BULLETIN DE L’INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE ENTOMOLOGIE, 79: 59-72, 2009 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN ENTOMOLOGIE, 79: 59-72, 2009

Seven new Neotropical species of the genus Ardistomis PUTZEYS (Coleoptera: Carabidae: : ): notes about classification and a checklist of species names of that genus

By Pavel VALDÉS R.

Abstract (1977) proposed that Semiardistomis be ranked as a genus, in which he included Ardistomiellus (that name The ardistomine complex includes the genera Ardistomis Putzeys, becoming thereby a junior subjective synonym of 1846, Semiardistomis Kult, 1950 and Aspidoglossa Putzeys, 1846. Semiardistomis). NICHOLS (1988a, 1988b), following Seven new species of the genus Ardistomis Putzeys are described and illustrated: A. drumonti n. sp. from French Guiana, A. minutus n. sp. Whitehead’s proposals, formally ranked Semiardistomis from Bolivia, A. samyni n. sp. from Honduras, A. dostali n. sp. from as a genus, and included therein the species originally Costa Rica and A. onorei n. sp., A. bulirschi n. sp., and A. vergelae assigned to Ardistomiellus; members of the genus n. sp. from Ecuador. Two monophyletic groups are defined: group Ardistomis inhabit West Indies where arranged here in muelleri and group ovatus, both with montane flightless species. A checklist includes valid names, new synonymies and distributional 3 “major” lineages. records for Ardistomis. In this paper I accept the Whitehead/ Nichols proposals for classification of the species of the Key words: Coleoptera, Carabidae, Clivinini, Ardistomina, new ardistomine complex at the supraspecific level. For species, Neotropic Ardistomis, I define just those monophyletic groups that include the new species described with a clear set of autapomorphies. The complete revision of the genus Introduction Ardistomis is under preparation. The major focus of this paper, however, is description and illustration of This represents the first paper in a series of studies diagnostic features of new species of Ardistomis from devoted to elucidation of the taxonomic structure of the Central and northern South America. Included also is ardistomine complex, which comprises three genera: a checklist of the specific names of the species of this Ardistomis PUTZEYS, 1846; Semiardistomis KULT, genus, bringing up to date the list by LORENZ (2005). 1950; and Aspidoglossa PUTZEYS, 1846. The group is currently placed in the tribe Clivinini. Previously it was recognized as Ardistomides (PUTZEYS, 1866), Material and Methods Ardistomina (CSIKI, 1927) or Ardistomini (JEANNEL, 1946). This complex is confined to the Western More than 2200 specimens of the ardistomine Hemisphere, ranging through the Neotropical Region, complex were examined, representing 103 named including the West Indies northward to southeastern species. Most specimens were borrowed, but some are United States in the Nearctic Region. represented in my personal collection (PVCC). Listed PUTZEYS (1866) arranged the species of Ardistomis below, with acronyms used in the text, are names and in three numbered groups. His first group was addresses of the lending institutions (names of curators nominotypical; in his second and third groups he placed in parentheses) and owners of private collections species now included in the genus Semiardistomis ADVA = Alexander Dostal Collection (includes Kult KULT 1950. The latter author (1950) “provisionally” Collection), Vienna, Austria. placed the species in 13 groups, distributed among BMNH = The Natural History Museum. London, UK. three subgenera: Ardistomis s. str. with 4 groups, (Max Barclay and Christine Taylor) Semiardistomis with 8 groups from which 3 are true CMN = Canadian Museum of Nature, Aylmer, Ardistomis and one group as Ardistomiellus. WHITEHEAD Québec, Canada (R.S. Anderson, F. Génier). 60 P.R. VALDÉS

HECO = Hope Entomological Collection, Oxford usually crowned by a basal crest (bc); (2) a median University, UK. (James Hogan.) portion (Mp), on its dorsal side the ostium (o) can be IES = Instituto de Ecología y Sistemática, opened from different levels of this portion or from Ciudad de la Habana, Cuba. (A.Lozada) the basal bulb to base of apical portion, and ventrally IRSNB = Institut Royal des Sciences Naturelles a median ridge (mr) along its axis can be developed de Belgique, Bruxelles, Belgique. (A. or not; (3) the apical portion (Ap). Illustrations of the Drumont). phallus are oriented to the left (sinistral) or to the right MNHP = Muséum National d’Histoire Naturelle, side (dextral). Paris, France. (T. Deuve and A. The surface of the endophallus is either densely or Tagavian). sparsely covered by an armature of different kinds of MNHNCu = Museo Nacional de Historia Natural de spines, hairs and microtrichia in specific areas. Inside Cuba, Ciudad de la Habana, Cuba (E. the endophallus a basal sclerite (bs) is present in males Gutiérrez) of some species. PBPC = Petr Bulirsch Collection. Prague, Czech One of the parameres (P1) is larger than the other Republic. one and has three apophyses: one lateral, one basal and PMGT = Pier Mauro Giacchino Collection, Torino, one ventral. The second paramere (P2) is slender and Italy. acute apically. Both are setose apically or glabrous, the UASM = Strickland Museum, University of Alberta, number of setae various. Edmonton, Alberta, Canada. (G. E. Ball and D. Shpeley). Taxonomic treatment Dissections of adults were made using standard techniques. The genitalia were preserved in glycerine Genus Ardistomis PUTZEYS, 1846 in microvials pinned beneath the specimens. Mouthparts were glued on small cards pinned beneath TYPE SPECIES: A. fasciolatus PUTZEYS, 1846 the specimens. Observations were made under a stereobinocular microscope (50X) and a compound RECOGNITION: Most species with a submarginal band of microscope (100X). All line drawings were made microsculpture on proepisternum. Mandibles elongate, from digital microphotographs using Corel Draw 13X terebral ridges almost straight. Anterior margin of software. clypeus with lateral lobes indistinct. Antennomere The following measurements were made using an 2 subequal in length to antennomere 3. Pronotal ocular micrometer: head length (HL): linear distance disc ovate to subglobose (Figs 1, 9), proepipleura from apical margin of clypeus to posterior margin of visible from above or not. Elytral striae complete and right eye; length of pronotum (PL): linear distance from inpunctate. Protibia ventral surface with only one seta anterior to posterior margin along the midline; pronotal along midline. Abdominal sternum VII with a lateral width (PW): greatest linear transverse distance; elytral projection that fits into preapical elytral plica. Union length (EL): linear distance from basal ridge to apex of abdominal sterna III and IV obliterated at middle. along the suture; elytral width (EW): greatest linear Abdominal sterna with or without accessory setae. transverse distance across both elytra. The standardized Phallus (Figs 2, 8, 21) with a defined basal bulb which body length (SBL) is the sum of the lengths of head, produces distally only one lamella with a median and pronotum and elytra. an apical portions, the ostium is opened dorsally; Most terms used for structural features are found in parameres well differentiated each other, with reduced previous works on Carabidae: ALLEN & BALL (1980) number of setae. Gonocoxae unsegmented, elongate for adult microsculpture; ACORN & BALL (1991) for with a distinctly differentiated wide basal portion. adult mandibles; LIEBHERR & WILL (1998) for female genitalia. MATERIAL EXAMINED: Most currently recognized species The phallus in genus Ardistomis (see Figs. 2, 8, 22) (see checklist, below) except: A. alticola DARLINGTON, consists of three parts: (1) a basal bulb (Bb) defined A. annona PUTZEYS, A. hispaniolensis Nichols at its distal end by the basal apophysis (ba) where (manuscript name), A. profundestriatus PUTZEYS, A. the parameres are attached, which includes the basal rufoclarus DARLINGTON, A. tropicalis PUTZEYS and A. opening (bo), its proximal end is usually developed unicolor PUTZEYS. in a canaliculated plate named basal projection (bp), New species of the genus Ardistomis PUTZEYS 61

