exist in the West Indies, ranging from tropical moist broadleaf forests to xeric cactus scrublands. The West Indian flora and fauna are rich and highly endemic. Seventy-two percent of 11,000 plant species of the West Indies are endemic (BirdLife International 2009). Among vertebrates, 99% of amphibians and 93% of reptiles are endemic (Hedges 2001). To date, 351 species of flea beetles in 53 genera are known to occur in the West Indies (Table 1). Compared to other regions of the World, these relatively small land masses are home to considerable species richness. For example, a much larger territory of the European part of the former U.S.S.R. (stretching from the White Sea on the North to the Black Sea on the South, including the Caucasus) has 321 species in 24 genera of flea beetles; none of the genera are endemic, and only 20% of the species are endemic (Konstantinov 1991). In the West Indies, 92% of flea beetle species and 20% of genera (marked bold in the Table 1) are endemic. In addition, only 13% of species occur on more than one island. Epitrix parvula Fabricius is the most widespread flea beetle species, being found on most West Indian Islands and in North and Central America. While the number of flea beetle species in the European part of the former U.S.S.R., including the Caucasus, and the West Indies are similar (321 and 351 respectively), the number of genera in the West Indies is more than twice that in the European part of the former U.S.S.R. and the Caucasus, which makes the fauna of the significant part of the Palearctic more of an "island" fauna than the island fauna itself ). As a rule, an island fauna is characterized by a relatively large number of species per higher taxon as a result of a relatively small number of introductions followed by extensive specieslevel radiation (Magnacca and Danford 2006). The fauna of the European part of the former U.S.S.R. (being part of the Palearctic) was recently dramatically changed by the Tertiary aridization and Quaternary glaciation ), while the West Indian fauna remained relatively intact.
The flea beetles of the West Indies are comparatively well studied. Extensive collecting and publications during the first half of the 20 th and early 21 st century reported many unusual and endemic flea beetle taxa (Blake 1928, 1931, 1934, 1937, 1938, 1944, 1947, 1960, 1964, Konstantinov 2002, Konstantinov and Chamorro-Lacayo 2006. The three most species rich genera in the West Indies are Aedmon Clark with 36 species, Homoschema Blake with 26 species, and Monomacra Chevrolat with 22 species. Two of these most speciose genera are West Indian endemics. The distribution of species in the endemic genera reveals some aspects of the faunistic relationships between different islands (Table 1). For example, the moss-inhabiting Kiskeya Konstantinov and Chamorro-Lacayo has three species in the West Indies, two in the Dominican Republic and one in Puerto Rico. Kiskeya is morphologically similar to the Oriental Clavicornaltica Scherer, alluding to broader biogeographical patterns of moss and leaf litter inhabiting flea beetles. Normaltica Konstantinov has a similar pattern with one species in the Dominican Republic and one in Puerto Rico. The apparent absence of  Kiskeya and Normaltica in Cuba probably reflects more a lack of collecting in that largest West Indian Island than to actual biogeographical patterns. Three currently known species of Monotalla Bechyné (two of which are undescribed) occur only in the Lesser Antilles. The only two species of Bonfilsus Scherer occur in Hispaniola and Guadeloupe (one species per island). Recent collecting efforts in Puerto Rico revealed unique flea beetles with several features rarely observed among flea beetle genera.

Materials and methods
Various collecting methods were used in Puerto Rico. Among them, beetles from sifted and unsifted moss samples by Berlese extraction represent most of the new taxa described here. Collecting in moss cushions was one of the most effective methods for uncovering a previously unknown fauna.
Dissecting techniques and terminology for most internal and external structures follow Konstantinov (1998Konstantinov ( , 2002. Terminology for the thoracic structures follows Konstantinov and Lopatin (1987) and Chamorro-Lacayo and Konstantinov (2004). We follow a format in which we provide a detailed description of a genus with relatively short species descriptions mentioning characters that are useful for species separation. For the new species of Distigmoptera Blake, the description is relatively long and contains characters that are helpful in determining generic placement given un- Description. Body length 1.08-1.18 mm, width 0.70-0.81 mm, elongate, relatively flat in lateral view (2.18 times as long as thick). Color brown without metallic luster, legs slightly lighter and antennae, except last antennomere, darker, almost black.
