George E. Ball

Conspectus of Carabid Classification: History, Holomorphology, and Higher Taxa

The purpose of an historical presentation is to develop in a tempral context events and circumstances that led to the state of affairs perceived at a particular time. History is, collectively, a record of the activities of Homo sapiens. In preparing an account of a portion of this record, one necessarily picks and chooses, and generally these activities are based on the viewpoint espoused by the author of the history. Or, put another way, a history is a reconstruction of events. As one knows, no two builders are apt to develop their structures in exactly the same way though both may use very similar materials. So, what follows is not the history of carabid classification, but a history as seen by one who appreciates and respects the workers of the past, who believes that science advances by evolution rather than by revolution, and who, as an evolutionist, believes that phylogeny and classification must be linked more or less closely.

A Taxonomic Review of the Genus Apenes Leconte (Coleoptera: Carabidae: Lebiini) in the West Indies, with Descriptions of New Species and Notes About Classification and Biogeography

ABSTRACT Primarily a taxonomic review of the West Indian elements of Apenes LeConte, this paper includes a classification, a key to species, descriptions of taxa, re-rankings, and new synonymy. In total, 45 species and subspecies are treated, 14 (11 species, three subspecies) of which are described as new. The taxa are arranged in the following sequence, with type localities of new species and subspecies in parentheses following the name of the taxon. The seven species of the Apenes variegata species group are included in two species complexes: A. variegata species complex, with A. variegata (Dejean), A. marginalis (Dejean), A. coriacea (Chevrolat), and A. sulcicollis (Jacquelin du Val); and A. morio species complex, with A. parallela (Dejean) (including the subspecies A. p. parallela (Dejean), A. p. inaguae Darlington, and A. p. sublaevis Darlington), A. portoricensis Darlington, and A. morio (Dejean). The A. sculpticeps species group is monobasic in the West Indies, including A. sculpticeps, new species (Barro Colorado Island, Panamá). Also monobasic is the A. prasina species group, including only A. prasina Ball and Shpeley. The A. purpurata species group includes seven species: A. kathleenae Ball and Shpeley; A. purpurata Fleutiaux and Sallé; A. chalumeaui Ball and Shpeley; A. ovalis Darlington; A. aptera Darlington; A. youngi, new species (13.5 km SE Puerto Escondido, Independencia Province, Dominican Republic, Hispaniola); and A. pecki, new species (Gran Piedra, Santiago Province, Cuba). The A. lucidula species group includes A. lucidula (Dejean), represented in the West Indies by the subspecies A. l. dulculia Ball and Shpeley, and A. l. michelii Ball and Shpeley, and A. lucia, new species (Anse La Raye, St. Lucia, Windward Islands). The 16 species and six subspecies of the A. fasciata species group are arranged in three species complexes: the monobasic A. farri species complex, including A. farri Ball and Shpeley; the A. fasciata species complex, including A. iviei Ball and Shpeley, A. darlingtoni Ball and Shpeley, A. davidsoni, new species (along Rio Mulito, 13 km N. Pedernales, Pedernales Province, Dominican Republic), and A. fasciata Chaudoir; and the A. lata species complex, including A. lata Darlington, A. philipi, new species (2 km N. Bayahibe, La Altagracia Province, Dominican Republic), A. dominica Ball and Shpeley, the polytypic A. delicata, including A. d. delicata Darlington, A. d. laevior Darlington, and A. d. cayman Ball and Shpeley, the ditypic A. thomasi, new species, including A. t. thomasi, new subspecies (Great Inagua Island, Bahamas) A. t. swearingenae, new subspecies (Salt Cay, Turks and Caicos Islands), A. toussainti, new species(swamps N. Dessalines, Haiti), A. lepidula Darlington, A. laevicincta Darlington, A. lachauxi, new species (Etang Lachaux, Departement du Sud, Haiti), A. rawlinsi, new