Group fasciolatus clypeus emarginate, pronotal disc subglobose and elytra This group is composed of different monophyletic units with humeri rounded and preapical elytral spot smaller. united by having labial palpomere 3 subequal in length to palpomere 2, abdominal sterna without accessory DESCRIPTION: Habitus as in Fig. 1. For measurements see setae and endophallus usually with abundant and varied Table I. armature. (Figs. 2, 3) Color. Body color mostly piceous; antennae testaceous, legs and mouthparts ferruginous; pronotum and elytra with slight greenish luster, elytra preapically Ardistomis drumonti n. sp. spotted, spots testaceous. (Figs 1, 2) Microsculpture. Frontoclypeus with most of sur- face smooth, mesh pattern isodiametric laterally; TYPE MATERIAL: HOLOTYPE male labelled: “Leprieuri supraantennal lobes smooth; vertex with mesh pattern Buquet Cayenne (C. Chd.)” [handwritten on green isodiametric; gena with mesh pattern isodiametric; paper]; “Soc. Ent. Belg. Coll. PUTZEYS” [printed]; gula with mesh pattern transverse. Mandibles smooth; “Syntype” [printed]; “Paralectotype Dyschirius submentum and mentum with mesh pattern isodiametric. leprieuri Chd. Des. S. W. Nichols 1984” [handwritten]. Pronotal disc in anterior two thirds with mesh pattern Holotype deposited in IRSNB. isodiametric, transverse in posterior third; proepisternum with mesh pattern isodiametric, submarginal band of ETYMOLOGY: Specific epithet a Latinized noun, microsculpture present; prosternum with mesh pattern masculine genitive case, based on the surname of Dr. transverse. Metasternum with mesh pattern transverse. Alain Drumont, curator of the collection of the Elytra with shallow longitudinal microlines; epipleuron Royal Belgian Institute of Natural Sciences, to whom with longitudinal microlines. Abdominal sterna with this species is dedicated. mesh pattern transverse. Chaetotaxy. Anterior marginal setae on pronotal disc RECOGNITION: This specimen was misidentified since closer to anterior angle than to posterior setae. Elytral PUTZEYS (1846) as Ardistomis leprieuri (CHD.) (Fig. disc with 5 setae in interval 3. Abdominal sternum 3). From other members of this group it is readily VII without accessory setae near base; preapical setae distinguished by less prominent eyes, anterior margin of equidistant from each other.

Figs 1-3. – 1. Ardistomis drumonti n. sp., habitus (dorsal view), scale bar 1 mm. 2—3 Male genitalia (a) ventral view, (b) right lateral view, scale bar 0,1 mm. 2. Ardistomis drumonti n. sp. Ap, apical portion; ba, basal apophysis; Bb, basal bulb; bc, basal crest; bo, basal opening; bp, basal projection; Mp, median portion; mr, median ridge; o, ostium; P, parameres. 3. Ardistomis leprieuri (CHAUDOIR). 62 P.R. VALDÉS