Head (Figs 4,5) flat in lateral view. Frons and vertex forming nearly straight line (Fig. 5) in lateral view. Facial part of head elongate. Supraorbital pore situated near outer corner of antennal callus, poorly visible. Antennal calli well developed, slightly longer than wide, oblique, separated from each other by wide midfrontal sulcus. Supracallinal sulcus deep, convex. Suprafrontal and supraantennal sulci well developed, deep. Supraorbital sulcus slightly impressed. Orbit as wide as transverse diameter of eye. In-terantennal space nearly as wide as transverse diameter of eye and as transverse diameter of antennal socket. Frontal ridge narrow, lowering in front of anterofrontal ridge. Anterofrontal ridge not separated from frontal ridge, long and swollen. Two ridges situated laterally of frontal ridge from lower margin of antennal socket to outer corner of mouth. Long seta situated at beginning of each ridge under antennal socket. Another long seta located on both sides of frontal ridge. Eyes small, slightly protruding laterally, 0.72 times as wide as long. Vertex covered with few large and deep punctures. Labrum with six setiferous pores, apically slightly incised. Labium with three palpomeres per palpus, distal palpomere longer than wide (Fig. 6). Maxillary palpus with four palpomeres, distal palpomere conical, slightly longer than preapical, sensilla patch with three setae (Fig.  8). Antenna with 11 antennomeres. First antennomere slightly wider and much longer than second and rest of antennomeres separately. Third and fourth antennomeres much thinner than second. Antennomeres gradually widening distally (Fig. 7). Pronotum ( Fig. 1) 1.34 times wider than long (measured in middle), without impressions, covered with large, deeply impressed punctures. Sides weakly rounded and relatively narrowly explanate, with maximum width in front of middle. Marginal anterolateral callosity situated perpendicularly to midline of body, 3.71 times shorter than lateral margin. Posterolateral callosity protruding laterally. Basal margin evenly convex, slightly extends posteriorly, with distinct border. Procoxal cavity open. Intercoxal prosternal process relatively narrow and parallel-sided in middle, with longitudinal ridge bordered by two deep grooves laterally, abruptly expanding beyond procoxae. Scutellum flat, wider than long, apex sharply triangular, sides straight. Mesocoxae separated by both meso-and metasterna. Mesosternum not covered by metasternum, horizontal ( Fig. 9). Metasternum ( Fig. 9) protruding anteriorly between mesocoxae, wide, nearly flat at apex. Elytron ( Fig. 1) widest near mid-length. Humeral callus absent. Elytral punctures arranged in nine rows not counting scutellar row. Punctures large, about as large as space between rows. Elytral apex narrowly rounded, surrounded by distinct border. Epipleura broad, slightly oblique, gradually narrowing posteriorly, not attaining sutural margin of elytron. Elytron with sensilla distributed evenly throughout surface, others concentrated in single sensilla patches (Fig. 14). Elytra fused. Elytral lock consists of longitudinal groove along its suture (Fig. 14). Wings absent.
Abdomen with five distinctly visible sternites. Apical sternite shorter than three preceding sternites combined, without appendages basally. Basal sternite longer than four following sternites together.
Median lobe of aedeagus ( Fig. 12) simple, robust, slightly and evenly curved in lateral view, without any sculpture ventrally.
Type species. Borinken elyunque Konstantinov and Konstantinova, new species. Etymology. This genus is based on the native Taino Indian name for the Island of Puerto Rico, Borinquen. The name is masculine.
Diagnosis and comparison. Borinken is very different from other flea beetle genera that are known to occur in mosses in the New World (Kiskeya in the West Indies, Nicaltica Konstantinov, Chamorro-Lacayo and Savini in Nicaragua, and Ulrica Scherer in the West Indies and Central and South America). Based on the general shape of the body, shape of the base of the pronotum without a lobe extending posteriorly, general shape of the metatibia and tarsal claw, Borinken is similar to Benedictus Scherer, which inhabits mosses in Asia and does not occur in the New World (Sprecher-Uebersax et al. 2009). It can be easily distinguished from that genus by the unique shape and details of the head, subquadrate apical antennomeres, and absence of the prebasal impression on the pronotum.
Borinken is also very different from any other West Indian or New World flea beetle genera. Among New World genera it is somewhat similar to Centralaphthona Bechyné based on the presence of antennal calli, lack of the prebasal groove on the pronotum, regular elytral striae and open procoxal cavities. Borinken can be easily differentiated from Centralaphthona by the following features: elongate facial part of head (normally short in Centralaphthona); antennal calli longer than wide (usually shorter than wide in Centralaphthona); apical antennomeres much wider than basal (about same width in Centralaphthona); vertex, pronotum, and elytra strongly punctured (punctation normally small in Centralaphthona; overall, this kind of coarse punctation is rare among flea beetles); apex of metatibia convex up to tarsomere (flat in Centralaphthona). Description. Body length 1.08-1.18 mm, width 0.70-0.81 mm, elongate, relatively flat in lateral view (2.18 times as long as thick). Color brown without metallic luster, legs slightly lighter and antennae, except last antennomere, darker, almost black. Vertex covered with large punctures, shiny, without wrinkles. Oblique fold situated between orbit and antennal callus. Proportions of antennomere lengths in male: 14:9:6:6:6:4:5:5:5:7:10. Antennomeres widened abruptly beginning from antennomere 7 (it is 0.71 times as long as wide). Pronotum evenly covered with large punctures, their diameter much larger than distance between them. Ventral side of body without many setae. Elytron with nine complete rows of punctures. Additional scutellar row incomplete. Punctures large, about as large as space between rows. Interspaces shiny with wrinkles or punctures. Proportions of tarsomere lengths of male as follows: protarsomeres 5:4:4:11; mesotarsomeres 5:4:4:11; metatarsomeres 9:4:5:11.

