Phylogenetic assessment of the Caribbean weevil genus Lachnopus Schoenherr (Coleoptera : Curculionidae : Entiminae)

Transcription

1 CSIRO PUBLISHING Invertebrate Systematics, 2012, 26, Phylogenetic assessment of the Caribbean weevil genus Lachnopus Schoenherr (Coleoptera : Curculionidae : Entiminae) Jennifer C. Girón A,C and Nico M. Franz A,B A Department of Biology, Call Box 9000, University of Puerto Rico, Mayagüez, PR 00681, USA. B Current address: School of Life Sciences, PO Box , Arizona State University, Tempe, AZ , USA. C Corresponding author. entiminae@gmail.com Abstract. The genus Lachnopus Schoenherr (Coleoptera : Curculionidae : Entiminae : Geonemini Gistel), with 66 species described to date, is the largest and most widespread entimine weevil genus in the Caribbean region. The monophyly and internal phylogenetic composition of Lachnopus are tested, using a cladistic analysis of 61 adult morphological characters of 31 ingroup species and five outgroup taxa representing the genera Diaprepes Schoenherr, Exophthalmus Schoenherr, Pachnaeus Schoenherr (all Eustylini), Ischionoplus Chevrolat (Geonemini) and Apodrosus Marshall (Polydrusini). The analysis produced two most parsimonious cladograms with a length of 167 steps, a consistency index of 36 and a retention index of 73. According to the strict consensus and preferred character state optimisations, the examined species of Lachnopus do not form a monophyletic entity, even though most species are placed in a major L. coffeae L. guerinii ingroup clade, which stands in sister relationship to Ischionoplus. This major clade is constituted by at least three well-recognisable subgroups, each associated with a particular geographic range in the Caribbean region: (1) a widely distributed L. coffeae L. lineicollis grade ; (2) the Hispaniolan L. proteus L. mercator clade; (3) and the L. hispidus L. guerinii clade with species from Cuba, Florida and Jamaica. This study provides a sound phylogenetic basis for future revisions of Lachnopus and related geonemine genera. Received 9 September 2011, accepted 16 January 2012, published online 19 June 2012 Introduction Lachnopus Schoenherr (1840: 380) constitutes the most speciose and one of the most widespread genera of Caribbean entimine weevils (Coleoptera : Curculionidae : Entiminae sensu Alonso- Zarazaga and Lyal 1999). At present, 66 species are recognised within the genus (O Brien and Wibmer 1982; de Zayas 1988). Many of these show narrow distributional ranges and high rates of inter- or intra-island endemism; for instance, 26 out of 27 recorded Hispaniolan species apparently only occur on this Greater Antillean island. Overall, Lachnopus ranges from the south-eastern United States to Saint Vincent in the Lesser Antilles; however, the highest species diversity is concentrated in the Greater Antilles (O Brien and Wibmer 1982; de Zayas 1988). Although the natural history of most Lachnopus species is not well known, several species are considered economically relevant as herbivores of cultivated plants; for example, L. inconditus Rosenschoeld on citrus trees in the Dominican Republic (Woodruff 1985) and L. coffeae Marshall on coffee plants in Puerto Rico (Wolcott 1948). Lachnopus is classified in the predominantly New World tribe Geonemini Gistel, as circumscribed in Alonso-Zarazaga and Lyal (1999) and Franz (2012). Species placed in the genus vary in size from 5 to 25 mm, with body shapes ranging from elongate to oval, and display a wide spectrum of colour and scale patterns; including combinations of small, pyriform, dark and mostly Journal compilation CSIRO 2012 glabrous species to large, elongate species that are uniformly and densely covered with brilliant iridescent scales, or whose scale coverage is arranged in specific patterns. Van Emden s (1944) key constitutes the most recent reference for identifying the genus, which, given its geographic range, was not treated in Champion s(1911) Biologia Centrali-Americana. Accordingly, Lachnopus is diagnosed by the following suite of traits: rostrum weakly and evenly convex throughout; antennal scape extending to, or slightly passing beyond middle of eye; frons between eyes conspicuously narrower than dorsal surface of rostrum; head not constricted posteriad of eyes; eyes only moderately convex; humeri only slightly wider than posterior margin of pronotum; femora unarmed; tibiae ventrally denticulate; and metatibial corbel (i.e. bevel sensu Thompson 1992) lacking scales. However, several of the species currently assigned to the genus are not subsumed under van Emden s (1944) concept of Lachnopus. As is typical for other Caribbean entimine genera (Woodruff 1985; Franz and Girón 2009; Franz 2010a, 2010b, 2012; Girón and Franz 2010), Lachnopus has had a complex nomenclatural and taxonomic history characterised by numerous isolated additions of species and a lack of comprehensive taxonomic revisions (see also O Brien and Wibmer 1982). The valid genus name was originally introduced by Schoenherr in 1840; it is a replacement name for Menoetius Dejean (1821: 94; type species

