A taxonomic teview of the subtribe Pericalina (Coleoptera: Carabidae: Lebiini) in the Western Hemisphere, with descriptions of new species and notes about classification and zoogeography

Abstract

A taxonomic review ofthe lebiine subtribe Pericalina in the Western Hemisphere, this paper includes a treatment of the genus-groups, a key to the genera, keys to subgenera, species groups, and species of each polybasic genus, descriptions of new species and new subgenera, new locality records for previously described species, re-rankings, and new synonymy. In total, 111 species and subspecies are treated, 26 of which are described as new. A review of the taxonomically useful mandibular structure, based on stereo-electron (SEM) photographs, requires changes in previously published names of various structures. A review of adhesive vestiture on the front tarsi of males shows 3 different types each characteristic of different taxa. In the eastern Brazilian montane genus Oreodicastes Maindron, 5 species are recognized, of which 3 are described as new (with type localities): O. aeaeus and O. minos (Campos de Jordao, State of Sao Paulo, Brazil); and O. rhadamanthus (Serra do Caraca, State of Minas Gerais, Brazil). In the wide-ranging genus Stenognathus Chaudoir, 19 species are recognized, and arranged in the following subgenera: the monobasic Prostenognathus, new subgenus (type species S. onorei, new species); the monobasic Gnathostenus new subgenus (type species S. dentifemoratus, new species); the polybasic Stenognathus (s. str.) (type species Anchomenus melanarius Dejean), with 16 species; and the monobasic Pristolomus Chaudoir (type species S. dentifer Chaudoir). Stenognathus cayennensis Buquet 1835 is a nomen dubium. The species of subgenus Stenognathus are arranged in 4 species groups: the S. stricticollis species group, with 4 species; the monospecific S. procerus species group; the S. nigropiceus species group (including Phloeotherates Bates and Ferus Chaudoir), with 6 species; and the S. melanarius species group, with 5 species. New species (with type localities) described herein: S. (Prostenognathus) onorei (West of Chiriboga, Pichincha Province, Ecuador); S. (Gnathostenus) dentifemoratus ("Colombia"); S. (sensu stricto, nigropiceus group) plaumanni (Nova Teutonia, State of Santa Catarina, Brazil); and S. (sensu stricto, nigropiceus group)jauja (Sani Beni, Jauja Province, Departamento Junin, Peru). Descriptions and new locality records are reported for the species known previously. For the principally Middle American genus Phloeoxena Chaudoir, the new monobasic subgenus Oxephloena is described: type species, P. turrialba, new species (Turrialba, Cartago Province, Costa Rica). Ochropisus Bates is included as a subgenus in Phloeoxena, and the following 5 new species (with type localities) are described: P. (O.) davidsoni (15 km. northwest of El Paraiso, State of Guerrero, Mexico); P. (O.) henryi (northeast slope of Volean Tacana, State of Chiapas, Mexico); P. (O.) nevermanni (western slopes of Volcan Irazu, Costa Rica.); P. (O.) turnbowi (near the trailhead of the Continental Divide Trail, Chiriqui Province, Panama); and P. (O.) lamuralla (Parque Nacional La Muralla, Olancho Department, Honduras). In Phloeoxena (Oenaphelox), the following 4 new species are described: P. (O.) totontepec (16.6 km south of Totontepec, State of Oaxaca, Mexico); P. (O.) brooksi (El Guisayote, 24 km east of Ocotepeque, Ocotepeque Department, Honduras); P. (O.) ashei (Parque Nacional La Muralla, 14 km north of La Union, Olancho Department, Honduras); and P. (O.) viridis (El Guisayote, 24 km east of Ocotepeque, Ocotepeque Department, Honduras). In Phloeoxena (s. str.), the following 2 new species are described: P. (P.) nitida (51.5 km south of Valle Nacional, State of Oaxaca, Mexico); and P. (P.) obscura (vicinity of Coscomatepec, State of Veracruz, Mexico). Based on study of the holotype, Phloeoxena biundata Steinheil is included in the subgenus Phloeoxena. The status of P. (P.) picta batesi Ball is changed from subspecies to species, based on new locality data indicating extensive range overlap of P. picta (sensu lato) and P. batesi. New locality records are provided for the following previously