Revision and phylogeny of the Caribbean weevil genus Apotomoderes Dejean , 1834 ( Coleoptera , Curculionidae , Entiminae )

Th e weevil genus Apotomoderes Dejean, 1834 (Curculionidae: Entiminae: Geonemini) is revised, including a redescription of the only previously known species, A. lateralis (Gyllenhal, 1834), and descriptions of fi ve new species: A. anodontos sp. n., A. menocrater sp. n., A. sotomayorae sp. n., A. chariedris sp. n., and A. hadroprion sp. n. Th e monophyly of Apotomoderes is supported by multiple synapomorphic character states including the two-segmented labial palps, a postocular constriction on the head, a sexually dimorphic, globular pronotum in males, and the presence of setae in the dorsal subapical region of the aedeagus. In addition, all species of Apotomoderes except A. anodontos have a large, knife-like cuticular tooth on the profemur and a toothed ridge along the anteromesal margin of the protibia. Illustrations of external and internal morphological traits are provided, along with a key to the six constituent species. A cladistic analysis of 12 taxa (6 outgroup, 6 ingroup) and 22 characters yielded a single most parsimonious cladogram (L=33, CI=75, RI=90) with the topology (A. anodontos, (A. menocrater, (A. sotomayorae, (A. lateralis, (A. chariedris, A. hadroprion))))). A species of Artipus Sahlberg (Naupactini) was placed as the most immediate relative of Apotomoderes; however, the state of phylogenetic knowledge of Caribbean entimine weevil is still too incomplete to warrant any higher level rearrangements. All species of Apotomoderes occur on Hispaniola with the exception of A. sotomayorae which is endemic to Mona Island, Puerto Rico. A historical biogeographic reconstruction yielded the taxon-area cladogram (southwestern Dominican Republic, (eastern Dominican Republic, Mona Island)), suggesting two successive eastbound colonization events in the Miocene/Pliocene, originating from the southern Hispaniola peninsula. Reliable host plant records are unavailable although adults of A. menocrater have been found on allspice (Pimenta Lindley; Myrtaceae) and lignum vitae (Guaiacum Linnaeus; Zygophyllaceae). ZooKeys 49: 33–75 (2010) doi: 10.3897/zookeys.49.303 www.pensoftonline.net/zookeys Copyright N.M. Franz . This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. RESEARCH ARTICLE Launched to accelerate biodiversity research A peer-reviewed open-access journal


Introduction
Th e entimine weevil genus Apotomoderes Dejean (1834: 253) was originally proposed to accommodate a single species, A. lateralis (Gyllenhal 1834: 45), from Hispaniola.Dejean had transferred this species out of Schoenherr's Apotomus (1834: 44), a name which was published just weeks earlier (Alonso-Zarazaga and Lyal 1999) yet which constitutes a junior homonym of Apotomus Illiger (1807: 348), a genus of Carabidae.Consequently, Eurilia Laporte (1840: 308) is an unnecessary replacement name for Apotomus Schoenherr, and also a synonym of Apotomoderes.Both Dejean (1834) and Schoenherr (1840) incorrectly attributed the valid genus and species name to Carl Gustav Mannerheim who had described the species in a letter to Schoenherr and then donated the type specimen to Schoenherr's museum.For a relatively short period of time, a second species, A. albicans (Lacordaire 1863a: 21) was placed in this genus, until it was transferred to Megalostylus Schoenherr (1840: 114) by Chevrolat in 1878 (p. LXVI;as M. farinosus [junior synonym]) and then by Pascoe 1881 (p.42; as M. expansus [junior synonym]), as reviewed in Champion (1911: 241-246).Th us in effect, no other species of Apotomoderes have been described since 1834 (O'Brien and Wibmer 1982;Pérez-Gelabert 2008).Schoenherr (1834) provided the most detailed description of Apotomoderes to date, emphasizing the postocular constriction of the head, prominent eyes, expanded pronotum, subovate elytra, and conspicuously toothed profemora as diagnostic features.He placed the genus in his division Brachyderides, and particularly in the vicinity of the Old World genus Cratopus Schoenherr which is presently assigned to the tribe Cratopodini Hustache (Lyal and Alonso-Zarazaga 2006).Lacordaire (1863b: 25-27, 64-66) classifi ed Apomotoderes in the Naupactides (= Naupactini Gistel), but distinguished the genus from all other members of this tribe in light of the posteriorly constricted head.Lacordaire (1863b: 81-82) also mentioned a possible affi nity of Apotomoderes with Megalostylus.Interestingly, his description of "A.lateralis" from Haiti ('uniform opal grey, epipleura of elytra chalk white') applies to another species, described below.Subsequently, the genus was placed in the tribe Barynotini Lacordaire (e.g. Blackwelder 1947: 796, O'Brien andWibmer 1982: 41) whose current valid name is Geonemini Gistel (Alonso-Zarazaga and Lyal 1999).However, none of the past or present tribal placements of Apotomoderes are well corroborated given that the mid-level classifi cation of entimine weevils remains poorly understood (Th ompson 1992;Marvaldi 1997;Oberprieler et al. 2007).
Field work in the Dominican Republic and on Mona Island, Puerto Rico, has brought to light several new species of Apotomoderes.Th ese species are herein described and illustrated, and the genus is revised, including a generic redefi nition, redescription of the type species, key to all species, phylogenetic analysis, and historical biogeographic reconstruction.