species (3 km west of Cueva de Arena, Parque Los Haitises, El Seibo Province, Dominican Republic), the polytypic A. scobifera including A. s. scobifera Darlington, A. s. melaena Ball and Shpeley, A. s. steineri, new subspecies (1.5 km S. Hutland, 19°20′N 81°13′W, Grand Cayman Island), and A. s. turnbowi, new subspecies (Great Inagua Island, Bahamas). The A. postica species group includes A. faber, new species (St. Vincent, Windward Islands), and A. plaumanni (Liebke). The name A. faber is used in place of A. steinheili Ball and Shpeley, which in turn was a replacement name for A. fasciata Steinheil (not Chaudoir), a species known only from Colombia. Apenes laevis Liebke is a junior synonym of A. coriacea (Chevrolat), new synonymy, and Apenes strandi Liebke is a junior synonym of A. delicata delicata Darlington, new synonymy. Ecologically, the species of Apenes are primarily geophilic mesophiles and xerophiles, occupying primarily the forest floor, ranging from lowland dry seasonal to montane wet cloud forest and elfin woodland. A few species occupy also human-disturbed situations or open areas adjacent to sea beaches. Adults are active on the ground and in flight during darkness. During daylight, adults can be found in leaf litter, or under other debris on the ground, such as fallen tree trunks. Adults are active in every month of the year but principally during spring and early summer (March—June). Elevational range of occupied habitats extends from sea level to 3000 m, but species are most abundantly distributed at lower elevation (500 m or less). Brachyptery (nine of 45 taxa) is associated with, but not confined to, life at higher elevations, principally in wet montane forest or elfin woodland. Prominent aspects of a postulated biogeographical history of the West Indian species of Apenes are the following: - From the tropical American source areas of northern South America and southern Middle America (or Central America), 12 Apenes stocks invaded the West Indies over a time span that extended from the early Oligocene to the Pleistocene, or about 37 MYBP. - The Greater Antilles were populated by six ancestral stocks now represented only by West Indian precinctive taxa from South American and Middle American source areas. Early arrivals could have come by way of an emergent Aves Ridge that was attached at one end to South America, and at the other to the emergent proto-Greater Antilles. Later arrivals likely came by flight or were carried by flotsam, probably using temporarily emergent (now submerged) islands along the Nicaragua Rise that extended from Central America eastward to western Jamaica, and from there across the Jamaica channel to Hispaniola. A second overseas portal for Miocene-age stocks to enter the Greater Antilles was western Cuba. - Evolution in the Greater Antilles included early vicariance (as the proto-Greater Antilles evolved into its present form), later inter-island dispersal, inter- and complex intra-island differentiation (principally in Hispaniola), extinction, and ecological shifts, as exemplified by elevational distribution and brachyptery. - Thus the history of the Greater Antillean Apenes assemblage is dominated by relatively old arrivals and extensive evolution in structural and ecological features. - The Lesser Antilles have three stocks with precinctive species, two of whose ancestors originated in the Greater Antilles, and one on the South American mainland. Only one of these stocks differentiated and is represented by two species. Three monobasic stocks confined in the West Indies to the Lesser Antilles are represented on the mainland by conspecific populations and are regarded as adventive, relatively recent (Pleistocene) arrivals. - The ranges of two other adventive taxa extend from the tropical American mainland through the Lesser Antilles and deep into the Greater Antilles (probably also Pleistocene arrivals). - Thus the history of the Lesser Antillean Apenes assemblage is dominated by relatively recent arrivals and relatively minor evolution in structural and ecological features.