2 68 Invertebrate Systematics J. C. Girón and N. M. Franz Curculio valgus Fabricius 1775: 150) and Ptilopus Schoenherr (1823: column 1140, and Schoenherr 1826: 118; type species Curculio aurifer Drury 1773: 68) (see Alonso-Zarazaga and Lyal 1999). Thus, the first descriptions of species now pertaining to Lachnopus were published under the genus name Curculio Linnaeus (1758: 377), during the period of (O Brien and Wibmer 1982). Several subsequent species names and circumscriptions were published originally as members of Menoetius and Ptilopus; however, since the resolution of the International Commission of Zoological Nomenclature (ICZN 1987) these two names have been placed into synonymy with the valid genus name Lachnopus (see also Alonso-Zarazaga and Lyal 1999). In addition to the aforementioned nomenclatural changes, there has been considerable uncertainty with regards to the taxonomic distinctness of Lachnopus vis-à-vis other genera in the Geonemini as well as other taxa placed in the closely related tribe Eustylini Lacordaire (Lacordaire 1863; Kuschel 1955; Vaurie 1961; Woodruff 1985; O Brien and Kovarik 2001; Franz 2010a, 2011, 2012). In particular, Lachnopus has been poorly delimited from members of the eustyline genus Exophthalmus Schoenherr, which presently contains more than 40 Caribbean species (Morrone 1999; Franz 2012). Other Caribbean genera with presumed affinities to Lachnopus include Apotomoderes Dejean (see Franz 2010b), Ischionoplus Chevrolat, Pachnaeus Schoenherr, Tetrabothynus Labram & Imhoff and other as yet undescribed taxa (J. C. Girón and N. M. Franz, pers. obs.). Woodruff (1985: 371) aptly described this situation as a Pandora s Box of taxonomic confusion. The most recent species addition to Lachnopus was L. kofresi Wolcott (1941: 104). Hustache (1929) reviewed the taxonomy of the Guadeloupian species therein treated under the genus name Prepodes Schoenherr, a synonym of Exophthalmus whereas Wolcott (1948) provided a synopsis of the species of Puerto Rico. Marshall (1922, 1926a, 1926b, 1933, 1934) described 10 species from the Greater Antilles in a series of isolated treatments and de Zayas (1988) described nine new species from Cuba. Nevertheless, and in spite of the striking appearance, diversity and abundance of Lachnopus specimens in the field and in research collections, no author has attempted a comprehensive revision of the genus since Gyllenhal described nearly 20 species some 170 years ago (in Schoenherr 1834, 1840). As a result, much of the material available in research collections is simply identified as Lachnopus spp., and no reliable inferences about the taxonomic and biogeographic relationships within this diverse lineage are possible. Here we provide a first phylogenetic assessment of Lachnopus based on a morphological cladistic analysis of adult weevils pertaining to 31 putative ingroup species and six suitable outgroup taxa. In particular, we evaluate whether Lachnopus as presently composed (O Brien and Wibmer 1982; Morrone 1999; Franz 2012) is a monophyletic entity, and which main lineages and corresponding synapomorphies can be distinguished within this Caribbean group of weevils. Our study therefore presents a critical step towards a monographic treatment of all species of Lachnopus and a deeper understanding of its biogeographic patterns. Materials and methods Morphological characters The morphological terminology is generally in accordance with Torre-Bueno (Nichols 1989; see also Girón and Franz 2010), with specific terms used for characterising the apex of the rostrum (Vaurie 1963), ventral structures of the head (Lyal 1995), tibial apices and abdominal segments (Thompson 1992), and female and male terminalia (Howden 1995; Velázquez de Castro 1997; Wanat 2007). All observations and dissections were performed using Leica MS5 and MZ16 stereomicroscopes (magnification: 7 115) (Leica Microsystems, Wetzlar, Germany) or an Olympus BX41 compound microscope (magnification: ) (Olympus Corporation, Tokyo, Japan), each equipped with an ocular graticule for measurements of lengths and relations. Habitus photographs were taken with a Microptics XLT imaging system (Visionary Digital, Palmyra, VA, USA), whereas scanning electron micrographs were produced with a JEOL 5410LV system (Japan Electron Optics Laboratory Co., Tokyo, Japan). Line drawings of internal structures were initially traced from digital images taken through the compound microscope, and subsequently redrawn using an illustration software program. The figures are arranged first in accordance with the character sequence and second according to phylogeny. Examined taxa In light of practical limitations with regards to the availability of specimens for detailed dissections, an exemplar approach was used (Prendini 2001). In all, 31 species of Lachnopus occurring throughout the entire Caribbean region were studied (Table 1). The respective specimens were obtained on loan from the Charles W. O Brien Collection (CWOB, Green Valley, AZ, USA) and Invertebrate Collection, University of Puerto Rico (UPRM, Mayagüez, PR, USA) collections. Given the uncertain monophyly and relationships of Lachnopus to other geonemine and eustyline taxa, a representative set of five entimine outgroup taxa was chosen (Nixon and Carpenter 1993) including: Ischionoplus viridiguttatus Chevrolat, 1878: X (Geonemini); Diaprepes abbreviatus (Linnaeus) 1758: 386; Exophthalmus quadrivittatus (Olivier) 1807: 325, Pachnaeus litus (Germar) 1824 : 431 (all Eustylini); and Apodrosus argentatus Wolcott, 1924: 130 (Polydrusini Schoenherr). In accordance with emerging phylogenetic results on Neotropical eustyline and geonemine genera (Franz 2012), the cladogram was rooted with A. argentatus. A synoptic set of the examined taxa has been placed in the University of Puerto Rico at Mayagüez Invertebrate Collection (UPRM-INVCOL; Franz and Yusseff Vanegas 2009). Phylogenetic analysis The character matrix was compiled and edited using ASADO (Nixon 2008). The most parsimonious cladograms and character state optimisations were identified via a comprehensive search using the parsimony ratchet (Nixon 1999) as implemented in TNT (Goloboff et al. 2008; spawned out of ASADO), based on the following commands: (1) ratchet settings 200 iterations per replication, 4% up-/down-weighted; (2) drift settings 100

3 Phylogeny of the Caribbean weevil genus Lachnopus Invertebrate Systematics 69 Table 1. Names and distributional information for the analysed species of Lachnopus and suitable outgroup, mainly according to O Brien and Wibmer (1982) Bold font indicates the locality of provenance of the examined material; FL, Florida; CR, Costa Rica; GT, Guatemala; MX, Mexico; NI, Nicaragua; PA, Panama; CU, Cuba; DR, Dominican Republic; HA, Haiti; HI, Hispaniola: Dominican Republic and Haiti; JM, Jamaica; PR, Puerto Rico, including Mona and Vieques Islands; AN, Anguilla; BA, Barbados; DO, Dominica; GU, Guadeloupe; MA, Martinique; MO, Monserrat; NE, Nevis; SB, St Barthélemy; SC, St Croix; SK, St Kitts; SL, St Lucia; ST, St, Thomas; SV St Vincent; TO, Tortola Taxon Pachnaeus litus (Germar), (Cyphus) Apodrosus argentatus Wolcott, Diaprepes abbreviatus (Linnaeus), (Curculio) Exophthalmus quadrivittatus (Olivier), (Curculio) Exophthalmus sulcicrus Champion, Ischionoplus viridiguttatus Chevrolat, 1878 X Lachnopus acuticollis (Gyllenhal), (Ptilopus) Lachnopus albomaculatus (Gyllenhal), (Ptilopus) Lachnopus argus (Reiche), (Ptilopus) Lachnopus atramentarius (Gyllenhal), (Ptilopus) Lachnopus aurifer (Drury), (Curculio) Lachnopus campechianus Gyllenhal, Lachnopus chlorophanus (Gyllenhal), (Ptilopus) Lachnopus coffeae Marshall, Lachnopus curvipes (Fabricius), (Curculio) Lachnopus floridanus Horn, Lachnopus gowdeyi Marshall, Lachnopus guerinii Jacqelin du Val, Lachnopus hispidus (Gyllenhal), (Ptilopus) Lachnopus histrio (Marshall), (Ptilopus) Lachnopus inconditus Rosenschoeld, Lachnopus kofresi Wolcott, Lachnopus lineatoguttatus Perroud, Lachnopus lineicollis (Chevrolat), (Diaprepes) Lachnopus luctuosus (Klug), (Sitona) Lachnopus mercator (Olivier), (Curculio) Lachnopus mundus (Gyllenhal), (Ptilopus) Lachnopus niveoirroratus Jacqelin du Val, Lachnopus planifrons Gyllenhal, Lachnopus proteus (Olivier), (Curculio) Lachnopus seini Wolcott, Lachnopus sparsimguttatus Perroud, Lachnopus splendidus Boheman, Lachnopus spretus (Gyllenhal), (Ptilopus) Lachnopus valgus (Fabricius), (Curculio) Lachnopus vittatus (Klug), (Ptilopus) Lachnopus yaucona Wolcott, Distribution FL, MX, CU, JM DR, PR, SC BA, DO, GU, HI, MA, MO, PR, SL, SV HI CR, GT, NI, PA, SA DR CU HI (HA) FL, CU HI (DR) CU, JM GU HA PR DR, GU, JM, NE, PR, SB, SK, ST, SV, TO FL JM CU FL, CU HA HI (DR) PR (Mona Is.) CU GU CU HI DR CU DR HA PR CU CU DR AN, PR, SB, SC CU PR iterations per replication; (3) tree fusion settings 10 rounds, 200 MB max. RAM; (4) general settings 1000 tree to hold; (5) analyses ratchet, drift, sectorial search, tree fusion, TBR-max; and (6) xmult settings three hits, five consense. The resulting cladograms and character state transformations were examined in ASADO under various optimisations. Bremer branch support values (Bremer 1994) were calculated in NONA (Goloboff 1999) with the commands hold , suboptimal 15 and bsupport 15. Results Most parsimonious cladograms A comprehensive search of the cladistic matrix containing 37 terminal taxa (5 outgroup; 31 ingroup) and 61 parsimonyinformative characters (Table 2) yielded two most parsimonious cladograms, each with a length (L) of 167 steps, a consistency index (CI) of 36 and a retention index (RI) of 73, respectively (Farris 1989). The strict consensus tree (L = 168; CI = 36; RI = 73) collapsed two nodes (Fig. 1). The first of these affected the phylogenetic arrangement of P. litus (Germar) and L. histrio (Marshall), grouped either sequentially or as a separate clade and in sister-group relationship to the I. viridiguttatus L. guerinii ingroup clade; whereas the second affected the placement of L. sparsimguttatus Perroud, either in sister-group relationship to the L. floridanus L. luctuosus clade or forming a trichotomy with the former two taxa. The L. coffeae L. yaucona clade also formed a trichotomy due to lack of node support. Bremer support values were mapped onto the internal nodes of