Methods
Morphological studies.Th e descriptive sequence used in this study is in accordance with Franz and Girón (2009).Morphological terms usually follow Torre-Bueno (Nichols 1989); with additional specialized terms used for the mouthparts (Ting 1936;Morimoto and Kojima 2003), apex of rostrum (Vaurie 1963), metendosternite (Velázquez de Castro 1998), wings (Zherikhin and Gratshev 1995), tibial apices and abdominal segments (Th ompson 1992), and male and female terminalia (Howden 1995;Velázquez de Castro 1997;Wanat 2007).Specimens were observed with a Leica MZ16 stereomicroscope (magnifi cation: 7-115×) and an Olympus BX41 compound microscope (magnifi cation: 20-400×), each equipped with an ocular graticule for length measurements.Th e overall body length was measured (in dorsal view) from the anterior margin of the eye to the posterior margin of the elytra; whereas the length of the rostrum was measured from its apex to the anterior margin of the eye.Th e number of individuals measured is written once in parentheses for all measurements of external structures; another separate number refl ects the number of measurements for the length/width relations of the wings and the median lobe.Th e numbering of the abdominal ventrites refl ects their homology within Curculionoidea (Th ompson 1992).Th e original species accounts were trimmed to emphasize diagnostic features and characters with potential phylogenetic relevance.Features that are shared between the male and female are usually mentioned only once (in the male), as are similar traits on serially homologous structures such as the legs.
Th e habitus photographs were taken with a Microptics XLT imaging system.All drawings of internal structures were prepared with a drawing tube attached to the compound microscope.Th e initial, simplifi ed line sketches were scanned and redrawn using an illustration software program, thus highlighting features with diagnostic signifi cance.
Type material.Th e labels for new type specimens include the genus name and species epithet, a gender symbol, and the author and year.Th ey are colored red for the holotypes and yellow for all paratypes.Th e insect collection codens are based on Arnett et al. (1993) Lanteri 1990; though see Scataglini et al. 2005 for a later perspective).Th e selection was informed in part by preliminary results from phylogenetic studies of Caribbean entimine weevils (NMF, unpublished), and the cladogram was rooted accordingly with E. quadrivittatus.Autapomorphies for species of Apotomoderes are presented in the species accounts yet were excluded from the cladistic analysis.Reductive coding (Strong and Lipscomb 1999) was use in select instances to represent homology assessments pertaining to particular species groups.Th e character matrix was compiled, edited and refi ned using the matrix and tree interfaces in WinClada (Nixon 2002).Th e characters were numbered based on their sequence of appearance in the taxonomic descriptions.Th e most parsimonious cladogram and character state optimizations were identifi ed using NONA (Goloboff 1999).An exhaustive search of the tree space for the 12-taxon matrix was performed using the commands "whennig" and "mswap+".Finally, branch support values (Bremer 1994) were calculated in NONA with "hold 10000", "suboptimal 20" and "bsupport 20".
Historical biogeography.A historical vicariance analysis of Apotomoderes was performed through integration of the species phylogeny with the inferred areas of endemism (Morrone 2009).In light of an obvious alignment of the phylogeny and geographic areas, only assumption 0 sensu Nelson and Platnick (1981;component analysis) was employed to generate a taxon-area cladogram (see also Page 1990).Th e resulting scenario was reconciled with available geological information (Iturralde-Vinent 2006).Type species: Apotomoderes lateralis (Gyllenhal, 1834: 45), by monotypy (originally as Apotomus lateralis).