described species of Phloeoxena: P. (Tacana) herculeano Ball; P. (Ochropisus) concolor Ball (substantial range extension in western Mexico); P. (Ochropisus) caudalis Bates; P. (Oenaphelox) newtoni Ball (a second record); P. (Oenaphelox) undata Chaudoir; P. (Oenaphelox) geniculata Chaudoir; P. (Oenaphelox) signata Dejean; P. (s. str.) dealata Darlington; P. (s. str.) imitatrix Darlington; P. (s. str.) schwarzi Darlington; P. (s. str.)portoricensis Darlington; P. (s. sir.) megalops erioinorum Ball (range extension from Costa Rica to Panama; P. (s. sir.) limbicollis Bates; P. (s. sir.) nigricollis Ball; P. (s. str.) picta franiae Ball; P. (s. sir.)picta unicolor Chaudoir; P. (s. str.)picta picta Chaudoir; P. (s. str.)picta apicalis Ball; and P. (s. str.) batesi Ball. Two monobasic species and 1 dibasic species of the pantropical genus Catascopus Kirby are recognized. New synonymy is as follows: C. obscuroviridis Chevrolat 1835 = C. mexicanus Chaudoir 1877 = C. angulicollis Bates 1878; C. validus Chaudoir 1854 = C. guatemalensis Bates 1883; C. brasiliensis Dejean 1825 includes the South American C. b. brasiliensis (new status) = C. cayennensis Chaudoir 1872 and the Middle American C. b. chontalensis Bates 1878 (new status). New locality records for the 3 Middle American species, C. obscuroviridis, C. validus, and C. b. chontalensis show extensive range overlap. Nonetheless, C. validus seems to be isolated altitudinally from the other 2 species. Catascopus b. brasiliensis is a rather varied species and wide-ranging in South America, from the Brazilian Atlantic Forest to the Upper Amazon Basin, in Peru and Ecuador. The monospecific Chilean-Argentinian genus Catascopellus Straneo (type species C. crassiceps Straneo) is removed from the Somotrichus genus-group and placed in the monogeneric Catascopellus genus-group, primarily on the basis of divergence in the combination of structural details of the mouthparts and adhesive vestiture of the front tarsi of males. The Catascopellus genus-group in many of its diagnostic features is much like the Thyreopterus genus-group, but males of the 2 groups differ from one another in the adhesive vestiture of the fore tarsi. The known geographical range of C. crassiceps is extended from eastern Chile to western Argentina. The precinctive Neotropical Eurycoleus genus-group, including only Lelis Chaudoir and Eurycoleus Chaudoir, is recharacterized, with special reference to details ofthe mandibles and labium. The 4 valid species of Lelis, (with synonyms of the valid names) follow: the South American L. obtusangula (Chaudoir) 1852; the South American L. quadrisignata (Buquet) 1834 = L. polygona (Bates), 1869 = L. bifasciata Chaudoir 1869, new synonym = L. latipennis (Bates) 1869, new synonym; the South American-Middle American L. rutila (Bates) 1869 = L. viridipennis Chaudoir 1869 = L. cyanipennis Steinheil 1875, new synonym = L. insculpta (Bates) 1893, new synonym; and the Middle American L. bicolor Chaudoir 1869. Putative adelphotaxon relationships, based on structural details of adults, are: L. obtusangula (L. quadrisignata (L. rutila + L. bicolor)). Based on color pattern, the 8 species of the genus Eurycoleus Chaudoir are arranged in 2 species groups: the E. poecilopterus group with 2 species; and the E. tredecimpunctatus group, with 6 species. The species of the latter species group are arranged in 2 subgroups, the septemplagiatus subgroup, with 2 species, and the tredecimpunctatus subgroup, with 4 species, including E. erwini, new species (Estacion Sirena, Playa Sirena, Puntarenas, Costa Rica). Both species groups and both subgroups are represented in South America and Middle America by precinctive species. New locality records for Eurycoleus poecilopterus Buquet extend its range to northern Argentina, the southernmost area for the genus. The sole genus in the Western Hemisphere of the Pericalus genus-group, Copiodera Dejean, another pantropical genus, is represented by 44 species. New locality records are recorded for the following: C. elongata Putzeys; C. schaurni Chaudoir; C. megalops Bates (range extension into Middle America, from South America); C. championi Bates; C. versicolor Bates; C. nigrostriata (Reiche); C. lineata (Bates); C. transversa (Reiche); C. relucens Bates; C. festiva Dejean; C. aeneorufa Bates; C. tripartita Chaudoir; C. cupreotincta Bates; C. chalcites Bates; C. acutipennis Buquet (all recently acquired males were assigned to previously established morphs based on details of the armature of the internal sac); C. picea Dejean; C. nitidula Buquet; and C. brunnea Shpeley and Ball. A review of the classification of the Neotropical indigenous Eucheila genus-group, based on characters of newly discovered species, and on re-evaluation of relationships, indicates recognition of a single genus, Eucheila Dejean, with the following newly ranked taxa as subgenera: Hansus Ball and Shpeley (type species Hansus reichardti Ball and Shpeley); Bordoniella Mateu (type species Bordoniella lucida Mateu): Inna Putzeys (type species Inna punctata Putzeys); and Eucheila Dejean (type species Eucheila flavilabris Dejean). Pseudoinna Mateu (type species I. boliviana Mateu), described as a subgenus of Inna, is recognized as a subgenus of Eucheila. In subgenus Eucheila, a new species group is proposed: E. erwini species group, with 2 species. New species are: E. (Hansus) kiplingi (Puerto Misahuali, Napo Province, Ecuador); E. (Bordoniella) marginate. (Rio Tambopata Reserve [12°50'S 069°20'W], Departamento Madre de Dios, Peru); E. (Pseudoinna) mateui (Pakitza, Departamento Madre de Dios, Peru); E. (P) surinamensis (Rainville, Suriname district, Surinam); E. (Eucheila) erwini (Rio Napo Explomapocamp [03°15'S 072°55'W], Loreto Department, Peru); and E. (E.) pilosa (about 75 km south-southeast of Apoera, Nickerie District, Surinam). New distribution records are provided for the following species: E. (H.) reichardti (Ball and Shpeley) (range extension to Upper Amazon Basin in Peru, from Guyana in northeastern South America); E. (I.) purpurea (Ball and Shpeley); E. (I.) breviformis (Chaudoir); E.(I.)costulata (Chaudoir); E. (I.) boyeri (Solier) (range extension from mainland to Aruba, Dutch West Indies); E. (I.) nevermanni (Liebke) (range extension from Costa Rica to Panama); E. (I.) megala (Reichardt); E. (P) inpa (Ball and Shpeley) (range extension from Central to Upper Amazon Basin); E. (P) boliviana (Mateu) (range extension from Bolivia to Peru and Ecuador); E. (E.) strandi Liebke; E. (E.) cordova Ball and Shpeley (range extension from a single locality in Veracruz State, Mexico to Belize, in Middle America); E. (E.) adisi Ball and Shpeley (range extension from Central to Upper Amazon Basin, in Peru and Ecuador). Study of the mouthparts suggests that their evolution in the Pericalina of the Western Hemisphere has involved changes in form, reductions through loss or consolidation, and gains represented by additional setae. Overall, 2 basic types of mouthparts are recognizable: Type A, characteristic of the Eucheila genus-group; and Type B, characteristic of the Thyreopterus, Catascopellus, Pericalus, and Eurycoleus genus-groups. Type Aexhibits more numerous modifications than Type B, but the markedly derived features of Type A are combined with retention of plesiotypic features, suggesting that the Eucheila genus-group separated early from those genusgroups with Type B mouthparts. The basic Type B pattern has substantial modifications, useful for taxonomic recognition; both within and between genus-groups that exhibit this type of mouthparts, and probably for phylogenetic analysis. From a functional perspective, the less derived forms of mouthparts, as in, for example, the thyreopteroid Stenognatlius onorei, or the eucheiloid Eucheila marginata, indicate general predaceous feeding, involving particulate matter. The most striking modifications of the mouthparts, as seen in the subgenus Eucheila, suggest a profound change in food or feeding mechanisms, from consumption of particulate matter to consumption of liquids, or liquefied tissue. The distribution pattern of genera and species is fitted to a general model purporting to explain such patterns, in terms of plate tectonics, refuge theory, and climatic change during the Tertiary Period. Concluding remarks offer explanation of and justification for the taxonomic changes, including proposal of monobasic taxa and recognition of species based on limited material.