4 70 Invertebrate Systematics J. C. Girón and N. M. Franz Taxa Table 2. Character matrix for the cladistic analysis of the species of Lachnopus, with select outgroup taxa, arranged in alphabetic sequence, inapplicable character states;?, missing information (see also text) Characters Apodrosus argentatus Diaprepes abbreviatus Exophthalmus quadrivittatus Pachnaeus litus Ischionoplus viridiguttatus Lachnopus acuticollis Lachnopus albomaculatus ????????? Lachnopus argus Lachnopus atramentarius Lachnopus aurifer Lachnopus campechianus Lachnopus chlorophanus ? ????00?????????????? Lachnopus coffeae Lachnopus curvipes Lachnopus floridanus Lachnopus gowdeyi Lachnopus guerinii Lachnopus hispidus Lachnopus histrio ? ????00?????????????? Lachnopus inconditus Lachnopus lineatoguttatus Lachnopus lineicollis Lachnopus kofresi Lachnopus luctuosus ? ????01????????????? Lachnopus mercator ????????? Lachnopus mundus ? ????00?????????????? Lachnopus niveoirroratus ????????? Lachnopus planifrons Lachnopus proteus Lachnopus seini Lachnopus sparsimguttatus ????????? Lachnopus splendidus ????????? Lachnopus spretus Lachnopus valgus Lachnopus vittatus Lachnopus yaucona the strict consensus (Fig. 1), which also served as the reference tree for addressing phylogenetic groups and relationships. In addition, one of the two most parsimonious trees (Fig. 2) is presented to illustrate the preferred character state optimisations: 50 unambiguous, 3 ACCTRAN and 8 DELTRAN (Agnarsson and Miller 2008). Character analysis The characters, states and inferred optimisations are presented simultaneously in this section. The implications of alternative optimisations are assessed in the corresponding character accounts. Individual lengths, consistency and retention indices (l, ci and ri) are provided for all characters that are inferred to be homoplasious in the present analysis. The subsequent discussion of synapomorphies and relevant homoplasious features is restricted to the ingroup taxa (see Franz 2012 for a more inclusive analysis on eustyline and geonemine taxa). The characters correspond to observations made on male specimens unless stated otherwise. Characters 20, and were coded as missing (? ) for L. chlorophanus (Gyllenhal), L. histrio, L. luctuosus (Klug) and L. mundus (Gyllenhal), due to a lack of male specimens available for study. Similarly, characters were coded as missing (? ) for L. albomaculatus (Gyllenhal), L. mercator (Olivier), L. niveoirroratus Jacquelin du Val, L. sparsimguttatus and L. splendidus Boheman due to a lack of female specimens. 1. Scales, surface microsculpture: (0) sculptured (usually ribbed) (Fig. 3A); (1) smooth (Fig. 3B). Synapomorphy for the I. viridiguttatus L. guerinii clade. 2. Head, arrangement of smooth scales on dorsal surface: (0) variously sparse and/or scarce (to patchy), appressed (Fig. 4A); (1) mostly uniform and dense, with curved (elevated) scales (Fig. 4B). Coded as inapplicable for taxa with ribbed scales (see character 1). Convergently

5 Phylogeny of the Caribbean weevil genus Lachnopus Invertebrate Systematics 71 Fig. 1. Strict consensus tree for Lachnopus species and select outgroup taxa (L = 168; CI = 36; RI = 73). Numbers displayed above at the left end of each branch represent Bremer support values.

6 72 Invertebrate Systematics J. C. Girón and N. M. Franz Fig. 2. Preferred most parsimonious cladogram and character state optimisations (50 unambiguous, three ACCTRAN and eight DELTRAN; see Agnarsson and Miller 2008). Black rectangles represent single, non-homoplasious character state transformations; whereas white rectangles represent multiple, homoplasious character state transformations. The small numbers above and below each rectangle correspond to character numbers and states, respectively. present in the L. proteus L. mercator and L. hispidus L. niveoirroratus clades (l = 2; ci = 50; ri = 88). 3. Head, scales on dorsal surface restricted to inner margin of eyes, thereafter continuing anteriorly as longitudinal lateral stripes along rostrum, yet mesal area of rostrum glabrous: (0) absent; (1) present (Fig. 4C). Synapomorphy for the L. lineatoguttatus L. guerinii clade. 4. Head, surface of inner margin of eyes: (0) flat (Fig. 4D); (1) depressed (Fig. 4E). Convergently present in D. abbreviatus, the L. coffeae L. yaucona clade, the L. floridanus L. sparsimguttatus clade and in L. kofresi (l = 4; ci = 25; ri = 62). 5. Head, eyes in frontal view large and elongate in species where luctuosus-type of scale coverage is present (see character 25): (0) absent; (1) present (Fig. 4A). Coded as inapplicable in species with other types of scale coverage. Convergently present in the L. coffeae L. yaucona clade, in L. curvipes (Fabricius) and in the L. campechianus L. lineicollis clade (l = 3; ci = 33; ri = 60). 6. Head, presence of ocular sclerite: (0) well defined; (1) not defined (Fig. 4D). Synapomorphy for the L. curvipes L. guerinii clade, with a subsequent reversal in L. valgus (l = 2; ci = 50; ri = 90).