Borinken elyunque
Median lobe of aedeagus (Fig. 12) robust in ventral view, with apex evenly convex without apical denticle. Apex slightly swollen in lateral view. Ventral side flat, without membranous window.
Etymology. The specific epithet is a noun in apposition based on the type locality. Ecology. Unidentified moss samples that contained B. elyunque were collected in the forest from a variety of substrates (rocks, tree stumps, trunks and branches) (Figs 59, 60) Blake, 1943, by original designation).
Discussion. Distigmoptera was first recorded in the West Indies by Medvedev (2004) who described a new species from the Dominican Republic. Fourteen previously described species of this genus are known to occur in the USA, Canada, Mexico, and Costa Rica. Among the West Indian genera of Monoplatini, Distigmoptera is mostly similar to Apleuraltica Bechyné. Apart from characters mentioned in the key (see below), Distigmoptera can be differentiated from Apleuraltica by the antennae that are not clearly clubbed, antennomere six in males is only slightly different from antennomere seven (the antennae are clearly clubbed, antennomere six in males is markedly different from antennomere seven in being much shorter and narrower than seven in Apleuraltica) and by the metatibial apex without a sharp denticle (the metatibial apex has a sharp denticle in Apleuraltica). better separated in female and wider and poorly visible in male. Lighter parts of elytron covered with setae lighter in color, darker parts covered with setae darker in color. Head (Figs 16, 17) slightly convex in lateral view, evenly and strongly rugose and pubescent. Frons and vertex forming slightly convex line in lateral view. Supraorbital pore indistinguishable. Antennal callus clearly visible, nearly quadrate, its surface situated above surface of vertex. Midfrontal sulcus wide and deep. Supracallinal and supraorbital sulci poorly visible. Suprafrontal and supraantennal sulci shallow. Orbit relatively narrow, 2.50 times narrower than transverse diameter of eye. Interantennal space as wide as transverse diameter of eye. Antennal socket rounded. Frontal ridge wide, parallel sided. Anterofrontal ridge merged with frontal ridge forming denticle in middle. Eyes small, slightly protruding laterally, inner margin curved. Labrum deeply notched in middle, with six setiferous pores. Apical maxillary palpomere as wide as long, conical, much smaller than preceding palpomere. Labial palpomeres of equal length, apical conical. Antenna with 11 antennomeres, antennomeres widening apically. Antennomere four thinnest (Fig. 18). Proportions of antennomere lengths in male: 12:4:6:5:5:5:7:7:7:6:9; in female: 12:6:5:5:5:6:6:6:6:7:9.

Distigmoptera chamorrae
Pronotum (Figs 15, 18) 1.47 times wider than long. Pronotal disc anteriorly raised in two wide ridges separated by shallow and wide impressions. Anterior margin straight, with distinct border. Lateral margins subparallel, very slightly convex, without explanation. Posterior margin nearly straight, without distinct border. Anterolateral callosity globular and evenly rounded, bearing seta and not forming denticle posteriorly. Posterolateral callosity absent. Pronotal surface covered with large closely placed punctures and a few yellow, whitish and black setae. Scutellum triangular, densely covered with yellow setae. Prosternal surface densely covered with irregular punctures. Prosternal intercoxal process extended posteriorly beyond coxa and truncate posteriorly. Posterior end about twice as wide as middle. Procoxal cavities closed posteriorly. Mesosternum shorter than prosternal process, quadrate, rugose and pilose. Metasternum smooth and pilose, convex in lateral view, as long as pro-and mesosterna together. Posterior margin with sharp notch.
Elytral surface punctate (Figs 15,18), with punctures forming nine striae (not counting marginal and short scutellar striae), densely pilose with black setae near base and yellow setae in posterior half. Interspaces between puncture rows two and three, four and five, six and seven form convex ridges. Humeral callus absent. Base of elytron with callus situated between suture and humeral corner. Epipleura wide, nearly vertical, narrowing abruptly at elytral apex but not reaching it. Elytral apex narrowly truncate.
