Badister Clairville 1806: A new species and new continental record for the nominate subgenus in Amazonian Perú (Coleoptera, Carabidae, Licinini)

Abstract Badister (Badister) amazonus sp. n. is described from Perú, Loreto, 1.0 km SW Boca del Rio Samiria, Vigilante Post 1, 130m, “04°40.5'S, 074°18.9'W" its type locality. It is known also from two other localities in Loreto Department, Perú, in both the Varzea and Igapó river systems. This new species is sufficiently different that a new informal higher taxon, the amazonus species complex, is recognized. An updated key to the Western Hemisphere species of subgenus Badister is provided.

especie es suficientemente diferente que un nuevo taxón más elevado, el complejo de especies amazonus, se reconoce. Se presenta una clave actualizada de las species del subgénero Badister para el Hemisferio Occidental. Taxonomía, clasificación, Licinina, Baudia Ragusa, 1884, clave de especies, región Neotropical, Varzea, Igapó introduction Unlike the movements of the comic book character Superman, taxon range extensions rarely come to our notice in "leaps and bounds." And, almost never does a newly discovered member of a subgenus show up on a new continent quite apart from its well recorded and quite distant range. Ball (1959) noted that the southernmost occurrence of the subgenus Badister (s. str.) Clairville, 1806 in the Western Hemisphere was in Oaxaca, México "on or near the edge of the Neotropical Region." Since then, Ball (1992) also recorded Badister (Baudia) reflexus LeConte from Quintana Roo, Badister (s. str.) flavipes mexicanus Van Dyke from Chiapas and Tabasco, México, somewhat south and east of Oaxaca, and a single specimen of subgenus Baudia Ragusa 1884 from Tucumán, Argentina. In the collection at the Strickland Museum, University of Alberta, Edmonton, Canada, an undescribed brachypterous species from Veracruz, México is represented and at the National Museum of Natural History, Washington, DC, seven undescribed species of the subgenus Baudia are represented from various places in southern México, Belize, Honduras, and Costa Rica. Before the discovery reported in this paper, the southern limits in the Western Hemisphere of the subgenus Badister was known to be in Chiapas and that of subgenus Baudia in the Province of Puntarenas, Costa Rica, and according to Ball (1992) in Argentina (noted above). Another licinine monobasic genus, (the rather distantly related though con-subtribal) Eutogeneius Solier, 1849, is known from Chile. Between that country and Argentina and Costa Rica, nothing had been recorded for Licinini and the tribe was considered amphi-tropical.

Palabras clave
To the senior author's great surprise on the night of May 21, 1990, at Cocha Shinguito, Loreto Department, Perú, he encountered two adult members of a relatively large species of Badister in a swamp forest at the edge of a standing pond in wet leaf litter overlaying gray clay soil (Kaolin). On a subsequent expedition to the same watershed the following year with the junior author, many more adults were encountered at two localities in low damp or wet places near the mouth and at the midpoint of the Río Samiria in the Pacaya-Samiria National Reserve (noted without discussion by Ball 1992). This river lies in a vast subsidence zone on the western edge of the Amazon Basin. The watershed is composed of "black water;" this includes the Igapó main river, many smaller 2 nd order streams, and hundreds of small to large oxbow lakes ("cochas" in Spanish and on some country maps). On the subsequent expedition mentioned above, this species was again found (September 2, 1991) along a major white water river, the Río Ucayali. We take this opportunity, as part of the Bell Fest Symposium volume, to describe the new species, its way of life as far as is known, and its place in a biogeographic context of the Genus Badister Clairville.
Ross and Joyce Bell, to whom this volume is dedicated, have lived in Vermont, USA, throughout their long and productive careers in taxonomic studies of Rhysodini and other carabids. Coincidentally, they have lived in an area of maximum species richness for Badister, with no less than five species occurring in their surroundings.