7 Phylogeny of the Caribbean weevil genus Lachnopus Invertebrate Systematics 73 A B Fig. 3. Microsculpture of scale surface (SEM). (A) Ribbed scales of Apodrosus argentatus, 1000;(B) smooth scales of Lachnopus coffeae, 350. A B C D E Fig. 4. Head in frontal view. (A) Lachnopus campechianus; (B) L. hispidus; (C) L. lineatoguttatus; (D) L. aurifer; (E) L. luctuosus. Scale bar = 1 mm. 7. Rostrum, occipital suture (as circumscribed in Lyal 1995): (0) not visible to shallow (Fig. 5A); (1) visible, deep (Fig. 5B). Synapomorphy for the L. albomaculatus L. guerinii clade, with a subsequent reversal in the L. proteus L. guerinii clade, and an apparent regain in the L. argus L. guerinii clade (l = 3; ci = 33; ri = 86). 8. Head, lateral margins of rostrum: (0) parallel to apically convergent (Fig. 4B, D); (1) widened at or apicad of antennal insertion (Fig. 4A, C, E). Convergently present in the L. albomaculatus E. quadrivittatus and L. coffeae L. guerinii clades, with three subsequent reversals in L. valgus and in the L. planifrons L. mercator and L. hispidus L. gowdeyi clades, respectively (l = 5; ci = 20; ri = 73). 9. Head, dorsal outline of rostrum in lateral view: (0) rounded; (1) straight. Convergently present in the L. albomaculatus E. quadrivittatus clade, in L. histrio and in the L. curvipes L. guerinii clade, with a subsequent reversal in L. valgus (l = 4; ci = 25; ri = 50). 10. Rostrum, direction of scrobe: (0) passing below eye (Fig. 5A); (1) directed towards (or passing over ) eye (Fig. 5B). DELTRAN optimisation preferred (see Fig. 2), taking into account the differences in head shape between P. litus and L. histrio. Thus, convergently present in P. litus, L. histrio and in the L. kofresi L. guerinii clade (l = 3; ci = 33; ri = 84). 11. Rostrum, posterior region of dorsal margin of scrobe: (0) carinate (Fig. 5B); (1) weakly defined (Fig. 5A). Convergently present in the L. albomaculatus E. quadrivittatus clade (with a subsequent reversal in D. abbreviatus), in L. valgus, the L. kofresi L. lineicollis clade and in the L. atramentarius L. mercator clade, with a subsequent reversal in the L. chlorophanus L. mercator clade (l = 6; ci = 16; ri = 50). 12. Rostrum, carinae formed by the posterior region of dorsal margin of scrobe: (0) sharp (Fig. 5B); (1) rounded. Coded as inapplicable for species in which the posterior region of dorsal margin of the scrobe does not form a well-defined carina (see character 11). DELTRAN optimisation preferred (see Fig. 2), assuming a late reversal to a sharp carina in the L. chlorophanus L. mercator clade. Synapomorphy for the L. albomaculatus L. guerinii clade, with three subsequent reversals in the L. chlorophanus L. mercator clade, the L. hispidus L. gowdeyi clade and in the L. argus L. guerinii clade (l = 4; ci = 25; ri = 62). 13. Antennae, apex of antennal scape: (0) slightly and/or gradually widened (Fig. 4C); (1) strongly and somewhat abruptly widened (Fig. 4E). Convergently present in L. albomaculatus, I. viridiguttatus, and in the L. floridanus L. sparsimguttatus and L. splendidus L. guerinii clades (l = 4; ci = 25; ri = 57).

8 74 Invertebrate Systematics J. C. Girón and N. M. Franz A B Fig. 5. Head in lateral view (SEM). (A) Lachnopus inconditus, 35; (B) L. curvipes, Mouthparts, left mandibular scar (see Thompson 1992): (0) equate with surface to slightly raised (Fig. 4C); (1) strongly protruded and sharply raised (Fig. 4D). Synapomorphy for the L. aurifer L. gowdeyi clade. 15. Mouthparts, number of setae on each side of labial prementum: (0) 0 5; (1) 6 or more. Convergently present in L. albomaculatus, E. quadrivittatus, P. litus and in the L. argus L. guerinii clade (l = 4; ci = 25; ri = 25). 16. Pronotum, mesal longitudinal impression: (0) absent or not visible; (1) present (Fig. 6A). Convergently present in the D. abbreviates E. quadrivittatus clade, P. litus, L. gowdeyi Marshall and in the L. lineatoguttatus L. guerinii clade (l = 4; ci = 25; ri = 62). 17. Pronotum, surface of posterior margin: (0) simple (Fig. 6B); (1) carinate (Fig. 6A). Synapomorphy for the L. hispidus L. guerinii clade. 18. Pronotum, angle formed at posterior corners: (0) acute (Fig. 6C); (1) right to obtuse (Fig. 6B). Synapomorphy for the P. litus L. guerinii clade, with a subsequent reversal in the L. hispidus L. guerinii clade and an apparent regain in the L. argus L. guerinii clade (l = 3; ci = 33; ri = 81). 19. Elytra, development of humeri: (0) present, well developed (Fig. 6D), where the width of the elytra, measured at the humeri, is clearly larger than the posterior width of the pronotum; (1) reduced to absent (Fig. 6C), where the width of the elytra, measured at the humeri, is either only slightly larger or as wide as the posterior width of the pronotum. Convergently present in L. valgus, L. kofresi and in the L. hispidus L. guerinii clade, with a subsequent reversal (= well developed) in the L. argus L. guerinii clade (l = 4; ci = 25; ri = 62). 20. Elytra, lateral margins on male: (0) oriented in parallel from base to declivity, either straight or rounded (Fig. 6D); (1) clearly and continuously converging from base (minimally) to declivity (Fig. 6A, B, C). Synapomorphy for the L. proteus L. guerinii clade. 21. Elytra, scale coverage uniform and dense, composed mainly of green iridescent scales (at least in lateral regions: (0) absent; (1) present (Fig. 6B). Synapomorphy for the L. proteus L. mercator clade, with a subsequent reversal in the L. atramentarius L. inconditus clade (l = 2; ci = 50; ri = 80). 22. Elytra, presence of splendidus-type of scale coverage: dorsal surface with alternating large glabrous areas of orange to reddish brown integument and dark to light turquoise scales forming patches or stripes along odd strial intervals (Fig. 6A): (0) absent; (1) present. Synapomorphy for the L. lineatoguttatus L. guerinii clade. 23. Elytra, distribution of splendidus-type of scale coverage (see character 22): (0) mainly along elytral intervals 1, 3, 5 and 7, continued as longitudinal stripes along pronotum; (1) along elytral intervals 3, 5 and 7, with an anterior and a posterior pair of mesal patches of scales on pronotum (Fig. 6A). Coded as inapplicable for species that lack the splendidus-type of scale coverage (see character 22). Synapomorphy for the L. argus L. guerinii clade. Lachnopus splendidus is coded as 0 as this species lacks scales along elytral interval 1, yet presents longitudinal stripes along the pronotum. 24. Elytra, presence of aurifer-type of scale coverage on dorsal surface: composed of oval overlapping scales and limited to a transverse stripe covering anterior region of intervals 2 5 accompanied by longitudinal stripes along intervals 3 and 5: (0) absent; (1) present (Fig. 6C). Synapomorphy for the L. aurifer L. gowdeyi clade. 25. Elytra, presence of luctuosus-type of scale coverage: dorsal surface with yellow to white or bluish, sesame seed-like scales arranged at varying densities, not forming distinct patterns of distribution beyond patches of ~20 scales: (0) absent; (1) present (Fig. 6D). Convergently present in L. acuticollis (Gyllenhal) and in the I. viridiguttatus L. guerinii clade, with a subsequent reversal in the L. proteus L. guerinii clade (l = 3; ci = 33; ri = 83). 26. Elytra, presence of regularly arranged straight and erect setae along intervals: (0) absent; (1) present (Fig. 7A). Convergently present in the L. planifrons L. mercator and L. hispidus L. niveoirroratus clades. In remaining taxa, short, curved and decumbent seta-like scales arranged regularly along the strial intervals may be present, yet such scales are not considered homologous to state (1) since they are not setae (l = 2; ci = 50; ri = 87).