Head. Clypeus with anterior margin emarginate. by smaller size and subglobose pronotal disc. Vertex with transverse groove between posterior supraorbital setae. Eyes normal. Antennomeres 4–10 DESCRIPTION: Habitus as in Fig. 4. For measurements see about 1.6 X longer than wide. Table I. Mouthparts. Labrum with anterior margin projected Color. Body ferruginous; elytra unspotted; antennae, medially. Mandibles with basal portion about 4/5 length mouthparts and legs testaceous. of terebra. Labium with palpomere 3 subequal in length Microsculpture. Frontoclypeus with most of to palpomere 2. surface smooth; supraantennal lobes smooth; vertex Thorax. Pronotum subglobose; anterior transverse with mesh pattern isodiametric; gena with transverse and median longitudinal impressions distinct. microlines; gula with transverse microlines. Proepipleura visible from above. Metasternum posteriad Mandibles smooth; submentum and mentum with mesocoxa longer than metacoxa. mesh pattern isodiametric. Pronotal disc with mesh Elytra. Oval, humeri somewhat rounded; striae pattern transverse; proepisternum with mesh pattern distinct throughout their length; intervals convex. isodiametric, submarginal band of microsculpture Metathoracic wings: Fully developed. present; prosternum with mesh pattern transverse. Male genitalia: (Fig. 2) Phallus dextral, slender Metasternum with mesh pattern transverse. Abdominal in frontal view (Fig. 2a), curved in lateral view (Fig. sterna with transverse microlines. Elytra smooth. 2b); basal bulb distinct, basal orifice (bo) oval; basal Chaetotaxy. Anterior marginal setae on pronotal disc projection (bp) developed, sclerotized, apical margin closer to anterior angle than to posterior setae. Elytral rounded; basal crest (bc) present; basal apophysis (ba) disc with 5 setae in interval 3. Abdominal sternum VII moderate, sclerotized; ostium (o) opened from half of with ambulatory setae near base, one on each side; ventral basal portion to base of apical portion; median preapical setae equidistant each others. ridge (mr) pronounced; apical portion (ap) plate like, Head. Clypeus with anterior margin slightly arcuate. moderate, projected rightward, margin rounded, micro Vertex with transverse groove between posterior canaliculated. Endophallus normal in length; armature supraorbital setae. Supraantennal lobes with basal on apical area covered with microtrichia; basal sclerite median depressions. Eyes prominent, hemispherical . absent. Paramere 1 (P1) about 2 times longer than wide, Antennomeres 4–10 about 1.7 X longer than wide. asetose; lateral apophysis normal, blunt; basal apophysis Mouthparts. Labrum with anterior margin slightly acute; ventral apophysis developed. Paramere 2 (P2) projected medially. Mandibles with basal portion about slender, 7/10 length of paramere 1, apex acute, 3 apical 7/10 length of terebra. Labium with palpomere 3 longer setae. than palpomere 2. Thorax. Pronotum subglobose; anterior transverse impression distinctly impressed and median longitudinal Group venustulus impression distinct. Proepipleura hardly visible from This group includes several monophyletic units that above. Metasternum posteriad mesocoxa subequal in share labial palpomere 3 longer than palpomere 2, length to metacoxa. abdominal sterna with accessory setae and endophallus Elytra.Oval, humeri distinct; striae distinct small, usually with reduced armature. throughout their length; intervals convex. Metathoracic wings. Fully developed Female genitalia. Gonocoxa (Gc) (Fig. 5), slender; Ardistomis minutus n. sp. apical portion 1/2 of total gonocoxite length, folded (Figs. 4, 5) medially, two apical setae; basal portion ends in a notable perpendicular prolongation with two small TYPE MATERIAL: HOLOTYPE female labeled: “Bolivia. 24- seta apically. Laterotergite (Lt) rectangular in outline. 27. IX. 1996, Dpto Santa Cruz, Perseverancia, leg. W. Spermathecal duct moderate in length, reservoir Rossi” [printed]. Holotype deposited in PBPC. distinct; spermathecal gland insert in half portion of spermathecal duct. ETYMOLOGY: Specific epithet a Latinized masculine adjective, referring to the small size of this species. Group muelleri RECOGNITION: This species is close to Ardistomis This is a monophyletic group including species that venustulus PUTZEYS (Fig. 6) from which it is recognized share the following autapomorphies: abdominal New species of the genus Ardistomis PUTZEYS 63

Figs 4-6. – 4. Ardistomis minutus n. sp., habitus (dorsal view), scale bar 1 mm. 5—6 Female genitalia (dorsal view), scale bar 0,1 mm. Gc, gonocoxae; Lt, lateroterguites. 5. Ardistomis minutus n. sp. 6. Ardistomis venustulus.

sternum VII with an irregular row of accessory setae Key for identification of adults of the muelleri group near base, and phallus elongate, tubular, with reduced from the Neotropical mainland. and blunt apical portion, reduced endophallus with few sparse microtrichia and distinctly developed 1 Eyes normal, elytral humeri slightly rounded, basal sclerite. Other characters like color fuscous to metathoracic wings full …………………………… 2 ferruginous, elytra unspotted or labial palpomere 3 - Eyes small, humeri rounded, metathoracic wings longer than palpomere 2 suggest close relationship vestigial ………………………………………………3 with members of the venustulus clade. KULT (1950) 2 Total length more than 5 mm, anterior margin of described for A. muelleri (Figs 17-19) “proepisterna clypeus arcuate. Costa Rica …………….…………… without submarginal furrow”. I examined the Kult types …………………..…………………… A. dostali n. sp. of muelleri (2 paratypes teneral), as well 6 additional - Total length less than 4,6 mm. anterior margin of specimens from UASM collection. All exhibit a band clypeus emarginate. Ecuador ……… ………………… of proepisternal microsculpture, but it is indistinct. The ……………………………………. A. bulirschi n. sp. members of the muelleri group that inhabit montane wet 3 Eyes hemispherical, temples not developed. Ecuador forest habitats show a tendency to metathoracic wing ………………….. ……………………A. onorei n. sp. and eye reduction. Some West Indian montane flightless - Eyes flat, temples developed overall behind eyes …… species probably share a common ancestor with this ………………………………………………………4 mainland group. 4 Total length more than 4,5 mm. Pronotal disc ovate. Geographical distribution: see Fig. 20 Mexico …….…………………………A. muelleri KULT - Total length less than 3,5 mm. Pronotal disc globose. Honduras ………..……………………A samyni n. sp. 64 P.R. VALDÉS

Figs 7-19. – 7, 9, 11, 14, 17. Habitus (dorsal view), scale bar 1 mm. 8, 10, 12, 15, 18. Male genitalia (a) ventral view, (b) right lateral view, scale bar 0,1 mm. 13, 16, 19. Female genitalia (dorsal view), scale bar 0,1 mm. Gc, gonocoxae; Lt, lateroterguites 7—8 Ardistomis dostali n. sp. Ap, apical portion; ba, basal apophysis; bc, basal crest; bo, basal opening; bp, basal projection; bs, basal sclerite; o, ostium; P, parameres. 9—10. Ardistomis samyni n. sp. 11—13. Ardistomis bulirschi n. sp. 14—16. Ardistomis onorei n. sp. 17—19. Ardistomis muelleri KULT. New species of the genus Ardistomis PUTZEYS 65