Diagnosis.
Apotomoderes keys to Artipus Sahlberg in Anderson (2002), by virtue of the absence of a postocular lobe and vibrissae, the laterally positioned scrobe, the long median sulcus on the rostrum, laterally positioned eyes, and the conspicuous epistoma.However, Apotomoderes can be distinguished from this and other genera of Caribbean entimine weevils (cf.O'Brien and Wibmer 1982) by a unique combination of synapomorphic and/or diagnostic features; viz.scale coverage complete; labial palps 2-segmented (Fig. 3B); rostrum dorsally with a transverse depression (e.g.Fig. 13B); antennal scape only moderately stout; head with a postocular constriction (e.g.Fig. 9B); eyes protruded, globular; pronotum sexually dimorphic, larger and more globular in males (e.g.Fig. 11A); metatibial apex simple, without a bevel or fl ange ("corbel open"), margins lined with spiniform setae, surface surrounding condyle partially covered with scales; presence of setae in the dorsal subapical region of the aedeagus (e.g.Fig. 5D); aedeagus with a pair of weakly sclerotized rami along ostium; and spermatheca with collum and ramus subcontiguous (e.g.Fig. 17C).In addition, nearly all species of Apotomoderes have a large, knife-like cuticular tooth only on the profemur and a conspicuously toothed ridge along the anteromesal margin of the protibia (in idealized orientation of legs) in both males and females (e.g.Fig. 13C).Th e genera and species most closely related to Apotomoderes are likely still undescribed (NMF, personal observation), but diff er from this taxon minimally by the shape of the head and pronotum.
Antennae 11-segmented.Scape moderately stout, nearly straight, clavate; extending to region between posterior margin of eye and anterolateral margin of pronotum, directed ventrad of eye in idealized position; covered with linear scales.Funicle 7-segmented, slightly longer than scape; funicular antennomeres progressing from elongate to equilateral, clavate, covered with appressed and suberect, linear scales; I and II (usually) similar in length.Club 3-segmented, similar in length to funicular antennomeres III-VI, nearly 3× longer than wide, dark brown, pubescent; I and II similar in length; III slightly longer, conical, transversely sutured.
Scutellum exposed, semi-circular, slightly wider than long, anterior margin straight, posterior margin rounded, covered with creamy white scales.
Prosternum (Figs. 1C, 14B) longer than mesosternum, with a transverse sulcus on each side anteriorly and posteriorly of procoxal cavities; procoxal cavities positioned at midpoint, contiguous, prosternal process short, elevated.Mesosternum slightly shorter than metasternum, strongly angulate, mesocoxal process slightly elevated; anterior half coved with plumose scales, posterior half with longer, suberect, linear scales; mesocoxal cavities separated by distance nearly half as long as width of each mesocoxal cavity.Metasternum with median sulcus present as a large, transverse fovea positioned anteriad of posterior margin, surface slightly undulate, posterolateral regions set off (with small posterior "face"), and with linear scales (see above); metacoxal cavities separated by distance slightly shorter than width of each mesocoxal cavity.Metendosternite (Fig. 4A) with stalk slightly shorter than furcal arms, ventral margin 2-3× wider than dorsal width of stalk; hemiducts wide, truncate; dorsal margin of sheath undulate; anterior tendons positioned near midpoint between median keel and base of furcal arms; furcal arms diverging at 30-45° in relation to medial keel.
Legs.Prothoracic legs longer than mesothoracic legs; scale colors variously interspersed, though forming a more uniform light/dark/light pattern on femora, metallic scales often present on anterior sides of profemora and near tibial apices; ventral sides of tibiae with rows of longer, whitish or yellowish, suberect setae.Profemur/pronotum length ratio 0.80-1.08;profemur moderately stout, in cross-section subcircular ; (usually) with 1 very large, knife-like cuticular tooth inserted at apical 2/5 on anteromesal margin, tooth ventrally directed (absent in one examined species).Protibia/profemur length ratio 0.84-1.02;protibia moderately stout, nearly straight, in cross-section elliptical, apically slightly expanded; anteromesal margin with a distinct, laminate, rounded or triangular projection near basal 2/5 (absent in one examined species), from thereon to apex with row of 10-15 smaller, apically rounded or (oblique) truncate cuticular teeth (teeth larger along laminate projection), each tooth distally with an aurate, spiniform seta; protibial apex with anterior margin truncate, setal comb absent; mucro similar in length to tarsal claw, surpassed by tufts of aurate setae.Protarsus with tarsomere I nearly 2× as long as II, elongate, clavate; II and III similar in length, equilateral to transverse; and jointly similar in length to V; claw paired, separate, simple.Meso-and metathoracic legs slighter shorter and longer than prothoracic legs, respectively; meso-and metafemora unarmed; meso-and metatibiae lacking laminate projec- tions, teeth along anteromesal margin less pronounced; metatibial apex simple, with anterior margin curved (on side opposed to mucro), projected, margins with a comb of spiniform setae of increasing length, yet without a bevel or fl ange ("corbel open"), surface surrounding metatarsal condyle glabrous to partially covered with (metallic) scales "invading" surface from dorsal side.