specimens and methods
Included in this study are a total of 58 specimens from the National Museum of Natural History, Washington, DC (NMNH, Terry L. Erwin, Curator) and 14 specimens from the Strickland Museum, University of Alberta, Edmonton, Canada (UASM, George Ball and Danny Shpeley, Curators). Some of these specimens will be deposited in the California Academy of Sciences (CAS, David H. Kavanaugh, Curator), the Carnegie Museum of Natural History (CMNH, Robert Davidson, Collection Manager), the Museum of Comparative Zoology (MCZ, Philip Perkins Collection Manager), and the holotype in the Museo de Universidad de San Marcos, Lima, Perú (MUSM, Gerardo Lamas, Curator).
Methods and species concepts follow those previously described (Ball 1959;Erwin and Kavanaugh 1981;Kavanaugh and Erwin 1991). The species validation and diagnosis format follows as closely as possible that suggested in Erwin and Johnson (2000). Measurements of length (ABL, SBL) and width (TW) follow those of Ball (1972) and Kavanaugh (1979): ABL (apparent body length), measured from apex of labrum to apex of longer elytron; SBL (standardized body length), equals the sum of the lengths of the head (measured from apex of clypeus to a point on midline at level of the posterior edge of compound eyes), PL (pronotal length ), measured from apical to basal margin along midline, and LE (elytron length), measured from apex of scutellum to apex of the longer elytron; and TW (total width), measured across both elytra at their widest point with suture closed.
Terms used for the pygidial gland system were taken from Will et al. (2000). For the ovipositor sclerites, terms used are those of Liebherr and Will (1998). For explanation of orientation of the surfaces of gonocoxite 2, see Ball and Shpeley (2009: 92). Note also that Ball (1959) referred to gonocoxite 2 as the "stylus" and its blade as the "digitus." The terms for the parts of the male genitalia are either standard among carabidologists, or are taken from Ball and Shpeley (2009: 92). Attributes of the abdominal ventral sterna are referred to using the numbering system generally accepted in Carabid studies, i.e., the sternum divided medially by the hind coxae is sternum II (the first being hidden) and the last visible is sternum VII (Liu et al. 2011).
The habitus image ( Fig. 1) of the adult beetle portrays most of the character states referred to in the key provided. Male and female genitalic digital photo-illustrations are standard for descriptive taxonomy of carabid beetles. The images of the adult and its parts were made with a Visionary Digital TM high resolution imaging system. Figure  captions include an ADP number, which is a unique identification number for the specimen that was illustrated or imaged and links the specimen and associated illustrations and/or image to additional information in electronic databases at the NMNH.
Throughout this paper, a hyphen (-) is used for its normal purpose; a dash (-) is used to indicate a continuum between entries (e.g., between measures, altitudinal ranges, Figure numbers or letters, or across months). Geographical data are presented for the new species based on all known specimens available at the time of manuscript preparation. Georeferences have been determined from locality information provided on specimen labels. Latitude and longitude are reported in decimal degrees. A distribution map is provided for the species (Fig. 7). Here, an English vernacular name is proposed, as vernacular names are becoming increasingly needed in conservation and/or agricultural and forestry applications, as well as for the Encyclopedia of Life (www.eol.org). Geographic Distribution. Members of this genus occur in the Antillean subregion of the Neotropical Region, Amazon Basin of Perú, the Holarctic Region, Afrotropical Region, in the Cape sub-region, and on the island of Madagascar (Ball 1959), Oriental Region eastward to Java, and the Australian Region, including New Guinea and eastern Australia (Darlington 1968: 15).