Ardistomis dostali n. sp. medially. Mandibles with basal portion about 3/5 (Figs. 7,8) length of terebra. Labium with palpomere 3 longer than palpomere 2. TYPE MATERIAL. HOLOTYPE male labelled: “Costa Rica, Thorax. Pronotum subovate; anterior transverse and Cartago, Tapanti N P, Road Rio Orosi, 1200- 1500m median longitudinal impressions distinct. Proepipleura 9°42- 44’N, 83°46’W, 13- 15. 5. 2006, leg. Barries, visible from above. Metasternum posteriad mesocoxa Cate & Nagy” [printed]. PARATYPE (1) labeled: “6/ longer than metacoxa. Costa Rica, Puntarenas, Est. Tres Colinas, 1800- Elytra. Elongate- oval, humeri rounded; striae 2000m, La Amistad N. P., 21- 23. 4. 2005, 9°07’11»N, distinct throughout their length; intervals convex. 83°04’06»W, leg. Barries, Cate & Nagy» [printed]. Metathoracic wings. Fully developed. Holotype deposited in INBIO Collection: Apdo. Male genitalia. (Fig. 8) Phallus dextral, notably Postal 22-3100 Sto. Domingo de Heredia, Costa Rica; elongate, somewhat tubular, slender in frontal view Paratypes deposited in ADVA. (Fig. 8a), slightly curved in lateral view (Fig. 8b); basal bulb indistinctly defined from medial part, basal orifice ETYMOLOGY: Specific epithet a Latinized noun, (bo) fusiform; basal projection (bp) distinct, sclerotized, masculine genitive case, based on the surname of Dr. apical margin blunt; basal crest developed; basal Alexander Dostal from Vienna Austria, specialist in apophysis (ba) moderate; ostium (o) wide, opened from Carabidae, to whom this species is dedicated. basal beginning of ventral median portion to base of apical portion; median ridge absent; apical portion (ap) RECOGNITION: See the key to species of the muelleri short, projected leftward, margin blunt. Endophallus group, above. short; armature consist on one small apical area covered with micro setae; basal sclerite (bs) present, very DESCRIPTION: Habitus as in Fig. 7. For measurements see elongate, almost the same length as median portion, Table I. length of apical prolongations about 2/5 of total sclerite Color. Body fuscous; elytra bicolored, posterior length. Paramere 1 (P1) about 3,2 times longer than third ferruginous, unspotted; antennae, mouthparts and wide, asetose; lateral apophysis acute; basal apophysis legs ferruginous. blunt; ventral apophysis normal. Paramere 2 (P2) Microsculpture. Frontoclypeus with most of surface slender, about 1/2 length of paramere 1, apex acute, one smooth; supraantennal lobes smooth; vertex with mesh apical seta. pattern transverse; gena with mesh pattern transverse; gula with transverse microlines. Mandibles smooth; submentum and mentum with mesh pattern isodiametric. Ardistomis samyni n. sp. Pronotal disc with faith mesh pattern slightly (Figs. 9, 10) transverse, microlines very shallow; proepisternum with mesh pattern isodiametric, submarginal band of TYPE MATERIAL: HOLOTYPE male labelled: “Honduras, microsculpture present; prosternum with mesh pattern Olancho, La Muralla N. Pk., 14 km. N. La Union, transverse. Metasternum with mesh pattern transverse. 1450m, 17. VIII. 1994, S. Peck, 94- 39, wet mont. for. Elytra with microlines longitudinally oriented. litter” [printed]. Holotype deposited in CMN. Abdominal sterna with mesh pattern transverse. Chaetotaxy. Anterior marginal setae on pronotal ETYMOLOGY: Specific epithet a Latinized noun, disc equidistant between anterior angle and posterior masculine genitive case, based on the surname of Dr. setae. Elytral disc with 5 setae in interval 3. Abdominal Yves Samyn, specialist on holothurian echinoderms, sternum VII with ambulatory setae near base, 5 on each at the Royal Belgian Institute of Natural Sciences to side in an irregular row; inner pair of preapical setae whom this species is dedicated. separated 2 times distance between inner and outer seta. Head. Clypeus with anterior margin arcuate. RECOGNITION: See the key to the species of the muelleri Vertex with deep transverse groove between posterior group, above. supraorbital setae. Supraantennal lobes with basal median depressions. Eyes normal. Antennomere 2 DESCRIPTION: Habitus as in Fig. 9. For measurements see subequal in length to antennomere 3; antennomeres Table I. 4–10 about 1.6 X longer than wide. Color. Body fuscous; elytra bicolored, posterior Mouthparts. Labrum with anterior margin projected third ferruginous, unspotted; antennae testaceous, 66 P.R. VALDÉS mouthparts and legs ferruginous. Ardistomis bulirschi n. sp. Microsculpture. Frontoclypeus with most of (Figs. 11-13) surface smooth; supraantennal lobes smooth; vertex with mesh pattern isodiametric; gena with mesh TYPE MATERIAL: HOLOTYPE male labelled: “Ecuador, pattern transverse; gula with transverse microlines. Pichincha, Alluriquin, Rio Toachi, 9- 11.III.1999. lgt. Mandibles smooth; submentum and mentum with Šebela, Prouza, Bašta” [printed]. PARATYPE (1), female mesh pattern isodiametric. Pronotal disc with mesh labeled: “ Ecuador, Guayas, Milagro, 14- 17.III.2000, pattern transverse, microlines shallow; proepisternum lgt. Šebela, Prouza .” [printed] Holotype deposited in with mesh pattern isodiametric, submarginal band of PBPC, Paratype deposited in PVCC microsculpture present; prosternum with mesh pattern transverse. Metasternum with mesh pattern transverse. ETYMOLOGY: Specific epithet a Latinized noun, Elytra with longitudinal microlines. Abdominal sterna masculine genitive case, based on the surname of Dr. with mesh pattern transverse. Petr Bulirsch from Prague, Czech Republic, specialist Chaetotaxy. Anterior marginal setae on pronotal disc on Carabidae, to whom this species is dedicated. equidistant between anterior angle and posterior setae. Elytral disc with 5 setae in interval 3. Abdominal sternum RECOGNITION: See the key to the species of the muelleri VII with ambulatory setae near base, 5 on each side in an group, above. irregular row; inner pair of preapical setae separated 1.5 times distance between inner and outer seta. DESCRIPTION: Habitus as in Fig. 11. For measurements Head. Clypeus with anterior margin arcuate. see Table I. Vertex with deep transverse groove between posterior Color. Body fuscous; elytra bicolored, posterior supraorbital setae. Supraantennal lobes with basal third ferruginous, unspotted;; antennae testaceous, median depressions. Eyes small, temple moderate. mouthparts and legs ferruginous. Antennomere 2 subequal in length to antennomere 3; Microsculpture. Frontoclypeus with surface smooth; antennomeres 4–10 about 1.6 X longer than wide. supraantennal lobes smooth; vertex with mesh pattern Mouthparts. Labrum with anterior margin projected isodiametric; gena with mesh pattern isodiametric; medially. Mandibles with basal portion about 3/5 gula with transverse microlines. Mandibles smooth; length of terebra. Labium with palpomere 3 longer than submentum and mentum with mesh pattern isodiametric. palpomere 2. Pronotal disc with faith mesh pattern transverse; Thorax. Pronotum subglobose; anterior transverse proepisternum with mesh pattern isodiametric, and median longitudinal impressions distinct. submarginal band of microsculpture present; prosternum Proepipleura visible from above. Metasternum with mesh pattern transverse. Metasternum with mesh posteriad mesocoxa subequal in length to metacoxa. pattern transverse. Abdominal sterna with mesh pattern Elytra. Oval, humeri rounded; striae distinct transverse. Elytra with longitudinal microlines. throughout their length; intervals convex. Chaetotaxy. Anterior marginal setae on pronotal disc Metathoracic wings. Vestigial. closer to anterior angle than to posterior setae. Elytral Male genitalia. (Fig. 10) Phallus dextral, elongate, disc with 5 setae in interval 3. Abdominal sternum VII somewhat tubular, slender in frontal view (Fig. 10a), with ambulatory setae near base, 6 on each side in an curved in lateral view (Fig. 10b); basal bulb defined irregular row; preapical setae equidistant from each from medial part, basal orifice fusiform; basal projection other. distinct, sclerotized, apical margin blunt; basal crest Head. Clypeus with anterior margin emarginate. developed; basal apophysis moderate; ostium wide, Vertex with deep transverse groove between posterior opened from 1/10 of ventral median portion to base supraorbital setae. Supraantennal lobes with basal of apical portion; median ridge absent; apical portion median depressions pronounced. Eyes normal. short, projected leftward, margin blunt. Endophallus Antennomere 2 subequal in length to antennomere 3; short; armature consist on one small apical area covered antennomeres 4–10 about 1.6 X longer than wide. with micro setae; basal sclerite present, elongate, length Mouthparts. Labrum with anterior margin projected of apical prolongations about 1/2 of total sclerite length. medially. Mandibles with basal portion about 3/5 Paramere1 about 2.1 times longer than wide, asetose; length of terebra. Labium with palpomere 3 longer than lateral apophysis acute; basal apophysis blunt; ventral palpomere 2. apophysis normal. Paramere 2 slender, about 1/2 length Thorax. Pronotum subovate; anterior transverse and of paramere1, apex acute, one visible apical seta. median longitudinal impressions distinct. New species of the genus Ardistomis PUTZEYS 67