Elytra.Length/width ratio 1.52-1.90;widest at humeri (to anterior third); anterior margins jointly wider than posterior margin of pronotum, sinuate; humeri obliquely rounded; lateral margins posteriorly slightly converging in anterior half, thereafter more strongly converging, rounded, posteriorly attenuate; posterior margins narrowly truncate.Elytra in lateral view slightly convex; posterior declivity apparent though not strongly angulate, nearly straight.Elytra with 9 complete striae and 1 incomplete stria; striae similar in width to intervals; stria X indistinct ("merging with stria IX") in second third of entire length; punctures separated by distance similar to, or slightly longer than, width of each puncture; intervals slightly elevated; scales covering entire integument, arranged in mostly irregular micro-patterns of creamy white to dark brown hues, though often locally also with iridescent metallic scales, including (in most species) a single colored or metallic scale in each strial puncture.Wings.Wings (Fig. 4B) fully developed (absent in one examined species), elongate (linear), wing/body length ratio 1.15-1.35,wing length/width ratio 3.30-3.78;anterior margin slightly angulate near midpoint; posterior margin nearly straight, undulate in anal area; anal lobe absent.Alar veins well defi ned: C, Sc, R, Rr and rs distinct, radial cell present; M r and Cu 1 short, r-m reduced; 2A distinct, 3A and 4A weaker; R 3 distinct, M 1 reduced, only apparent distally.
Variation.Variation in size, shape (relative expansion) of the pronotum, "armature" of the profemur and protibia, and particularly in scale patterns is considerable within and among species of Apotomoderes.In spite of the remarkable palette of scale colors -ranging from creamy white to dark brown and including several hues of iridescent scales that can occur on many parts of the rostrum, pronotum, elytra and legs -these patterns are often too variable to clearly separate species.In older, worn specimens, large portions or nearly all of the scales are lost, or have turned transparent, exposing the underlying, dark reddish brown integument.Some minor intraspecifi c variation is apparent in the shape of the male aedeagus and female spermatheca.
Natural history.Apotomoderes is restricted to Hispaniola -with two apparently disjunct regions of occurrence in the southwestern and eastern parts of the islandand to Mona Island, Puerto Rico (Fig. 18).Th e sampled species tend to inhabit lower elevation coastal dry forests.No specifi c host plant associations are known.It is likely that the species have fairly broad host ranges within their particular habitats, as observed generally in many entimine lineages (e.g.Woodruff 1985;Oberprieler et al. 2007;Franz and Girón 2009).
Etymology.Th e name Apotomoderes is derived from the name Apotomus (see above), based on the Greek term apo-tomós = abruptly, curtly; and the Greek term deire = neck, throat.Th us the name likely refers to the postocular head constriction.Th e gender is masculine.
Apotomoderes anodontos Franz, sp.n. urn:lsid:zoobank.org:act:A4DF9315-045D-4684-9A7D-0C04BA623BD7Figs. 7, 8 Diagnosis.Apotomoderes anodontos is the smallest of the herein treated species and is readily diff erentiated from other species by the absence of a large, knife-like tooth of the profemur and the lack of a ridge-like, toothed projection along the anteromesal margin of the protibia (Fig. 7C).
Description -male.Length 4.32-6.28mm, width 1.65-2.58mm, length/width ratio 2.40-2.62(N=5), widest at humeri to anterior third of elytra or near midpoint of pronotum.Linear piliform scales moderately long and abundant, particularly on legs, elytra, and thoracic and ventral sterna, either transparent white or light (rusty) brown, recumbent to suberect, more appressed on pronotum.With characteristic though locally variable patterns of creamy white and (light) (rusty) brown or tan scales, lighter scales typically more abundant on lateral sides and on legs, creating a two-colored impression, some specimens with semi-regular micro-patterns on elytra, (very) pale blue, moderately iridescent, metallic scales primarily on rostrum and tibiae.Rostrum (Figs. 7B, 7C) short and wide, length 0.45-0.68mm, rostrum/pronotum length ratio 0.28-0.30,rostrum length/width ratio 0.60-0.66,depressed region basad of epistoma covered with creamy white (slightly iridescent) to (very) pale blue or light greenish metallic scales, thereafter increasingly with tan scales, lateral sulcus anteriad of eye reduced.Rostrum in lateral view nearly 2× wider (dorsoventrally) than long.Ventral margin of scrobe strongly angulate.Scape extending to anterolateral margin of pronotum, laterally compressed, covered with linear and subcircular scales.Funicular segment I longer than II.Head with eyes relatively large, moderately and almost evenly globular (posteriorly not abruptly curved), anterior, ventral, and posterior margins nearly straight, eyes separated (in dorsal view) by distance (much) less than 2× anterior-to-posterior length of each eye, scales on head predominantly creamy white and tan.Pronotum length/width ratio 0.88-0.98,pronotum/elytra length ratio 0.53-0.55,laterally strongly expanded (wide) though not strongly