Accounts of taxa
Habitat. Wet and moist areas including shaded margins of vernal pools in deciduous forests, open margins of alkaline lakes, secondary floodplains of river banks, at eutrophic marshes, pond and lakes, in disturbed places such as gravel pits, vacant lots, pastures, and cultivated fields (e.g., barley).
Description. Head ( Fig. 2) with two supraorbital setigerous punctures per eye, frontal impressions usually shallow, basin-like. Clypeus (cl) markedly and slightly asymmetrically emarginate, not divided into two portions by the emargination, deflected ventrally, forming an angle of somewhat less than 90 degrees with the front, bearing a single long seta on each side, each of which originates in a broad approximately triangular pit. Eyes prominent. Antenna with complex variety of sense organs beginning on antennomere 3 (a3). Labrum (lb) about 2/3 as wide as clypeus, deeply bilobed, cleft anteriorly almost extended to base, with three setae on each lobe. Mandibles very short and obtuse, terebra markedly reduced, apparently edentate; either right or left mandible deeply notched (mn), preceded by a prominent swelling, or boss (mb). Maxillary palpi long and slender, palpomere 4 (mp4) fusiform, slightly broader than palpomeres 1-3, narrowly truncate apically. Labial palpus with palpomere 2 bisetose; palpomere 3 (lp3) securiform, distal margin obliquely truncate.
Prothorax. Pronotum ( Fig. 1) wider than head, more or less cordate, a pair of setigerous punctures on each side, one of the pair at posterior lateral angle, the other just inside lateral margin about half way between base and apex; narrowly margined laterally, hind angles obtuse, slightly to markedly rounded. Proepisternum with microsculpture mesh pattern longitudinal; microlines fine, transverse on remaining lateral and ventral thoracic sclerites. Prosternum with apex of intercoxal process feebly margined, asetose.
Pterothorax. Metepisternum elongate, the outer margin about 1.5 times greater in length than the anterior margin, posterior margin about 0.75 times anterior margin.
Elytra. Oblong, wider than pronotum at widest point, apical margin entire or very slightly sinuous, interneurs fine, parascutellar interneur (Fig. 3A, pss) long, joined or not to apical portion of interneur 1; intervals flat to slightly convex with two discal punctures on interval 3, adherent to interneur 2. Lateral marginal (umbilical) series of 15-18 setae, concentrated and narrowly spaced in anterior and posterior thirds, with few widely spaced setae medially.
Hind wings. Macropterous or brachypterous. Venation not studied. Legs. Slender; tibial spurs very finely serrulate; posterior tarsi very slender, dorsal face dull and medially feebly sulcate; anterior tarsi of male (Figs 3B-C) with tarsomeres 1-3 markedly or slightly dilated, with extensive or limited patch of adhesive articulo-setae ventrally (Fig. 3c) (also, see Stork 1980: 195, 290, for detailed description of adhesive setae of Badister bipustutulatus (Fabricius)) Abdominal sterna. Surfaces with very fine, transverse microlines, very close together, surface quite iridescent. Abdominal sternum VII with two setigerous punctures on posterior margin in male, four in female, apical margin more rounded in the male, more truncate in the female.
Ovipositor and female reproductive organs (Figs 6A-D). Gonocoxite 2 (gc 2) falcate, base (b) about as long as blade (bl), latter relatively short, pointed distally; margins with several ensiform setae; with or without short preapical nematiform setae. Reproductive organs standard for Carabidae, with bursa copulatrix, common oviduct, spermatheca, and spermathecal glands (for details, see this topic below, in description of Badister amazonus, n. sp.). Diagnosis. In addition to the features presented in the generic diagnosis and description above, all of the Western Hemisphere species of the typical subgenus have the right Figure 2. Badister amazonus sp. n., dorsal aspect, female, ADP051544; type locality. Head, including eyes, antennomeres 1-3, mouthparts, and anterior part of pronotum. Legend: a3, antennomere 3; cl, clypeus; lb, labrum; lp3, labial palpomere 3; mb, mandibular boss; mn, mandibular notch; mp4, maxillary palpomere 4. mandible deeply notched (Fig. 2, mn), and a row of setae on each latero-ventral margin of tarsomere 5. Another useful attribute in distinguishing the subgenera is the relative length of the hind tarsi. In adults of Badister and Trimorphus, the posterior tarsi are more than 3/4 the length of the hind tibiae; in Baudia, the hind tarsi are perceptibly relatively shorter. Anterior margin of proepisternum and lateral margins of prosternum distinctly rugose; maxillary palpomere 3 distinctly shorter than palpomere 4; head and pronotum black, shining surface of pronotum smooth without microlines; head with mesh pattern isodiametric, microlines shallow, very fine,  Derivation of specific epithet. The epithet "amazonus" is a singular Latinized masculine noun in apposition, based on the name of the area in which these beetles were found.