Pro-and mesofemora slightly dilated and tibiae subcylindrical, somewhat enlarged towards apical edge (Fig. 17); pubescence sparsely distributed. Metafemur greatly enlarged, 1.82 times longer than wide and 1.72 times longer than metatibia. Pro-and mesotibiae without apical spurs. Metatibia straight in lateral view, slightly curved in dorsal view. Outer and inner lateral dorsal ridges more or less straight with apical third with numerous denticles. Metatibial spur well developed. First metatarsomere inserted preapically and about as long as two subsequent tarsomeres together. Claw tarsomere swollen. Claw split in male and appendiculate in female.
Abdomen pubescent, with five visible sternites. Apical sternite shorter than three preceding sternites combined, without appendages basally (Fig. 26). Basal sternite shorter than three following sternites together. Last abdominal tergite of female without groove in middle, evenly covered with long setae.
Median lobe simple, slightly curved in lateral view with more abrupt curvature near middle; in ventral view, with lateral margins almost parallel, apex subtriangular without denticle (Fig. 22). Ventral side apically flatter than basally.
In female genitalia, posterior part of sternite eight sclerotized along its entire margin (Fig. 25). Tignum with rounded anterior margin, evenly sclerotized, bearing many moderately long setae (Fig. 25). Vaginal palpi (Fig. 24) elongate, anteriorly and along middle strongly sclerotized and merged anteriorly for more than half of their length, each with about eight setae at apex, with posterior sclerotization shorter than anterior (Fig. 24). Spermatheca curved (Fig. 23), with receptacle and pump not differentiated from each other (pump about as wide as receptacle). Apex of pump with flattened projection. Spermathecal duct long, forming "S" coils.
Diagnosis and comparison. Distigmoptera chamorrae can be easily differentiated from all continental species of Distigmoptera by the bicolorous antennae with antennomeres five in the male and five and six in the female being yellowish, much lighter than the rest of the antennae. It can be distinguished from the only other West Indian species (D. antennata Medvedev) by the absence of wings (D. antennata is winged).

Kiskeya elyunque
Etymology. The specific epithet is a noun in apposition based on the type locality. Diagnosis and comparison. Kiskeya elyunque is the only species of Kiskeya known to occur in Puerto Rico. It can be easily differentiated from the other two known species of the genus based on the key below.
Ecology. Unidentified moss samples which contained K. elyunque were collected in the forest from a variety of substrates (rocks, tree stumps, trunks and branches) (Fig.  59, 60).
Ecology. Unidentified moss samples that contained U. eltoro were collected in the forest from a variety of substrates (rocks, tree stumps, trunks and branches) (Figs 59,60 (Figs 51, 52). Head surface dorsally shiny, ventrally with some wrinkles (Fig. 53). Vertex with several large punctures, supraorbital pore as large as a few punctures on vertex near it. Supracallinal sulcus not separating antennal calli and vertex medially. Frontal ridge wide, about as long as antennal calli (Fig. 53). Anterofrontal ridge making long denticle about as long as entire clypeus. Pronotum and elytron with coarse punctures. Interspaces of elytron flat on disc, slightly convex apically. Proportions of tarsomeres of male as follows: protarsomeres 5:4:5:9; mesotarsomeres 5:4:5:9; metatarsomeres 10:5:5:9. Median lobe of aedeagus with more or less curved sides in ventral view, with ridge in middle being wider at base, narrowing towards middle and widening towards apex. In lateral view slightly curved without bump on ventral side beyond middle (Fig. 55). Spermatheca with pump at base wider than receptacle and duct making coils (Fig. 56). Sternite eight nearly fully sclerotized with tignum sharply bent anteriorly (Fig. 58). Vaginal palpi merged at about apical one third (Fig. 57).
Etymology. The specific epithet is a patronym dedicated to Mike Ivie who collected the holotype. Head with supraorbital pore much larger than a few small punctures on vertex. Supracallinal sulcus separating antennal calli and vertex medially. Pronotum and elytron with fine punctures (Fig. 47)

Key to Monoplatini genera of the West Indies
In addition to Distigmoptera and Ulrica, six other Monoplatini genera are reported in the West Indies: Aedmon Clark, Apleuraltica, Bonfilsus Scherer, Homotyphus Clark, Hypolampsis Clark, and Physimerus Clark (Takizawa 2003). Aedmon, Apleuraltica, and Bonfilsus are West Indian endemics, relatively small (with the exception of Aedmon) and relatively well circumscribed. However, Homotyphus (with about 20 species mostly from South America and just one in the West Indies), Hypolampsis (with about 50 species from North, Central, and South America and just two in the West Indies), and Physimerus (with about 60 species from Central and South America and one in the West Indies) are poorly understood; their identity, distinguishing characters and composition need extensive review. Below we provide a key for Monoplatini genera of the West Indies based on the West Indian species.