9 Phylogeny of the Caribbean weevil genus Lachnopus Invertebrate Systematics 75 A B C D Fig. 6. Dorsal habitus. (A) Lachnopus guerinii, male; (B) L. mercator, male; (C) L. aurifer, male; (D) L. floridanus, male. Scale bar = 5 mm. A B Fig. 7. Lateral habitus. (A) Lachnopus chlorophanus, female; (B) L. gowdeyi, male. Scale bar = 5 mm. 27. Elytra, interval IX: (0) widened anteriad of metacoxae (Fig. 7B); (1) not particularly widened at any point (Fig. 7A). ACCTRAN optimisation preferred (see Fig. 2), based on the distinctness of the L. hispidus L. niveoirroratus clade among its relatives. Thus, convergently present in the L. acuticollis E. quadrivittatus and L. coffeae L. guerinii clades, with a subsequent reversal in the L. hispidus L. guerinii clade and an apparent regain in the L. hispidus L. niveoirroratus clade (l = 4; ci = 25; ri = 72). 28. Legs, presence of smooth scale coverage on metafemora: (0) scarce to irregular (mostly glabrous, with interspersed areas of dense and uniform coverage; see Fig. 7B); (1) dense and uniform. Coded as inapplicable in taxa with sculptured scales (see character 1). Convergently present in L. planifrons Gyllenhal, in the L. spretus L. mercator clade and in L. hispidus (Gyllenhal) (l = 3; ci = 33; ri = 50). 29. Legs, surface of male femora: (0) smooth; (1) with tiny conspicuous teeth (Fig. 6C). Synapomorphy for L. aurifer L. gowdeyi clade. 30. Legs, profile of male metatibiae: (0) straight (Fig. 8A); (1) curved (Fig. 8B). Synapomorphy for the L. curvipes L. valgus clade. 31. Legs, denticles on male ventral surface of metatibiae: (0) absent (Fig. 8A); (1) present (Fig. 8B). Convergently present in the L. albomaculatus E. quadrivittatus and L. coffeae L. guerinii clades (l = 2; ci = 50; ri = 50). 32. Legs, presence of a large simple tooth in proximal region of inner margin of male metatibiae: (0) absent; (1) present (Fig. 8B). Synapomorphy for the L. curvipes L. valgus clade. Males of L. kofresi present a relatively enlarged denticle in comparison with the remaining denticles along the metatibia; however, this denticle is positioned near mid length, as opposed to proximally, and does not develop to form the shape of the characteristic L. curvipes L. valgus tooth. On the other hand, L. campechianus Gyllenhal presents a proximal tooth with three cusps. Neither are considered homologous to the apomorphic condition of the L. curvipes L. valgus clade. 33. Legs, bevel of apex of metatibiae (see Thompson 1992): (0) glabrous (Fig. 8C); (1) covered with spines (strong acute setae) or scales. Coded as inapplicable in A. argentatus in which the apex of posterior tibia lacks a bevel (see Girón and Franz 2010). Convergently present in the L. acuticollis E. quadrivittatus and L. argus L. guerinii clades (l = 2; ci = 50; ri = 75).

10 76 Invertebrate Systematics J. C. Girón and N. M. Franz A B C Fig. 8. Ventral side of male left metatibia. (A) Ischionoplus viridiguttatus; (B) Lachnopus curvipes; (C) apex of metatibia of L. curvipes, Legs, premucro developed into a spine on apex of metatibiae (see Thompson 1992): (0) absent; (1) present (Fig. 8C). Coded as inapplicable in A. argentatus, in which the apex of the metatibia has a spur instead of a spine (see Snodgrass 1993). Synapomorphy for the I. viridiguttatus L. guerinii clade, with a subsequent reversal in the L. mundus L. mercator clade (l = 2; ci = 50; ri = 88). 35. Male terminalia, shape of apex of apodeme of spiculum gastrale (see Wanat 2007): (0) strongly expanded; (1) simple to slightly expanded. Convergently present in the L. albomaculatus E. quadrivittatus and L. curvipes L. guerinii clades, with a subsequent reversal in the L. hispidus L. guerinii clade (l = 3; ci = 33; ri = 85). 36. Male terminalia, outline of apical portion of aedeagus (see Wanat 2007): (0) continuous; (1) sinuate. Convergently present in L. albomaculatus and in the L. coffeae L. guerinii clade, with a subsequent reversal in L. floridanus Horn (l = 3; ci = 33; ri = 66). 37. Male terminalia, shape of apex of aedeagus (see Wanat 2007): (0) rounded (Fig. 9A); (1) truncate (Fig. 9E). Convergently present in the L. curvipes L. valgus clade, the L. campechianus L. lineicollis clade, the L. atramentarius L. mercator clade and in L. splendidus (l = 4; ci = 25; ri = 62). 38. Male terminalia, distance from apical margin of ostium to apex of aedeagus (see Wanat 2007): (0) width of aedeagus; (1) more than 1.5 width of aedeagus (Fig. 9G, H). Synapomorphy for the L. lineatoguttatus L. vittatus clade. 39. Male terminalia, width of single apical sclerite of aedeagus: (0) narrow, occupying one-third or less of width of aedeagus; (1) wide, occupying more than half of width of aedeagus (Fig. 9C). Coded as inapplicable for taxa that have more than one apical aedeagal sclerite. DELTRAN optimisation preferred (see Fig. 2), thus not presuming a homologous relationship with the particularly configured apical sclerites of the L. coffeae L. yaucona clade, which are composed of two lateral plates and one mesal plate, the latter apically truncate and slightly wider. Convergently present in D. abbreviatus and in the L. curvipes L. guerinii clade (l = 2; ci = 50; ri = 50). 40. Male terminalia, presence of fine small papillae in the endophallus (see Wanat 2007): (0) present (Fig. 9A D, G, H); (1) absent (Fig. 9E, F). Synapomorphy for the L. campechianus L. lineicollis clade. 41. Male terminalia, presence of a single endophallic sclerite (see Wanat 2007): (0) absent; (1) present (Fig. 9C F). Convergently present in L. acuticollis and in the L. campechianus L. lineicollis and L. argus L. guerinii clades (l = 3; ci = 33; ri = 50). 42. Male terminalia, presence of a single pair of matching sclerites in the endophallus (see Wanat 2007): (0) absent; (1) present (Fig. 9A, B). ACCTRAN optimisation preferred (Fig. 2), thus supporting a L. chlorophanus L. mercator sister relationship. Synapomorphy for the I. viridiguttatus L. guerinii clade, with subsequent reversals in L. curvipes, the L. campechianus L. lineicollis clade, L. proteus (Olivier), and the L. chlorophanus L. mercator and L. lineatoguttatus L. guerinii clades. The paired sclerites can vary in shape, ranging from rod-like structures to irregular plates (l = 6; ci = 16; ri = 66). 43. Male terminalia, degree of sclerotisation of dorsal surface of aedeagus: (0) membranous; (1) slightly lighter than or similar to sclerotisation of ventral surface. Synapomorphy for the L. albomaculatus L. guerinii clade, with subsequent reversals in the D. abbreviatus E. quadrivittatus clade, L. sparsimguttatus and in the L. atramentarius L. inconditus clade (l = 4; ci = 25; ri = 40). 44. Male terminalia, mesal longitudinal carina on dorsal surface of aedeagus: (0) absent; (1) present. Coded as inapplicable for taxa with a membranous dorsal surface of the aedeagus (see character 43). ACCTRAN optimisation preferred (Fig. 2), thus supporting a close L. chlorophanus L. mercator relationship. Convergently