globular (more plane than in other species), scales dorsally predominantly darker, rusty or tan, irregularly shaped, though often with a subquadrate patch of lighter scales mesally near posterior margin, laterally with creamy white scales, creating a two-colored impression.Metendosternite similar to A. lateralis though furcal arms shorter.Legs predominantly covered with creamy white, rusty, and tan scales (more rarely with pale blue metallic scales, profemur/pronotum length ratio 0.80-0.83,profemur unarmed (lacking anteromesal tooth), protibia/profemur length ratio 0.84-1.02,lacking laminate, rounded anteromesal projection, though with 6-10 alternating smaller and larger, triangular (apically subacute) teeth, less much less prominent on meso-and metatibiae.Elytra (Fig. 7A) length/width ratio 1.60-1.68,scales dorsally (~ striae I-VI) predominantly darker, rusty brown or tan, sometimes with irregular patches of lighter scales, laterally (~ striae VII-X) with creamy white scales more abundant, creating the impression of a lighter stripe, punctures also with a small, creamy white scale.Wings absent.
Variation.Larger males have a more strongly expanded pronotum.Some specimens are primarily covered with red or rusty brown scales, other specimens have larger region with an underlying pale blue or greenish metallic hue.
Etymology.Named for the unarmed profemora, with an-signifying "not" and odontos signifying "tooth" (Brown 1956).Th e epithet is treated as a noun in apposition.
Natural history.Apotomoderes anodontos is known to occur in the low to mid elevation dry forests (100-670 m) of the southwestern Pedernales province of the Dominican Republic: Sierra de Bahoruco, Cabo Rojo, and Oviedo (Fig. 18).Th e host plant associations remain unknown.
Apotomoderes menocrater Franz, sp.n. urn:lsid:zoobank.org:act:D196352B-4B83-4A30-9633-6A2BF03A25D6Figs. 9, 10 Diagnosis.Apotomoderes menocrater is most readily separated from other congeneric species by the larger, subfoveate punctures in the posterior half of the pronotum (Fig. 9A, 9C).Other diagnostic features include a more elongate shape and more tubular pronotum (particularly in males), the presence of grey metallic scales (though see comments on variation), only moderately protruded and more evenly convex eyes (Fig. 9B), the presence of a patch of suberect scales near the mesal anterior end of the declivity in females (Fig. 9D; shared with A. anodontos), the apparently V-shaped uncinate rami along the ostium of the aedeagus (Fig. 10A), and the strongly curved cornu of the spermatheca (Fig. 10C).
Description -male.Length 5.40-9.05mm, width 2.00-3.50mm, length/width ratio 2.58-2.70(N=10), more elongate than oval, widest at humeri.Linear piliform scales relatively sparse, transparent white or yellow to light brown.With variable patterns of creamy white, gray metallic (several hues), light rusty brown (less abundant), and variously grayish or light to dark brown scales, particularly the creamy white and gray metallic scales have an underlying, greenish-turquoise-pinkish-yellowish iridescence ("opal gray"; cf.Lacordaire 1963b: 82).Rostrum length 0.70-1.12mm, rostrum/pronotum length ratio 0.38-0.42,rostrum length/width ratio 0.78-0.80,depressed region basad of epistoma covered with pale pink metallic scales, thereafter increasingly with creamy white and variously brownish scales.Head (Fig. 9C) with eyes (only moderately) protruded, almost evenly globular, posteriorly not abruptly curved, rostrum and head in lateral view only moderately angulate.Pronotum (Fig. 9C) length/width ratio 1.00-1.22,pronotum/elytra length ratio 0.42-0.48,pronotum equilateral to elongate, relatively narrow and only slightly globular (subtubular in small males), anteriorly slightly constricted, punctures in posterior half of pronotum larger, subfoveate, scales dorsally predominantly darker (grayish and brown), laterally with wider, irregularly shaped and variously interspersed stripe of creamy white scales.Metendosternite more elongate than in A. lateralis (less laterally expanded), ventral margin nearly 2× wider than dorsal width of stalk, furcal arms diverging at nearly 30° in relation to medial keel.Profemur/pronotum length ratio 1.06-1.10,profemur with anteromesal tooth only moderately large (in comparison with males of congeneric species), triangular, protibia/profemur length ratio 0.96-1.00,anteromesal projection and associated teeth of protibia only weakly projected, basally rounded, scales predominantly creamy white and variously brown, interspersed with pale turquoise and pinkish metallic.Elytra length/width ratio 1.78-1.92,more narrowly triangular in general appearance, widest at humeri, thereafter slightly yet continuously narrowed (attenuate), elytra predominantly with creamy white, gray metallic, and brown scales, each color varying in abundance and micro-patterns, in some specimens interspersed with light rusty brown and pale blue metallic scales, punctures with or without a creamy to transparent white scale.Wing/body length ratio 1.23-1.35,wing length/ width ratio 3.30-3.62(N=2).