Pterothorax. Normal for genus.
Elytra. Elytron about same width as head across eyes (mean WH/WE: 0.998 for both sexes), moderately convex, intervals moderately convex and slightly more so laterally, interval 3 bisetose with each setigerous puncture adherent to interneur 2.
Abdominal sterna with normal setation for genus. Male genitalia (Figs 5 A-5C). Median lobe (ml) with basal lobe (bl) about half length of shaft (sh), basal opening (bo) large. Shaft slender, curved ventrally, dorsally membranous (om) except for two long sclerotized strips (ss) extended from basal lobe to ostial opening (oo); in ventral aspect tapered toward rather broadly rounded apex, preapically with prominent preapical ridge (par), in lateral aspect, ridge a prominently projected point. Parameres (lp, rp) broad, apices subtruncate left paramere (lp) longer than right paramere (rp) about three quarters length of shaft (measured in left lateral aspect). Internal sac with apical ring of rather densely distributed microtrichia, without preapical spines.
Classification. Ball (1959: 194-195) arranged the species of subgenus Badister in two "complexes" characterized principally by features of the male genitalia and ovipositor gonocoxite 2. The combination of these features in B. amazonus fits neither complex. Accordingly, we propose here the monobasic amazonus complex, characterized as follows: median lobe of male genitalia dorsally with two long sclerotized strips, internal sac without sclerotized plates or spines; gonocoxite 2 of ovipositor relatively broad basally, ensiform setae relatively short; nematiform setae absent. Additional diagnostic features are: male fore tarsomeres 1-3 (Fig. 3B) relatively slender, not much wider than tarsomeres 1-3 of middle and hind tarsi, fore tarsomere 1 longer than wide, ventral adhesive setae (Fig. 3C) concentrated in anterior half of each tarsomere.
Dispersal potential. These beetles are fully macropterous and are probably capable of flight; they are fast walkers. They also are climbers on grasses (Paspalum spp.).
Way of life. Adults are associated with water: in open grassy marshes, on beaches of black and white water rivers, and in forest swamps at the edges of standing black water ponds in leaf litter on grey clay soil (Kaolin). They are active at night and take cover in the day among grass stems and under flood debris. They are active in May and August-September; no teneral adults were encountered in these months.
Other specimens examined. Geographic distribution. (Fig. 7). This species is currently known from only three localities: two along the Río Samiria, and one along the Río Ucayali, Loreto Department, Perú.
Notes. We are not presenting an analysis of variation in body proportions because this species can be separated from all other members of the genus without resorting to such attributes. In regard to the location (latitude/longitude) above for "1.0 km SW del boca Rio Samiria," Google Earth allows us now to refine what is given on the labels. The actual site is 4.686779° S, 74.336711° W at an elevation of 101m. Also, the female specimen from the Río Ucayali was mistakenly labeled "Río Amazonas." The river changes names in the vicinity of Iquitos, Perú.

Concluding statement
Although we have demonstrated that the genus Badister in the Western Hemisphere is not amphi-tropical, we are left with many questions. Is it not anomalous that a distinct species, such as Badister amazonus, should be found so distant geographically from its taxonomically close relatives in southern México, and yet be so structurally similar to them? One might expect that this seeming isolate would be markedly differentiated within the subgenus in which it has been placed, in other words, some sort of a relict; however, that seems unlikely. Rather, B. amazonus gives the impression of a taxon that is a comparatively recent arrival in South America, and one could expect it to have a geographical range that extends along northern South American waterways, as well as those of the Amazon Basin. On the other hand, B. amazonus could be a species with a restricted distribution that has undiscovered close relatives along these waterways that, collectively, would close the seemingly extensive gap in the overall range of subgenus Badister. Additionally, the occurrence of subgenus Baudia in mid-northern Argentina begs the questions: 1) Given that many species of this subgenus are known to be tropical (from southern México to Costa Rica), why has it not been found in the northern South American tropics as it has in subtropical/ temperate Argentina? 2) Is the Argentine specimen mislabeled? 3) Is it an invasive species?
So, questions are posed that will be answered by hardy field workers prepared to endure the hardships and risk to life and limb that one faces from unwanted encounters with the crocodilian and reptilian denizens inhabiting the South American wetlands, the home grounds of Badister amazonus n. sp. Such exploration is the torch that senior carabidologists pass on to following generations of carabid beetle enthusiasts.
Appendix 1 table 1. Tables with measurements and ratios for Badister amazonus sp. n. corresponding to same in Ball (1959) for all other western hemisphere species of Badister. All measurements in millimeters.