11 Phylogeny of the Caribbean weevil genus Lachnopus Invertebrate Systematics 77 A B C D E F G H Fig. 9. Male terminalia, aedeagus. Lachnopus hispidus:(a) dorsal view, (B) lateral view; L. argus:(c) dorsal view, (D) lateral view; L. campechianus:(e) Dorsal view, (F) lateral view; L. vittatus: (G) Dorsal view, (H) lateral view. Basal membranes of the aedeagus not drawn. Scale bar = 1 mm. present in L. albomaculatus and in the I. viridiguttatus L. guerinii clade, with three subsequent reversals in the L. kofresi L. lineicollis clade, the L. planifrons L. mercator clade and in L. splendidus, and an apparent regain in the L. chlorophanus L. mercator clade (l = 6; ci = 16; ri = 37). 45. Male terminalia, shape of basal margin of ventral surface of aedeagus: (0) widely roundly emarginate (Fig. 9G); (1) narrowly mesally emarginate (Fig. 9A). Synapomorphy for the L. albomaculatus L. guerinii clade, with two subsequent reversals in E. quadrivittatus and in the L. coffeae L. guerinii clade, and with three apparent regains in L. proteus and in the L. hispidus L. gowdeyi and L. argus L. guerinii clades (l = 6; ci = 16; ri = 50). 46. Male terminalia, degree of sclerotisation (colouration) of base of ventral surface of aedeagus: (0) regularly sclerotised, colouration similar to remainder of aedeagus; (1) heavily sclerotised, darker than remainder of aedeagus. DELTRAN optimisation preferred (Fig. 2), thus assuming independent acquisitions in L. planifrons and in the L. atramentarius L. inconditus clade (l = 2; ci = 50; ri = 50). 47. Male terminalia, mesal longitudinal carina on ventral surface of aedeagus: (0) absent; (1) present. Convergently present in L. lineatoguttatus Perroud and in the L. argus L. guerinii clade (l = 2; ci = 50; ri = 50). 48. Male terminalia, lateral profile of ventral surface of aedeagus: (0) rounded (Fig. 9B, D, H) to slightly sigmoidal (Fig. 9F); (1) irregular (see Marshall 1922). Convergently present in L. albomaculatus and in the L. coffeae L. yaucona clade (l = 2; ci = 50; ri = 66). 49. Male terminalia, aedeagus elongate, longer than wide, with a complex of paired mesal sclerites, elongate dorsal apical plates and shorter anteapical ventral sclerites: (0) absent; (1) present (Fig. 9A, B). Synapomorphy for the L. hispidus L. gowdeyi clade. 50. Male terminalia, aedeagus nearly 6.5 longer than wide, with a complex of a short, basally positioned and basally bifurcated sclerite, paired mesal or anteapical ventral sclerites, and a mesal, dorsal apical plate that is basally bifurcate: (0) absent; (1) present (Fig. 9C, D). Synapomorphy for the L. argus L. guerinii clade. 51. Male terminalia, aedeagus apically continuously narrowed, apex truncate, with a mesal large sclerite (resembling a happy face in dorsal view), a dorsal apical plate basally bifurcate and paired elongate ventral apical sclerites: (0) absent; (1) present (Fig. 9E, F). Synapomorphy for the L. campechianus L. lineicollis clade. 52. Male terminalia, aedeagus with paired apicolateral sclerites and a single mesally positioned acute sclerite, apex of aedeagus dorsally produced: (0) absent; (1) present (Fig. 9G, H). Synapomorphy for the L. lineatoguttatus L. vittatus clade. 53. Abdomen, anterolateral foveae on female sternum VII: (0) absent or not externally visible; (1) present (Fig. 10A, B). Convergently present in the D. abbreviatus E. quadrivittatus and I. viridiguttatus L. guerinii clades, with a subsequent reversal in the L. spretus L. mercator clade (l = 3; ci = 33; ri = 60). 54. Abdomen, shape of female tergum VIII: (0) transverse (Fig. 10C, D); (1) laterally compressed (Fig. 10E, F). DELTRAN optimisation preferred (Fig. 2), thus assuming three independent acquisitions of the apomorphic state. Convergently present in the L. coffeae L. yaucona clade and in the L. curvipes L. sparsimguttatus and L. hispidus L. guerinii clades, respectively (l = 3; ci = 33; ri = 84).