Terminalia with sclerites of sternum VIII diamond-shaped, posterior margin strongly angulate.Spiculum gastrale with furcal arms apically strongly curved out- wards.Aedeagus (Fig. 10A) with median lobe length/width relation 7.08-8.00(N=5), lateral margins subparallel in basal 5/6 (greatest width near mid region), thereafter nearly straight and converging to very small, lobe-like, apically narrowly rounded projection.Median lobe in lateral view only slightly (though homogeneously) curved, mid region (second and third fourth of entire length) nearly straight, greatest width near base of ostium, apically with a small, very narrow lobe-like projection that is minimally expanded and reclined.Internal sac with 2 moderately sclerotized, uncinate, apically obliquely truncate rami, positioned in ostium and reclined mesally, creating the impression of a V-shaped transparent area in apical half of ostium.
Etymology.Named for the characteristic, subfoveate punctures on the pronotum that resemble a moon crater landscape, with mene signifying "moon" and krater signifying "vessel, crater" (Brown 1956).
Natural history.Apotomoderes menocrater is known to occur in the lower elevation coastal dry forest habitats of the southwestern Pedernales province of the Dominican Republic: Jaragua National Park, Cabo Roho, Los Tres Charcos, and Oviedo (Fig. 18).Th e species likely also occurs in southern Haiti (cf.Lacordaire 1963b: 82).Label information (G.Dominici) suggests an association of adults with "lignum vitae" ("guayacán"; Guaiacum offi cinale Linnaeus -Zygophyllaceae) and "pimenta" ("allspice"; Pimenta Lindley -Myrtaceae); although the specifi c host plant associations remain unknown.Diagnosis.Apotomoderes sotomayorae, the only species occurring on Mona Island, is most readily separated from other congeners by the presence of regularly appearing, creamy white scales along the elytral striae (Fig. 11A; less distinctive in females).Other diagnostic features include the presence of transparent scales, particular on the pronotum of males (Fig. 11B), the lack of a pronounced elytral declivity in males and females (Fig. 11C), the apically triangular, point-like aedeagus (Fig. 12A), and the relatively narrow triangular lamina of the sternum VIII in females (Fig. 12B).Large males of similarly shaped species (A.lateralis, A. chariedris, and A. hadroprion) have a basally triangularly projected (as opposed to rounded) protibial ridge.
Terminalia with tegmen similar in length to median lobe.Aedeagus (Fig. 12A) with median lobe length/width relation 6.12-6.85(N=3), basiventral margin strongly emarginate, lobe-like projections subtriangular, lateral margins very slightly diverging in basal 5/6 of entire length, thereafter nearly straight and triangularly converging towards point-like, through very narrowly rounded apex.Median lobe in lateral view apically with small, knob-like projection that is neither expanded nor reclined.Internal sac with 2 weakly sclerotized, angulate-uncinate rami, positioned in ostium and reclined mesobasally.
Female.Length 7.66-9.04mm, width 2.82-3.40mm, length/width ratio 2.58-2.72 (N=3).Pattern of scale colors varying considerably from that of males: rostrum, pronotum, sterna, legs, and elytra more homogeneously and densely covered with scales of multiple colors including creamy white, pale yellow (rare), light rusty brown, dark brown (as opposed to transparent), pale blue metallic, and yellow-light-green-pink metallic.Scale colors arranged in semi-consistent patterns though particularly variable among specimens on rostrum and pronotum, and transparent scales (present in males) absent on pronotum and elytra, therefore not appearing "glabrous" and with less conspicuous pronotal or strial scale stripes.Brown scales more prominent on rostrum (not just anteriad of eyes).Legs more consistently covered with creamy white, light rusty brown, and variously metallic scales.Region of elytra just posteriad of scutellum with patch of metallic scales.Rostrum length 0.85-0.98mm, rostrum/pronotum length ratio 0.39-0.43,rostrum length/width ratio 0.76-0.80.Pronotum length/width ra- tio 0.98-1.04,pronotum/elytra length ratio 0.36-0.38,pronotum considerably less globular than in males.Legs more slender than in males, profemur/pronotum length ratio 0.98-1.08,profemoral tooth (much) smaller and more triangular than in males, protibia/profemur length ratio 0.97-1.03,anteromesal projection and associated teeth of protibia less pronounced.Elytra length/width ratio 1.80-1.93,posterior declivity similarly unpronounced as in males.
Variation.Males vary considerably in size, and several presumably allometric characteristics -viz.expansion of the pronotum, profemoral and protibial teeth, posterior attenuation of elytra -are less pronounced in smaller males.In some males yellow scales are more abundant in the lateral pronotal area.Th e relative abundance of creamy white, light rusty brown, and dark brown scales varies in females, particularly on the elytra, thus making the appearance of alternating stripes of darker scales (intervals) and lighter scales (stria) more or less obvious.
Etymology.Named in honor of Sonia Maria Sotomayor, Associate Justice of the Supreme Court of the United States, whose personal story and academic and professional achievements are an inspiration for people in Puerto Rico and elsewhere.
Natural history.Apotomoderes sotomayorae is endemic to Mona Island, Puerto Rico (Fig. 18), where it is likely widespread, though not particularly abundant, in the plateau and depression forests (Cintrón and Rogers 1991).Most specimens were taken at night, on a variety of shrubs and trees.