The first instar larva of Eripus oaxacanus Straneo & Ball (Coleoptera: Carabidae: Peleciini): indicator of affinity or convergence?

Abstract. The first instar larva of Eripus oaxacanus Straneo & Ball is diagnosed and described, representing the first laboratory‐described larval stage for a species of the tribe Peleciini. The larva possesses the derived character states of: (1) stemmata lacking, (2) mandible lacking penicillus, (3) maxilla with setose lacinia, (4) labium lacking a ligula, (5) claws single, (6) urogomphi short, rounded. These derived character states, if considered synapomorphic, would place Peleciini as a close relative to Brachi‐nini. Classifications based on adult characters are not concordant with this interpretaton, and the weight of evidence suggests that convergence due to a similar parasitic lifestyle is the basis for the derived similarity of Eripus and Brachinus larvae.

Western Hemisphere Zuphiini: descriptions of Coarazuphium whiteheadi, new species, and Zuphioides, new genus, and classification of the genera (Coleoptera, Carabidae)

Abstract Based on small samples (exemplars) analyzed with morphological methods, including detailed descriptions and illustrations, this study treats primarily the Zuphium genus-group in the Western Hemisphere, which comprises two precinctive genera: Coarazuphium Gnaspini, Vanin & Godoy, 1998 (type species Parazuphium tessai Godoy & Vanin, 1990) and Zuphioides gen. n. (type species Zuphium mexicanum Chaudoir, 1863). The genus Coarazuphium includes six troglobitic species from Brazilian caves, and one probably hypogaeic (troglophilic) species from the mountains of Oaxaca, in Mexico (Coarazuphium whiteheadi, sp. n., type locality, ridge top, in western Oaxaca, Mexico, at 2164 m, 35 km north of San Pedro Juchatengo, 16.462N, 97.010W). The epigaeic genus Zuphioides includes 23 species, with its geographical range extended from Neotropical temperate Argentina in southern South America, northward through the tropics to north temperate southeastern Canada, in the Nearctic Region. Keys are provided to the species of Coarazuphium and to thegenera of Western Hemisphere Zuphiini.

Mandibles and labrum-epipharynx of tiger beetles: basic structure and evolution (Coleoptera, Carabidae, Cicindelitae)

Abstract Using for comparison with, and as outgroups for, supertribe Cicindelitae, we describe and illustrate the mandibles and labrum-epipharynx of the basal geadephagans Trachypachus gibbsii LeConte, 1861 (family Trachypachidae), and family Carabidae: Pelophila rudis (LeConte, 1863) (supertribe Nebriitae, tribe Pelophilini) and Ceroglossus chilensis (Eschscholtz, 1829) (supertribe Carabitae, tribe Ceroglossini). The range and pattern of variation in structure of mandibles and labrum-epipharynx within the supertribe Cicindelitae was assessed using scanning-electron (SEM) images of these structures in nine exemplar taxa: Amblycheila baroni (Rivers, 1890), Omus californicus (Eschscholtz, 1829) and Picnochile fallaciosa (Chevrolat, 1854) (representing the Amblycheilini); Manticora tuberculata (DeGeer, 1778) (representing the Manticorini): Tetracha carolina (Linnaeus, 1767) (representing the Megacephalini); Pogonostoma chalybeum (Klug, 1835) (representing the Collyridini); and Therates basalis Dejean, 1826, Oxycheila species, and Cicindela longilabris Say, 1824 (representing the Cicindelini). An evolutionary transformation series was postulated for the mandibles and labrum-epipharynx, based on a reconstructed phylogenetic sequence, which, in turn, was based on morphological and DNAevidence.Principal features of the transformation series for the mandibles included development of a densely setose basal face; wide quadridentate retinaculum; a lengthened incisor tooth; a multidentate terebra (one to five teeth; two-three most frequent), followed by subsequent loss of one or more such teeth; development of a diastema in the occlusal surface; development and subsequent loss of scrobal setae, and reduction and loss of the scrobe. Principal features of the transformation series for the labrum included evolution of form from transverse, sub-rectangular to elongate almost square, to triangular; position and number of setae evolved from dorsal to insertion on the apical margin, the number increased from 8-10 to as many as 36, and decreased to as few as four. The epipharynx broadened evolutionarily, the pedium evolving in form from narrow, triangular and nearly flat, to broad, palatiform, and markedly convex; anterior parapedial setae both increased and decreased in number, and in orientation, from a row parallel to the parapedial ridge to a setal row extended forward at about a right angle to the latter.

Hybopteroides, a new genus in the Cryptobatida group of subtribe Agrina, with three new species and notes on their way of life (Insecta: Coleoptera, Carabidae, Lebiini)