12 78 Invertebrate Systematics J. C. Girón and N. M. Franz A B C D E F Fig. 10. Female abdomen. Sternum VII of Lachnopus valgus: (A) dry specimen, (B) after KOH treatment; tergum VIII of L. campechianus:(c) dorsal view, (D) lateral view; tergum VIII of L. aurifer:(e) dorsal view, (F) lateral view. Scale bar = 1 mm. 55. Female terminalia, apical margin of coxites in lateral view: (0) sinuate (Fig. 11A); (1) mesally deeply emarginate, separating dorsal and ventral sections (Fig. 11B D). Coded as inapplicable for taxa in which the apical margins of the coxites are straight in lateral view. DELTRAN optimisation preferred (Fig. 2), thus assuming independent acquisitions of the apomorphic state. Convergently present in the L. curvipes L. luctuosus clade, the L. campechianus L. lineicollis clade, the L. proteus L. mercator clade and in the L. lineatoguttatus L. vittatus clade, with a subsequent reversal in L. spretus (Gyllenhal) (l = 5; ci = 20; ri = 69). 56. Female terminalia, shape of apex of ventral section of coxites in lateral view: (0) rounded (Fig. 11A C); (1) acute (Fig. 11D). Coded as inapplicable for taxa in which the apical margins of the coxites are straight in lateral view (thus lacking a separate ventral section, see character 55). Convergently present in the L. coffeae L. yaucona and the L. proteus L. mercator clades, with a subsequent reversal in L. spretus L. mercator clade (l = 3; ci = 33; ri = 71). 57. Female terminalia, width of ventral section of coxites in lateral view: (0) nearly as wide as dorsal section, and uniformly so (Fig. 11A, B); (1) apically narrower than dorsal section (Fig. 11C, D). Applicability as in character 56. Convergently present in D. abbreviatus, P. litus, the L. coffeae L. yaucona clade, the L. campechianus L. lineicollis clade and in the L. proteus L. mercator clade (l = 5; ci = 20; ri = 69). 58. Female terminalia, length of spermathecal cornu versus width of spermathecal corpus (see Velázquez de Castro 1997): (0) cornu less than 1.9 longer than width of corpus (Fig. 12A); (1) cornu more than 2 longer than width of corpus (Fig. 12B). Synapomorphy for the P. litus L. guerinii clade, with three subsequent reversals in L. luctuosus, L. campechianus and in the L. hispidus L. guerinii clade (l = 4; ci = 25; ri = 75). 59. Female terminalia, inner margin of spermathecal cornu (section close to corpus): (0) uniformly curved (Fig. 12B); (1) nearly straight (Fig. 12A). DELTRAN optimisation preferred (Fig. 2), thus supporting separate acquisitions of the apomorphic state in the L. hispidus L. gowdeyi and L. lineatoguttatus L. vittatus clades. Convergently present in D. abbreviatus, L. histrio, the L. coffeae L. yaucona clade, L. floridanus, the L. hispidus L. gowdeyi clade and in the L. lineatoguttatus L. vittatus clade (l = 6; ci = 16; ri = 50). 60. Female terminalia, annular sculpture on surface of spermathecal cornu: (0) absent; (1) present. DELTRAN optimisation preferred (Fig. 2), thus supporting independent acquisitions of this trait in L. histrio and in the I. viridiguttatus L. guerinii clade. Convergently present

13 Phylogeny of the Caribbean weevil genus Lachnopus Invertebrate Systematics 79 A C Fig. 11. Female terminalia, coxites, lateral view (membranes, internal sclerites and structural thickness omitted). (A) Lachnopus hispidus; (B) L. vittatus; (C) L. chlorophanus; (D) L. planifrons. Scale bar = 1 mm. A Fig. 12. Female terminalia, spermatheca. (A) Lachnopus vittatus; (B) L. chlorophanus. Scale bar = 0.05 mm. in L. histrio and in the I. viridiguttatus L. guerinii clade, with a subsequent reversal in the L. curvipes L. sparsimguttatus clade (l = 3; ci = 33; ri = 75). 61. Female terminalia, extension of annular sculpture on surface of spermathecal cornu: (0) along the whole cornu; (1) only at apex of cornu. Coded as inapplicable for species that lack annular sculpture on the surface of the spermathecal cornu (see character 60). Synapomorphy for the L. coffeae L. yaucona clade. Major inferred clades and relative support Under the current scheme of taxon sampling, the genus Lachnopus is rendered para- or even polyphyletic and is composed of a major monophyletic L. coffeae L. guerinii clade (Bremer support value (Bsv) = 2) that excludes L. albomaculatus, L. acuticollis and L. histrio (Figs 1, 2). Within the main L. coffeae L. guerinii clade there are several well-circumscribed subclades that are furthermore closely correlated with distinct biogeographic regions, as follows (Fig. 1; Table 1): (1) a well-supported Puerto Rican L. coffeae L. yaucona clade (Bsv = 5) placed in sister B D B relationship to the remaining species of Lachnopus (Bsv = 1); (2) a less well-supported clade with two subclades, (a) one including the widespread L. curvipes and L. valgus (Bsv = 2), and (b) the other including L. luctuosus and L. sparsimguttatus from Cuba and L. floridanus from Florida in an unresolved trichotomous relationship (Bsv = 2); (3) a well-supported Guadeloupian clade formed by L. campechianus and L. lineicollis (Chevrolat) (Bsv = 6), placed in a sister relationship with the distinctive and intriguing L. kofresi from Mona Island; (4) a speciose and reasonably well-supported L. proteus L. mercator clade with exclusively Hispaniolan species (Bsv = 3); and (5) a well-supported and essentially Cuban Jamaican clade (Bsv = 4) that contains two subclades, (a) the L. hispidus L. gowdeyi clade (Bsv = 2), which in turn includes the L. hispidus L. niveoirroratus and the L. aurifer L. gowdeyi clades, and (b) the L. lineatoguttatus L. guerinii clade (Bsv = 2), which harbours the strongly supported L. argus L. guerinii subclade (Bsv = 7). Discussion The correct tribal placement of the genus Lachnopus has been discussed by several authors, typically in reference to other Caribbean genera, such as Exophthalmus (see Kuschel 1955; Vaurie 1961; Franz 2010a). This study reaffirms the placement of Lachnopus within the Geonemini (sensu Alonso-Zarazaga and Lyal 1999) and thus separate from the Eustylini, in accordance with recent results based on morphological data (Franz 2012). Nevertheless, the Geonemini as a whole remain poorly circumscribed, requiring additional phylogenetic genuslevel analyses to corroborate the placement of Lachnopus and its closest relatives. In the following paragraphs we discuss the character evidence and geographic range for the inferred Lachnopus clade and subclades (Figs 1, 2). The main Lachnopus ingroup stands in sister relationship to the Caribbean genus Ischionoplus, for which two species have been described: I. niveoguttatus Chevrolat from Cuba and I. viridiguttatus from the Dominican Republic. Ischionoplus is readily separated from Lachnopus by the presence of a strong postocular constriction (see also Franz 2012), the presence of a spine on the ventral margin of the profemur and the absence of denticles on the tibiae (Fig. 8A; see van Emden 1944). The non-monophyly of Lachnopus is reflected in the diversity of the external appearance of their constituent species. In particular, L. albomaculatus and L. acuticollis are more closely related to the herein examined species of Eustylini, whereas the phylogenetic position of L. histrio, which furthermore exhibits several unique head features, is somewhat intermediate and perhaps closer to Pachnaeus (traditionally in Tanymecini, yet placed in Eustylini sensu Franz 2012). Given the limited outgroup sampling, we provisionally refrain from renaming and/or transferring these taxa to other tribes. The main Lachnopus clade shows no unreversed synapomorphies, and is only weakly supported by the following homoplasious traits: (1) the lateral margins of the rostrum are widened at, or apicad of, the antennal insertion (character 8; Fig. 4A, C, E; shared with the L. albomaculatus E. quadrivittatus outgroup clade and reversed to parallel to