Proepipleura visible from above. Metasternum Color. Body fuscous; elytra ferruginous, unspotted; posteriad mesocoxa longer than metacoxa. antennae testaceous, mouthparts and legs ferruginous. Elytra. Elongate- oval, humeri rounded; striae Microsculpture. Frontoclypeus with most of surface distinct throughout their length; intervals convex. smooth; supraantennal lobes smooth; vertex with mesh Metathoracic wings. Fully developed. pattern isodiametric; gena with mesh pattern transverse; Male genitalia. (Fig. 12) Phallus dextral, elongate, gula with mesh pattern transverse. Mandibles somewhat tubular, slender in frontal view (Fig. 12a), smooth; submentum and mentum with mesh pattern curved in lateral view (Figs. 12b); basal bulb poorly isodiametric. Pronotal disc with mesh pattern transverse; defined from medial part, basal orifice fusiform; proepisternum with mesh pattern isodiametric, smooth basal projection distinct, apical margin blunt; basal near prosternum, submarginal band of microsculpture crest developed; basal apophysis moderate; ostium present; prosternum with mesh pattern transverse. wide, opened from begin of ventral basal portion to Metasternum with mesh pattern transverse. Abdominal base of apical portion; median ridge absent; apical sterna with mesh pattern transverse. Elytra with portion plate like, projected leftward, margin rounded. longitudinal microlines. Endophallus short; armature consist on one small apical Chaetotaxy. Anterior marginal setae on pronotal disc area covered with micro setae; basal sclerite about 2/5 closer to anterior angle than to posterior setae. Elytral length of median portion, length of apical prolongations disc with 5 setae in interval 3. Abdominal sternum VII about 1/2 of total sclerite length, almost without basal with ambulatory setae near base, 5 on each side in an prolongations. Paramere1 about 2,5 times longer than irregular row; preapical setae equidistant from each wide, asetose; lateral apophysis acute; basal apophysis other. blunt; ventral apophysis normal. Paramere 2 slender, Head. Clypeus with anterior margin arcuate, slightly about 2/5 length of paramere1, apex acute, one visible emarginate. Vertex with deep transverse groove between apical seta. posterior supraorbital setae. Supraantennal lobes with Female genitalia. (Fig. 13) Gonocoxa (Gc) slender; basal median depressions pronounced. Eyes moderately apical portion elongate and curved with two apical setae; reduced, temple short. Antennomere 2 subequal in basal portion 2/5 of total gonocoxite length, 2 small seta length to antennomere 3; antennomeres 4–10 about 1.6 in inner curvature. Laterotergite (Lt) wide, rectangular X longer than wide. in outline. Spermathecal duct long, reservoir distinct; Mouthparts. Labrum with anterior margin projected spermathecal gland indistinct. medially. Mandibles with basal portion about 3/5 length of terebra. Labium with palpomere 3 longer than palpomere 2. Ardistomis onorei n. sp. Thorax. Pronotum subovate; anterior transverse and (Figs. 14-16) median longitudinal impressions distinct. Proepipleura visible from above. Metasternum posteriad mesocoxa TYPE MATERIAL: HOLOTYPE male labelled: “Ecuador, longer than metacoxa. Napo, via Jondachi- Loreto km 59, ex cave m 700, Elytra. Moderately oval, humeri rounded; striae 13.VIII.2006, G. Onore leg.” [printed] PARATYPE (1), distinct throughout their length; intervals convex. female labeled as the holotype. Holotype deposited in Metathoracic wings. Vestigial. PMGT, Paratype deposited in PBPC. Male genitalia. (Fig. 15) Phallus dextral, elongate, somewhat tubular, slender in frontal view (Fig. 15a), ETYMOLOGY: Specific epithet a Latinized noun, curved in lateral view (Fig. 15b); basal bulb poorly masculine genitive case, based on the surname of Dr. defined from medial part, basal orifice fusiform; basal Giovanni Onore, from Quito Ecuador, agronomist, projection distinct, apical margin blunt; basal crest entomologist and Catholic missionary of the developed; basal apophysis moderate; ostium wide, Marianistas Congregation, collector of this species, opened from 1/10 of ventral median portion to base of which is dedicated to him apical portion; median ridge absent; apical portion plate like, projected leftward, margin rounded. Endophallus RECOGNITION: See the key for the muelleri species group, short; armature of one small apical area covered with above. micro setae; basal sclerite about 4/10 length of median portion, length of apical prolongations about 1/2 of total DESCRIPTION: Habitus as in Fig. 14. For measurements sclerite length, almost without basal prolongations. see Table I. Paramere1 about 3 times longer than wide, asetose; 68 P.R. VALDÉS