Apotomoderes lateralis (Gyllenhal, 1834)
Figs. 1,2,3,4,5,6 = Apotomus lateralis Gyllenhal 1834: 45 (fi rst valid combination in Dejean 1834: 253) Diagnosis.Apotomoderes lateralis, the type species, most closely resembles A. chariedris and A. hadroprion though it is slightly smaller than the latter two species, and has a slightly less expanded male pronotum Fig. 1A).Th e scale patterns on the elytra are usually more homogeneous (initially appearing a one primary color) and less "spotty" (Figs.2A, 2B).In addition, the aedeagus is apically more widely rounded, and the internal sac lacks well sclerotized rami along the ostium (Fig. 5D).Th e spermatheca is C-shaped (as opposed to more strongly curved), and has a short, tubular (as opposed to almost completely reduced) ramus (Fig. 6C).See also the species accounts of A. chariedris and A. hadroprion.
Terminalia with lamina of sternum VIII (Fig. 6A) triangular, slightly longer than wide, anterior margins slightly projected, posterior margin narrowly rounded.Spermatheca (Fig. 6C) C-shaped, collum very short (reduced), collum subcontiguous with, and angled at nearly 90° in relation to, moderately long ramus, corpus very short and expanded (widest), cornu continuously curved to end (at nearly 150° in relation to proximal orientation), gradually narrowed, apically very narrowly rounded.
Variation.Th e examined specimens vary primarily in the abundance of scales on the integument.
Etymology.Th e Latin term lateralis means "of the side" (Brown 1956), and may refer to the relatively well defi ned lateral stripe of creamy white scales on the pronotum and elytra of the female type specimen.
Natural history.Apotomoderes lateralis is known to occur in low elevation habitats in the southeastern provinces of the Dominican Republic (Fig. 18): La Altagracia (Boca de Yuma) and San Pedro de Macoris/Santo Domingo (east of Boca Chica).Th e host plant associations remain unknown.
Description -male.Length 8.90-9.70mm, width 3.58-3.78mm, length/width ratio 2.38-2.60 (N=3).Linear piliform scales more dense on pronotum, very short, appressed, transparent white; setae (much) longer and suberect on thoracic and ventral sterna and coxae.Scales predominantly creamy white, with a subtle iridescence, creating an almost silver eff ect, variously interspersed with rusty brown (localized, rare), tan, dark brown (more frequent, patchy/spotty) and turquoise (green to light blue) metallic scales (particularly as a secondary color on pronotum, anterior side of profemora), depressed region basad of epistoma covered primarily with creamy white to turquoise metallic scales, thereafter increasingly with creamy white scales, lateral region anteriad of eye with patch of light brown scales, sulcus reduced, region adjacent to mandibular incision with pale blue metallic scales.Rostrum length 0.98-1.00mm, rostrum/pronotum length ratio 0.31-0.33,rostrum length/width ratio 0.75-0.80.Head with eyes "tilted" posteriad, particularly in large males.Pronotum length/width ratio 0.91-0.97,pronotum/elytra length ratio 0.48-0.52,globular and strongly convex, dorsally with irregularly shape scales of various colors including dark brown and pale blue metallic patches, in some specimens laterally (posterior half ) with a wide, more uniformly creamy white "stripe", fl anked by darker regions.Profemur/pronotum length ratio 0.90-0.98,profemur (Fig. 13C) with (minimally) anterior side often covered in part with turquoise scales, with anteromesal tooth very large, also completely covered with scales, protibia/profemur length ratio 0.98-1.02,anteromesal projection of protibia (Fig. 13C) strongly and triangularly elevated (not rounded), associated teeth large, apically rounded to truncate.Elytra (Fig. 13A) length/width ratio 1.66-1.68,scales predominantly creamy white, often with turquoise metallic undertones, mesal striae more beige or tan, and with distinct though irregularly positioned and shaped dark brown spots, punctures also with a small, turquoise metallic scale.
Variation.Th e examined males vary primarily in the abundance of darker brown scales on the pronotum (spotty to predominant) and elytra (near absent to very con- spicuous); one specimen has multiple patches of rusty brown scales on the legs.Th e females diff er more strongly in size and in the overall scale color appearance, with some females have a more bluish white or rusty orange brown pattern.Material examined.Male holotype "DOMINICAN REPUBLIC, RD-55 ~ 2 km N Bayahibe, La Altagracia Prov., 31.vii.2002, 18°23.423' N, 68°50.453' W, D. Perez, R. Bastardo, B. Hierro" (MHND).Paratypes, same label information as male holotype (MHND: 2 males, 3 females).
Etymology.Named for the visually appealing scale colors and patterns, with charieis signifying "graceful" (Brown 1956), and the inserted letters dr (charie-DR-is) representing the initials of the Dominican Republic where the species occurs.Th e epithet is treated as an adjective.
Natural history.Apotomoderes chariedris is known to occur in coastal, humid forest habitats of the southeastern La Altagracia Province (Bayahibe, Parque del Este) of the Dominican Republic (Fig. 18).Th e host plant associations remain unknown.Diagnosis.Apotomoderes hadroprion closely resembles A. chariedris though it has a more patchy beige/dark brown scale pattern on the elytra (Fig. 16A).Th e pronotum is strongly globular (Fig. 16B) and the protibial ridge is distinctly triangular near the basal 2/5, particularly in large males.In addition, the apex of the aedeagus is point-like and not just narrowly rounded, and in lateral view forms a small, lobe-like (as opposed to knob-like) projection that is neither expanded nor reclined (Fig. 17A).Two mesally reclined, uncinate rami are present along the aedeagal ostium.Th e cornu of the spermatheca is strongly curved at the basal 2/5 (Fig. 17C), and thereafter gradually curved towards the tip (as opposed to apically slightly defl ected).See also the species accounts of A. chariedris and A. hadroprion.
Terminalia with furcal arms of spiculum gastrale apically strongly curved outwards.Tegmen similar in length to median lobe.Aedeagus (Fig. 17A) with median lobe length/width relation 6.-44-6.88 (N=1), basiventral margin strongly emarginate, lobe-like projections elongate and subtriangular, lateral margins slightly diverging in basal 5/6 of entire length, thereafter slightly rounded and gradually (triangularly) converging towards point-like, through very narrowly rounded apex.Median lobe in lateral view only slightly curved along extended mid region (second and third fourth of entire length), thereafter (apical fourth) dorsal margins converging towards apex in a slightly concave line, apically with very small, lobe-like, narrowly rounded projection that is neither expanded nor reclined.Internal sac with 2 weakly sclerotized, angulateuncinate rami, positioned in ostium and reclined mesobasally.
Terminalia with lamina of sternum VIII (Fig. 17B) widely triangular, all sides almost exactly equilateral, anterior margins projected, anterior half with 2 lateral, poorly defi ned transparent regions.Spermatheca (Fig. 17V) nearly V-shaped, ramus and collum angled at nearly 60°, each very short (reduced) and wide, resulting in a subcontiguous, uniformly swollen structure (corpus reduced), cornu strongly curved at basal 2/5, thereafter more gradually curved and continuously narrowed towards end.
Etymology.Named for the particularly prominent row of teeth on the protibia, with hadros signifying "well developed", and prion signifying "saw" (Brown 1956).Th e epithet is treated as a noun in apposition.
Natural history.Apotomoderes hadroprion is known to occur in low elevation habitats in two southeastern provinces of the Dominican Republic (Fig. 18): La Altagracia (Hoyo Azul) and San Pedro de Macoris (Juan Dolio).Th e host plant associations remain unknown.