14 80 Invertebrate Systematics J. C. Girón and N. M. Franz apically convergent (Fig. 4B, D) in the L. planifrons L. mercator and the L. hispidus L. gowdeyi clades); (2) the elytral interval IX is not particularly widened at any point (character 27; Fig. 7A; shared with the L. acuticollis E. quadrivittatus clade, reversed in the L. hispidus L. guerinii clade and regained in the L. hispidus L. niveoirroratus clade); (3) the ventral surface of the male metatibiae bears denticles (character 31; Fig. 8B; shared with the L. albomaculatus E. quadrivittatus clade of the outgroup); (4) the outline of the apical portion of the aedeagus is sinuate (character 36; shared with L. acuticollis and reversed in L. floridanus); and (5) the basal margin of the ventral surface of the aedeagus is widely and roundly emarginate (character 45; Fig. 9G; except in L. proteus and in the L. hispidus L. gowdeyi and L. argus L. guerinii clades, in which the basal margin of the ventral surface of the aedeagus is narrowly mesally emarginate (Fig. 9A)). This combination of homoplasious characters is not sufficient to establish a strong and unified redefinition and a clear delimitation of the genus, especially because the subclades within the main Lachnopus clade are well differentiated in the morphology and their geographic occurrence. The Puerto Rican L. coffeae L. yaucona clade is characterised by a consistent type of aedeagus; the apical sclerites are arranged as two lateral plates and one mesal plate, the latter apically truncate and slightly wider. These species also share an irregular profile of the ventral surface of the aedeagus in lateral view (as schematised by Marshall 1922). They exhibit considerable differences in size and in the density and colouration of scales; with L. coffeae being more densely covered with white scales on the lateral areas of elytra and thorax (as described by Marshall 1922 for the montanus variety), whereas L. seini Wolcott and L. yaucona Wolcott are more uniformly covered with yellow scales. Indeed, at this point there is not sufficient evidence to confirm the status of L. seini and L. yaucona as separate species. Given their sympatric occurrence (Wolcott 1948), additional studies will likely result in merging them into a single entity. The L. curvipes L. valgus clade is supported by a curved male metatibia accompanied by the presence of a large simple tooth in the proximal region of its inner margin (characters 30 and 32; see Fig. 8B). According to Wolcott (1923), these species are possibly synonyms; however, in our assessment several characters of the head and the aedeagus clearly separate them. Both species have a wide-ranging distribution including Puerto Rico and St Barthélemy as sympatric areas. In addition, L. valgus has been recorded for St Croix and Anguilla; whereas L. curvipes, the most widespread species of Lachnopus, is known to occur in Jamaica, the Dominican Republic, St Thomas, Tortola, St Kitts, Nevis and Guadeloupe (O Brien and Wibmer 1982). Individuals of the latter species show great intra- and inter-island variation in size and in the characteristics of the tegument and scale coverage. This variability is deserving of further study, both to reassess the taxonomic boundaries of the L. curvipes/l. valgus species complex and to properly understand the respective species distributions. The relationships within the L. floridanus L. sparsimguttatus clade are not fully resolved. In particular, L. floridanus is inferred as the sister taxon of either L. luctuosus or L. sparsimguttatus, which can be attributed to the lack of male specimens of L. luctuosus for study. Curiously, none of the most parsimonious trees group together L. luctuosus with L. sparsimguttatus, the Cuban species. The well-supported L. campechianus L. lineicollis clade is characterised by two unreversed synapomorphies: (1) the aedeagus has a mesal large sclerite resembling a happy face in dorsal view, with a dorsal apical plate that is basally bifurcate and paired elongate ventral apical sclerites (character 51; see Fig. 9E, F); and (2) the endophallus lacks small papillae (character 40; see Fig. 9E, F). The two constituent species occur on Guadeloupe, and are inferred as sister to the enigmatic L. kofresi. The latter species is endemic to Mona Island (Franz et al. 2009) and presents a poor match with the traditional gestalt of Lachnopus. The L. proteus L. guerinii clade is supported by a single synapomorphy; namely, the lateral margins of the male elytra are continuously posteriorly converging (character 20; see Fig. 6A C). The clade is furthermore distinguished by the presence of a shallow occipital suture (character 7; Fig. 5A) and the absence of the luctuosus-type of scale coverage (character 25; see Fig. 6D), although both features are homoplasious. The L. proteus L. guerinii clade includes species reported from Jamaica, Cuba and Hispaniola. The therein contained L. proteus L. mercator subclade (Hispaniola) is characterised by a synapomorphic, uniform and mostly green iridescent scale coverage (character 21; Fig. 6B; not present in the L. atramentarius L. inconditus; in L. spretus the green iridescent scales are restricted to the sides of the body). Additional homoplasious characters supporting this clade are related to the configuration of the female coxites. The incorporation of male specimens of L. chlorophanus and L. mundus and female specimens of L. mercator may increase phylogenetic resolution in this clade. According to this study there is insufficient evidence to clearly separate L. atramentarius (Gyllenhal) and L. inconditus as different species. Moreover, a comprehensive representation of Lachnopus species throughout Hispaniola and surrounding islands may lead to the erection of the L. proteus L. mercator clade as a separate genus. Members of the L. hispidus L. guerinii clade include the majority of Cuban and Jamaican species treated in this study. The clade is supported by a single synapomorphy namely, the presence of a carinate posterior margin of the pronotum (character 17; Fig. 6A) and six homoplasious characters: (1) the acute posterior corners of pronotum (character 18; Fig. 6C); (2) the reduced to absent elytral humeri (character 19; Fig. 6C); (3) the not particularly widened elytral interval IX (character 27; Fig. 7A; except in the L. hispidus L. niveoirroratus clade); (4) the strongly expanded apex of the apodeme of the spiculum gastrale (character 35); (5) the laterally compressed female tergum VIII (character 54; Fig. 10A); and (6) the relatively short spermathecal cornu (character 58; Fig. 12A). The L. hispidus L. gowdeyi subclade is supported by a single synapomorphy i.e. the slender shape of the aedeagus (character 49; Fig. 9A) and four homoplasious characters. This clade is subdivided into two clades. The first of these, the L. aurifer L. gowdeyi clade, is supported by three synapomorphic characteristics: (1) a sharply raised mandibular scar (character 14; Fig. 4D); (2) a unique elytral scale pattern ( aurifer-type of scale coverage; character 24; Fig. 6C); and (3) the presence of tiny denticles on the surface of male metafemora (character 29;