Fig. 20. – Map of Middle America and northern part of South America showing positions of known localities for the species of the muelleri group.

lateral apophysis acute; basal apophysis blunt; ventral Ardistomis. Because the ovatus group is closely related apophysis normal. Paramere 2 slender, about 4/10 to members of the venustulus clade, the two sharing length of paramere1, apex acute, one apical seta. many apomorphic features, I prefer to consider these Female genitalia. (Fig.16) Gonocoxa (Gc), slender; groups as congeneric. Thus, I consider absence of the apical portion elongate and curved with two apical proepisternal submarginal band of microsculpture from setae; basal portion 2/5 of total gonocoxite length, the ovatus group as a loss rather than retention of a 5 small seta in inner curvature. Laterotergite (Lt) plesiomorphic state. wide, subquadrate in outline. Spermathecal duct long, reservoir distinct; spermathecal gland inserted in basal portion of spermathecal duct. Ardistomis vergelae n. sp. (Figs. 21-23)

Group ovatus TYPE MATERIAL: HOLOTYPE male labelled “Ecuador: Napo, Antisana Reserve road, 4.2kmSW. Cosanga, Ardistomis ovatus PUTZEYS forms, together with 2150m, 00°37´19´´S, 77°50´01´´W, 5.XI.1999-229e, A. vergelae n. sp. a monophyletic unit defined by R.Anderson, montane evergreen forest litter” [printed]. the absence of proepisternal submarginal band of 1st PARATYPE (teneral) labeled “Ecuador: Napo, 2.5km microsculpture, pronotal disc transversely ovate, W. Cosanga, 2150m 00°35´24´´S, 77°53´19´´W, proepipleura not visible from above and phallus with 5.XI.1999- 228a, R.Anderson, montane evergreen forest ostium open from about 1/3 to 2/3 of ventral median litter” [printed]. 2nd PARATYPE male and female labeled portion. A. rotundipennis PUTZEYS probably shares a “Ecuador. Napo, Sierra Azul, 15.0 km. W. Cosanga, common ancestor with the former species. All three 2350 m, 00°40´55´´S 77°56´09´´W, 5.XI.1999- 227b, species have eyes and metathoracic wings reduced. R. Anderson, montane evergreen forest litter” [printed]. A. ovatus and A. rotundipennis were included The holotype and one paratype are deposited in CMN, in Semiardistomis by KULT (1950) and BOUSQUET one paratype is in PBPC, and one paratype is in PVCC. (2006). The difference in classification is based on interpretation of one character state: the absence ETYMOLOGY: Specific epithet a Latinized feminine from the ovatus group species of the proepisternal noun, genitive case, based on the surname of my wife, submarginal band of microsculpture. When present, Marydena Vergel, to whom this species is dedicated. this feature is regarded as apomorphic for the genus New species of the genus Ardistomis PUTZEYS 69

RECOGNITION: This species is readily distinguished transverse. from its relatives, A. ovatus PUTZEYS (Fig. 23) and A. Chaetotaxy. Anterior marginal setae on pronotal rotundipennis PUTZEYS by its piceous color with bluish disc equidistant between anterior angle and posterior luster. setae. Elytral disc with 5 setae in interval 3. Abdominal sternum VII with ambulatory setae near base, one on DESCRIPTION: Habitus as in Fig. 21. For measurements each side; preapical setae equidistant each others. see Table I. Head. Clypeus with anterior margin straight. Color. Body piceous with a bluish luster; elytra Vertex without transverse groove between posterior unspotted; antennae testaceous, legs and mouthparts supraorbital setae. Supraantennal lobes with basal ferruginous. median depressions. Eyes reduced, temple moderate. Microsculpture. Frontoclypeus surface with mesh Antennomere 2 subequal in length to antennomere 3; pattern isodiametric; supraantennal lobes mostly antennomeres 4–10 about 1.5 X longer than wide. smooth; vertex with mesh pattern isodiametric; gena Mouthparts. Labrum with anterior margin projected with mesh pattern isodiametric; gula with transverse medially. Mandibles with basal portion about 3/5 microlines. Mandibles smooth; submentum and mentum length of terebra. Labium with palpomere 3 longer than with mesh pattern isodiametric. Pronotal disc with mesh palpomere 2. pattern isodiametric, microlines shallowly impressed; Thorax. Pronotum transversely ovate; anterior proepisternum with mesh pattern isodiametric, transverse impression moderate and median longitudinal submarginal band of microsculpture absent; prosternum impression distinct. Proepipleura not visible from with mesh pattern transverse. Metasternum smooth. above. Metasternum posteriad mesocoxa shorter than Elytra smooth. Abdominal sterna with mesh pattern metacoxa.