Abstract Hybopteroides gen. nov., with three new species in the Cryptobatida group of subtribe Agrina, Lebiini, living in the lowland rain forest canopy of Ecuador and Perú are diagnosed, described and illustrated. The new genus is likely the adelphotaxon of Hyboptera Chaudoir 1872 + Thoasia Liebke 1939. Hybopteroides biolat sp. nov., the type species, is now known from the Río Manu watershed in Perú; Hybopteroides karolynae sp. nov. is now known from the Río Tambopata watershed in Perú; and Hybopteroides penrosei sp. nov. is now known from the upper and mid Río Napo watershed in Ecuador. The species of this new genus are known from insecticidal fogging collections from the rainforest canopy and upper understory and from two specimens collected by E. S. Ross at his ecotourism lodge at Alinahui, Ecuador, found in his live embiid colonies. Resumen Un nuevo género, Hybopteroides gen. nov., con tres nuevas especies del grupo Cryptobatida de la subtribu Agrina, Lebiini, que viven en los bosques lluviosos de tierras bajas de Ecuador y Perú se diagnostican, describen e ilustran. El nuevo género es probablemente el adelfotaxón de Hyboptera Chaudoir 1872 + Thoasia Liebke 1939. Hybopteroides biolat sp. nov., la especie tipo, se conoce de la cuenca hidrográfica del río Manu en Perú; la especie Hybopteroides karolynae sp. nov. se cononce de la cuenca hidrográfica del Río Tambopata en Perú; y la especie Hybopteroides penrosei sp. nov. se conoce de la parte superior y media de la cuenca hidrográfica del Río Napo en Ecuador. Las especies de este nuevo género se conocen de colecciones obtenidas del dosel del bosque lluvioso utilizando la técnica de termonebulización con insecticida y de dos especímenes colectados por E. S. Ross en sus cabañas ecoturísticas en Alinahui, Ecuador, en sus colonias vivas de Embioptera. Palabras claves.Ecuador, Perú, dosel de bosque lluvioso, sotobosque de bosque lluvioso Hyboptera Chaudoir, Embioptera

On the Provenance of Boheman's “Eugenies Resa” Carabidae (Coleoptera) Allegedly Described from Hawaii

ABSTRACT Five carabid beetle species described from the Hawaiian Islands, Polynesia—Calleida gracilis Gemminger and de Harold, 1868 (= Calleida amoenula Boheman, 1858), Calleida sanguinicollis Dejean, 1831 (= Calleida insularis Boheman, 1858), Lebia insularis Boheman, 1858, Selenophorus insularis Boheman, 1858, and Selenophorus picinus Boheman, 1858—are shown to be properly attributed to the American biogeographic region, with their type localities corrected to either Central America or Pacific South America. Two of the five names are newly synonymized: L. insularis Boheman = L. analis Dejean, 1825 (new synonymy), and S. insularis Boheman = S. chalcosomus Reiche, 1843 (new synonymy). Calleida gracilis is shown to be a member taxon of the janthina species-group of South America, most similar to Calleida tibialis Brullé, 1837. However, taxonomic uncertainty surrounding C. gracilis, C. tibialis, and an undescribed taxon precludes definitive species circumscription in this group, and C. gracilis is retained as a valid member of the janthina species-group pending future revision. Selenophorus picinus is placed as species incertae sedis within Selenophorus subgenus Gynandropus Dejean, alternately treated as the hylacis species-group. The biogeographical and ecological consequences obtained from removing these taxa from the Polynesian fauna include: 1) recognition of more natural biogeographical distributions for the genera Calleida Latreille and Dejean, Lebia Latreille, and Selenophorus Dejean; 2) elimination of all examples that could demonstrate extinction of a non-native carabid beetle species subsequent to its accidental introduction into the Hawaiian Islands.

'Bellography': Life and Contributions of Ross and Joyce Bell, two New England Naturalists.

Abstract The lives and contributions of Ross and Joyce Bell are described with particular attention to studies of invertebrate natural history in the state of Vermont and carabid beetles of several groups, including the world rhysodine fauna. Their work, all done at the University of Vermont, was mainly taxonomic in nature and included aspects of the biology of the species considered. During their careers they described more than 75% of the c. 340 rhysodine species known to science. Ross Bell also wrote a number of seminal papers about the basal relationships of the Adephaga and the comparative anatomy of carabid coxal cavities. Ross and Joyce inspired several generations of students at UVM to take up advanced work in entomology and natural history.

Introduction—Three Leaders

Over the years, carabid beetles have attracted a variety of distinguished and highly able systematists, and during the last 50 years three of the ablest have devoted their energies to the study of this family. I want to record a few words about these men, and about their contributions. They are Rene Jeannel, Philip Darlington, and Carl Lindroth. The last two are extant, and contributed to the program of this Symposium.