Key to the species of Apotomoderes
1.

Phylogenetic analysis
Th e 12-taxon matrix (Table 1) yielded a single most parsimonious cladogram (Fig. 19) with a length of 33 steps, a consistency index (CI) of 75 and a retention index (RI) of 90 (Farris 1989).Th e character states and inferred optimizations are presented simultaneously in this section.Due to limited outgroup representation, the discussion of synapomorphies is restricted to the ingroup taxa.Individual consistency and retention indices (ci, ri) are provided for all characters that are not shown as unreversed synapomorphies.

Historical biogeography
Th e known Apotomoderes species records (Fig. 18) suggest the presence of two areas of endemism with multiple species occurrences, i.e. the southwestern Dominican Re-public (Pedernales) and the eastern Dominican Republic (La Altagracia and San Pedro de Macoris), assuming that the habitat conditions of the latter area are suffi ciently similar to sustain a wider distribution of (minimally) A. lateralis than herein reported.
A third area of endemism, Mona Island (Puerto Rico), is inhabited by a single species, A. sotomayorae (Fig. 18).

Discussion
According to the preferred cladogram (Fig. 19), the monophyly of Apotomoderes is well supported by a series of putative synapomorphies; viz.two-segmented labial palps (char.2), a postocular constriction on the head (char.5), a more expanded pronotum in males (char.8), the presence of setae in the dorsal subapical region of the aedeagus (char.17), and the presence of (ancestrally) outwardly directed, uncinate rami along the ostium of the aedeagus (char.18).Additional homoplasious traits that characterize the genus include the "open" metatibial corbel (char.12), the presence of a patch of suberect scales on the female elytral declivity (char.15; reversed in the A. sotomayorae-A.hadroprion clade), and the subcontiguous collum and ramus of the spermatheca (char.21).Th is redefi nition of Apotomoderes is more comprehensive and also slightly more inclusive than Schoenherr's (1834) and Lacordaire's (1863b) generic concepts Although Apotomoderes, as presently redefi ned, is a readily recognizable and monophyletic group, its immediate relatives and tribal placements remain uncertain.Th e inferred phylogeny (Fig. 19) places the genus closer to other naupactine genera, i.e.Artipus and Pantomorus Schoenherr, than to examined members of the Geonemini (= current tribal placement sensu Alonso-Zarazaga and Lyal 1999) and Eustylini.Such a placement is also indicated when using Anderson's (2002) key to the North American genera of Entiminae.Nevertheless, our knowledge of the phylogenetic limits of these tribes is judged too incomplete and the available character support too weak to warrant a tribal reallocation of Apotomoderes at this time (see also Marvaldi 1997;Franz 2010;Franz and Girón 2009).Likewise, the herein presented phylogeny should not be interpreted as a strong hypothesis about the relationships among the outgroup taxa.In his recent review, Pérez-Gelabert (2008) lists more than 70 entimine species for Hispaniola; nevertheless the actual number of species may be twice as high based on the diffi culty to identify dozens of species found on the island during a single fi eld trip in 2008 (Charles W. O'Brien, personal communication).It is therefore likely that the closest relative to Apotomoderes remains undescribed.On the other hand, the outgroup selection is considered adequate for polarizing character states within Apotomoderes (Nixon and Carpenter 1993).
Th e distributions of A. anodontos and A. menocrater in the southwestern Dominican Republic on one side, and the A. lateralis-A.hadroprion clade in the eastern Dominican Republic on the other side (Fig. 18), are congruent with the geological separation of a (1) southern ("Tiburón") and ( 2) central (central + northern) peninsula constituting the paleoislands that make up Hispaniola (Iturralde-Vinent and MacPhee 1999).Th e two regions are separated by the Cul-de-sac Lake Enriquillo Valley (corresponding to the Muertos Trough), a very dry desert habitat that divides many components of the fauna of Hispaniola (e.g.Rosen 1985;Liebherr 1992;Liebherr and Godwin 2004).However, the age of origin of Apotomoderes is likely (much) younger than the Miocene collision of the southern and central/northern peninsulas (cf.Heubeck and Mann 1991).In particular, the A. sotomayorae / A. lateralis-A.hadroprion split (Fig. 20) suggests that all components of this clade are of late Miocene or early Pliocene origin (i.e.5-7 mya), or possibly even younger, given that Mona Island rose above sea level at this stage (González et al. 1997).Th is would suggest that Apotomoderes originated and speciated some time in the Oligocene-Miocene on the southern Hispaniola peninsula, followed by two individual and unidirectional colonization events of the central/northern peninsula and of Mona Island, respectively.On the other hand, the presence of multiple, apparently sympatric species of Apotomoderes within each of the Hispaniolan areas of endemism cannot be explained with the scarce available information on habitat preferences and host plant associations.

Figure 1 .
Figure 1.Habitus of A. lateralis: A male, dorsal view B male, lateral view C male, ventral view D female, venter, ventral view.

Figure 2 .
Figure 2. Habitus of female type specimen of A. lateralis, located in the Naturhistoriska Riksmuseet, Stockholm, Sweden: A lateral view B head, thorax and prolegs, dorsal view C specimen label.Photographs taken by Johannes Bergsten.

Figure 4 .
Figure 4. Metendosternite and hind wing of A. lateralis: A metendosternite, posterior view B right hind wing.

Figure 7 .
Figure 7. Habitus of A. anodontos, male: A dorsal view B lateral view C frontal view, showing unarmed profemora.

Figure 9 .
Figure 9. Habitus of A. menocrater: A female, dorsal view B female, lateral view C male, head and pronotum, dorsal view, showing subfoveate punctures D female, declivity with patch of suberect setae, lateral view.

Figure 11 .
Figure 11.Habitus of A. sotomayorae, male: A dorsal view B frontal view C lateral view.

Figure 13 .
Figure 13.Habitus of A. chariedris, male: A dorsal view, showing strongly expanded pronotum B head, dorsal view C frontal view, showing large profemoral tooth and toothed, triangularly projected protibial ridge.

Figure 18 .
Figure 18.Reported occurrences and posited areas of endemism for the species of Apotomoderes.

Figure 19 .
Figure 19.Phylogeny of the species of Apotomoderes and select outgroup taxa, according to the single most parsimonious cladogram (L=33, CI= 75, CI=90).Character 15 is mapped under ACCTRAN optimization, whereas character 16 is mapped under DELTRAN optimization.All other characters have unambiguous optimizations.Black rectangles represent single, non-homoplasious character state transformations, and white rectangles represent multiple, homoplasious character state transformations.Th e numbers above and below each rectangle correspond to character numbers and states, respectively; the numbers displayed at the left end of each branch represent Bremer support values.

Table 1 .
Agnarsson and Miller 2008)ladistic analysis of the species of Apotomoderes, with selected outgroup taxa.All multistate characters are coded as additive; "-" represents inapplicable character states (see also text).Agnarsson and Miller 2008), therefore convergently present in the two examined species of Lachnopus Schoenherr and in the in the A. fl oridanus-Apotomoderes clade, with a reversal in the A. lateralis-A.hadroprionclade (ci=33; ri=50).17.Male terminalia, presence of setae in dorsal subapical region of median lobe: (0) absent;(1) present.Synapomorphy for the species of Apotomoderes.18. Male terminalia, orientation of rami along ostium of median lobe: (0) rami sub-Coded as inapplicable in taxa where the collum and ramus are clearly separated (see character 21).Synapomorphy for the A. chariedris-A.hadroprion clade.