Figs 21-24. – 21—23. Ardistomis vergelae n. sp. 21. habitus (dorsal view), scale bar 1 mm. 22. Male genitalia (a) ventral view, (b) right lateral view, scale bar 0,1 mm. Ap, apical portion; ba, basal apophysis; bc, basal crest; bo, basal opening; bp, basal projection; bs, basal sclerite; o, ostium; P, parameres. 23. Female genitalia (dorsal view), scale bar 0,1 mm. Gc, gonocoxae; Lt, lateroterguites 24. Ardistomis ovatus PUTZEYS Male genitalia (a) ventral view, (b) right lateral view, scale bar 0,1 mm. 70 P.R. VALDÉS

Species N HL PL PW EL EW SBL PW/PL PW/EW PL/EL

A. drumonti n. sp. 1 0,67 1,15 1,31 2,72 1,92 4,54 1,14 0,68 0,42

A. vergelae n. sp. 4 0,60-0,65 1,15-1.30 1,33-1,50 2,75-3,10 1,75-2,00 4,50-5,05 1,12-1,15 0,75-0,80 0,41-0,42

A. minutus n. sp. 1 0,42 0,73 0,74 2,05 1,17 3,20 1,01 0,63 0,36

A. samyni n. sp. 1 0,47 0,90 0,95 2,10 1,32 3,47 1,06 0,72 0,43

A. dostali n. sp. 2 0,63-0,68 1,15-1,28 1,20-1,25 3,25 1,94-2,19 5,03-5,20 0,98-1,04 0,57-0,62 0,35-0,39

A. onorei n. sp. 2 0,55-0,62 0,95-1,06 0,97-1,05 2,45-2,65 1,55-1,60 3,95-4,33 0,99-1,02 0,63-0,66 0,39-0,40

A. bulirschi n. sp. 1 0,65 1,11 1,16 2,90 1,80 4,66 1,05 0,64 0,38

Table 1. – Variation of measurements (mm) and ratios for the specimens used in descriptions.

Elytra. Oval, humeri rounded; striae distinct synonyms. Known junior synonyms are in small type. throughout their length; intervals convex. Metathoracic wings. Vestigial. Genus Ardistomis PUTZEYS 1846 Male genitalia. (Fig. 22) Phallus dextral, slender in Ardistomus CSIKI 1927 frontal view (Fig. 22a), curved in lateral view (Fig. 22b); basal bulb undefined from medial part, basal orifice alticola DARLINGTON 1935- 173 (Haiti) (bo) fusiform; basal projection (bp) distinct, notably annona PUTZEYS 1846-639 (French Guiana) sclerotized, apical margin sharp; basal crest notably arechavaletae PUTZEYS 1866- 204 developed; basal apophysis (ba) moderate, markedly curtus PUTZEYS 1866- 205 (Argentina, Brazil, Uruguay) sclerotized; ostium (o) small, opened from 2/3 of ventral atripennis PUTZEYS 1866- 202 (Guadalupe) median portion to base of apical portion; median ridge batesi PUTZEYS 1866- 209 (Brazil) moderate; apical portion (ap) slender, projected leftward, brevis PUTZEYS 1866- 204 (Brazil) margin rounded. Endophallus short; armature consist on bulirschi n.sp. (Ecuador) one small apical area covered with micro setae follow by convexus PUTZEYS 1866- 202 (Mexico) a small area covered by microtrichia; basal sclerite (bs) dostali n.sp. (Costa Rica) present, elongate, length of basal prolongations about drumonti n.sp. (French Guiana) 1/2 of total sclerite length. Paramere1 (P1) with about dyschirioides PUTZEYS 1846- 644 (Colombia, the same length of phallus median and apical portion Nicaragua, Panama) together, its median portion slender, its apical portion eductus BATES 1881- 34 (Guatemala, Mexico) wide and hook like, one seta apically; lateral apophysis elongatulus PUTZEYS 1866- 208 (Cuba) prominent, hook like; basal apophysis blunt; ventral fasciolatus PUTZEYS 1846- 638 (Argentina, Bolivia, apophysis normal. Paramere 2 (P2) slender, 4/5 length Brazil, Chile, Paraguay) of paramere 1, apex acute, two apical setae. franki NICHOLS 1988a-79 (manuscript name) (Jamaica) Female genitalia. Gonocoxa (Gc) (Fig. 23) robust, guadeloupensis KULT 1950- 307 (Guadeloupe) trapezoidal in outline; apical portion short with two haemorrhoeus PUTZEYS 1866- 207 (Brazil) setae; basal portion 4/5 of total gonocoxite length, hispaniolensis NICHOLS 1988a- 83 (manuscript name) one small seta in inner curvature. Laterotergite (Lt) (Dominican Republic, Haiti, Jamaica) rectangular in outline. leprieuri (CHAUDOIR) 1843- 740 (Bolivia, French Guiana, Venezuela) lindrothi KULT 1950- 306 (Brazil) Checklist of the species of the genus Ardistomis mannerheimi PUTZEYS 1846- 645 (Puerto Rico, Virgin PUTZEYS Islands) marquardti KULT 1950- 305 Names are given in alphabetical order. For some names, minutus n.sp. (Bolivia) a question mark indicates they are possible junior New species of the genus Ardistomis PUTZEYS 71 muelleri KULT 1950- 308 (Mexico) herbivory. Canadian Journal of Zoology, 69: 638–650. nigroclarus DARLINGTON 1939- 83 (Dominican ALLEN, R.T. & BALL, G.E., 1980. Synopsis of Mexican taxa of Republic) the Loxandrus series (Coleoptera: Carabidae: Pterostichini). nitidipennis DARLINGTON 1934- 70 (Cuba) Transactions of the American Entomological Society, 105: obliquatus PUTZEYS 1846- 638 (USA) 481–576. ogloblini KULT 1950- 303 (Brazil) BATES, H. W., 1881. Insecta, Coleoptera, Carabidae, onorei n. sp. (Ecuador) Cicindelidae. In: GODMAN, F. D. and O. SALVIN (Eds.), ovatus PUTZEYS 1846- 644 (Colombia) Biologia Centrali- Americana. Coleoptera. Volume 1, Part 1. oxygnathus CHAUDOIR 1843- 738 (Brazil, French Taylor and Francis, London, pp 1- 40, pls. 1- 2. Guiana) BOUSQUET, Y., 2006. Review of the species of Ardistomina posticalis PUTZEYS 1866- 203 ( Brazil, Ecuador) (Coleoptera: Carabidae: Clivinini) in America north of profundestriatus PUTZEYS 1866- 201 (Brazil) Mexico. Zootaxa, 1308: 1–29. quadripunctatus KULT 1950- 304 (Brazil) BRULLÉ, G. A., 1838. Insectes de l’Amérique méridionale quixotei VALDÉS 2007- 26 (Cuba, Mexico) recueillis par Alcide d’Orbigny. In : Voyage dans l’Amérique ramsdeni DARLINGTON 1937- 120 (Cuba) méridionale… par Alcide d’Orbigny. Paris. Vol. 2, pt. 2, pp. rotundipennis PUTZEYS 1866- 206 (St. Vincent and 17– 56. Grenadines) rufoclarus DARLINGTON 1939- 82 (Dominican CHAUDOIR, M. de, 1843. Carabiques nouveaux. Bulletin de la Republic) Société Impériale des Naturalistes de Moscou, 16: 671– 795. samyni n.sp. (Honduras) CSIKI, E., 1927. Carabidae, Carabinae II (Pars 92), pp. 315- schaumi LECONTE 1857- 80 (Mexico, USA) 622 . In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum seriepunctatus BRULLÉ 1838- 41 Catalogus. Berlin. brasiliensis PUTZEYS 1846- 643 DARLINGTON, P. J., Jr., 1934. New West Indian Carabidae, with constrictus PUTZEYS 1846- 643 a list of the Cuban species. Psyche, 41: 66- 131. dubius PUTZEYS 1846- 643 DARLINGTON, P. J., Jr., 1935. West Indian Carabidae II. ssp. soror PUTZEYS 1846- 642 (Argentina, Brasil, Itinerary of 1934; forests of Haiti, new species; and new key Colombia, Panama) to Colpodes. Psyche, 42: 167- 215. tropicalis PUTZEYS 1846- 642 (?) (French Guiana) DARLINGTON, P. J., Jr., 1937. West Indian Carabidae III. New unicolor PUTZEYS 1846- 640 (?) (French Guiana) species and record from Cuba, with a brief discussion of venustulus PUTZEYS 1866- 207 (Brazil, French the mountain fauna. Memorias de la Sociedad Cubana de Guiana, Venezuela) Historia Natural „Felipe Poey“. 11: 115- 136. vergelae n. sp. (Ecuador) DARLINGTON, P. J., Jr., 1939. West Indian Carabidae V. New forms from the Dominican Republic and Puerto Rico. Acknowledgments Memorias de la Sociedad Cubana de Historia Natural „Felipe Poey“. 13: 79- 101.

I wish to thank Dr. George E. Ball for reviewing an early draft of JEANNEL, R., 1946. Coléoptères carabiques de la région the manuscript and Dr. Petr Bulirsch for reviewing the manuscript malgache (première partie). Faune de l’Empire français VI. and provide material for study. I also thank Dr. Alain Drumont, Dr. Office de la Recherche Scientifique Coloniale, Paris, 372 pp. Yves Samyn and Dr. P. Grootaert for their assistance during my visit to Brussels and Dr. Thierry Deuve and Dr. Azadeh Tagavian KULT, K., 1950. New Neotropical species of the group for their assistance during my visit to Paris. I acknowledge loans Ardistomina (Carabidae. Col.). Arthropoda, 1: 299–325. from curators of public or private collections mentioned above and my wife Marydena Vergel for her help during construction of LECONTE, J. L., 1857. Synopsis of the species of Clivina and manuscript. Financial support for this study came from Belgian allied genera inhabiting the United States. Proceedings of the Development Cooperation and the Royal Belgian Institute of Academy of Natural Sciences of Philadelphia, 9: 75- 83. Natural Sciences through the Belgian Focal Point of the Global Initiative held during 2007 and 2008 calls. LIEBHERR, J.K. & WILL, K.W., 1998. Inferring phy1ogenetic re1ationships within the Carabidae (Insecta, Co1eoptera) from characteristics of the fema1e reproductive tract. In: BALL, G.E., CASALE, A. & VIGNA TAGLIANTI, A. (Eds.), References Phy1ogeny and c1assification of Caraboidea (Co1eoptera: ). Museo Regiona1e di Scienze, Atti Torino, pp. ACORN, J.H. & BALL, G.E., 1991. The mandibles of some 107–170. adult ground : structure, function, and the evolution of LORENZ, W., 2005. Systematic list of extant ground beetles of 72 P.R. VALDÉS the world (Insecta Coleoptera “Geadephaga”: Trachypachidae, VALDÉS, P., 2005. Ardistomis quixotei (Coleoptera: Carabidae: and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). 2nd Clivinini), a new species from Cuba and Mexico: structural Edition. Privately published, Tutzing, 530 pp. and habitat features of adults, larvae and pupae, comparisons with previously described species, and notes about NICHOLS, S.W., 1988a. Systematics and biogeography of classification and biogeography.Zootaxa, 1497: 23–33. West Indian Scaritinae (Co1eoptera: Carabidae). Doctoral dissertation, Cornell University, Ithaca, New York, 393 pp. WHITEHEAD, D.R., 1977. [Genus] Ardistomis PUTZEYS, p. 392. In: REICHARDT, H. A synopsis of the genera of Neotropical NICHOLS, S.W., 1988b. Ka1eidoscopic biogeography of West Carabidae (Insecta: Coleoptera). Quaestiones Entomologicae Indian Scaritinae (Co1eoptera: Carabidae). In: LIEBHERR, 13: 346-493. J.K. (Ed.), Zoogeography of Caribbean . Comstock Publishing Associates, Cornell University Press, Ithaca and London, pp. 71–120. Pavel Valdés R. PUTZEYS, J. A. A. H., 1846. Monographie des Clivina et Gertrudis 365 apto5 genres voisins, précédée d’un tableau synoptique des genres e/ D’Strampes y Goicuria de la tribu des scaritides. Mémoires de la Société Royale des Sciences de Liège, 2: 521–663. Mcpio 10 de Octubre. Cp. 10 500 C. Habana PUTZEYS, J. A. A. H., 1866. Revision general des clivinides. Cuba Annales de la Société Entomologique de Belgique, 10: (e-mail: [email protected]) 1–242.