Beetles that live with ants (Coleoptera, Carabidae, Pseudomorphini): A remarkable new genus and species from Guyane (French Guiana), Guyanemorpha spectabilis gen. n., sp. n.

Abstract Among the extensive collections currently being made in Guyane (French Guiana), adults of a large and colorful species of pseudomorphine were encountered. The adults present, for the first time in the Western Hemisphere, elytra with a marked color pattern, and in addition a size considerably beyond that of the rest of the members of all other known genera in the Western Hemisphere. Both of these attributes, however, are well known in the Australian pseudomorphine fauna. This new species is described and illustrated and a revised key to the Western Hemisphere genera is included. The type locality of Guyanemorpha spectabilis gen. n., sp. n. is Guyane,Risquetout, PK20, 4.916°N, 52.516°W, 12m altitude.


Introduction
Surprising taxa of Carabidae continue to surface as collections from remote places and new habitats are explored (e.g., Erwin 2000Erwin , 2004Erwin and Geraci 2008). Another such taxon has been discovered in various parts of Guyane during ongoing biotic inventories in reserved areas of Guyane and exploration of that country's insect biodiversity by The Entomological Society Antilles-Guyane (SEAG) (cf. Erwin et al. 2012). The species is remarkable because the adults ( Fig. 1) present, for the first time in the Western Hemisphere, elytra with a marked color pattern, and in addition a size considerably beyond that of the rest of members of the other known genera in the Western Hemisphere. Both of these attributes, however, are well known in the Australian pseudomorphine fauna (Baehr 1992(Baehr , 1997 and it is likely a greater variety of color forms will be found in the future in South America.

Specimens and methods
Included in this study are a total of 4 specimens from other institutions and private collections (Appendix 1). "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 the abdomen; 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." "Habitus and attribute images of the adult beetles portray most of the character states referred to in the key provided. Male and female genitalic presentations are standard for descriptive taxonomy of carabid beetles, and in this case are digital photoillustrations (Erwin 2011). 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." "Geographical data are presented based on all known specimens of each species 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. 5].
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)," (Erwin and Amundson in press).

Western Hemisphere genera of Pseudomorphini Newman 1842
Guyanemorpha Erwin, gen. n. Guyane (French Guiana) Manumorpha Erwin & Geraci, 2008. Brazil, Ecuador, Guyane, Perú Notopseudomorpha (Baehr, 1997 (2008), Pseudomorphini appears as a branch of the higher Carabidae and associated with Graphipterini and Orthogonini; according to Erwin and Geraci (2008), the adelphotaxon is the tribe Orthogonini. All three tribes are associated in some way with ants or termites. Male genitalia of pseudomorphines have a bonnet-shaped phallobase as in the lebiomorphs, yet their accompanying parameres are large and nearly symmetrical (and in some species the parameres are sparsely setiferous), as in some primitive lineages of the family. Many known lineages of Pseudomorphini have been so highly selected for life with ants (and possibly termites) that external structures do not help much in discovering more normal carabid relatives (cf. Erwin and Amundson, in press).
Pterothorax. Metepisternum elongate and markedly narrowed posteriorly, the outer margin about 2 times greater in length than the anterior margin, posterior margin about 0.2 times anterior margin.
Legs. Short and depressed, femur posteriorly channeled for reception of tibia in repose; antennal comb notch very shallow; tibial spurs normal; anterior tarsi of male with tarsomeres 2-4 dilated slightly, ventrally each with two laterally placed rows of adhesive articulo-setae.
Abdomen. Abdominal sterna III-VII with patches of short setae and each of IV-VII with a single row of erect ambulatory setae numbering 2 to 8 setae; V and VI in male each with dense row of yellowish robust setae separated medially.
Female ovipositor and genital tract (Fig. 1). Gonocoxite 2 falcate, base about as long as blade, latter relatively short, pointed distally; margins without ensiform setae; with short preapical nematiform seta. Derivation of specific epithet. The epithet "spectabilis" is a Latin adjective describing the very large and colorful beetle species.

Guyanemorpha spectabilis
Proposed english vernacular name. Spectacular Guyane False-form beetle. Diagnosis. With the attributes of the genus as described above and color black and rufous with elytral spots (Fig. 1), color tone of head and pronotum uniform black; form broad and stout with tapered elytra; head with preapical lobe prominent but hidden in dorsal aspect beneath the frons, about 2/3 the length of the anterior margin of eye; pronotum (Fig. 1) coequal at base to elytra across humeri; elytron markedly tapered from humerus to narrower truncated apex and without interneurs or intervals, surface glabrous except parascutellar seta and 8 ombilicate setae near lateral margin.
Description. (Figs 1, 2, 3, 4; Appendix 3). Size: Very large for a Western Hemisphere species, ABL = 13.18 to 13.51 mm, SBL = 11.05 to 12.18 mm, TW = 6.36 to 6.86 mm. Dispersal potential. These beetles are macropterous and have been recorded from flight intercept traps (FITs), hence fully capable of flight; they are likely swift and agile runners as other species in the Tribe. Accordingly, this species may be expected to be more broadly distributed across a wider geographical range than current records indicate.
Way of life. Adults of other pseudomorphines in the Western Hemisphere are found in ant nests and the surrounding vicinity and possibly in termite nests (Ogueta 1967); female adults of species of Pseudomorpha are ovoviviparous (Liebherr and Ka- vanaugh 1985); Pseudomorpha and Notopseudomorpha larvae are ant nest inquilines (Erwin 1981, Lenko 1972. Members of G. spectabilis occur at lowland rainforest sites and most likely live with ants. They have been found in July and December.  Geographic distribution. (Fig. 5). This species is currently known only from Guyane.
Note. The holotype is currently held in trust at the National Museum of Natural History, Smithsonian Institution, Washington, DC until the planned Natural History Museum of Guyane is established, and at that time the specimen will be transferred there. See details of SEAG carabid collections in Erwin et al. 2012.

Concluding statement
Adults of Guyanemorpha share attributes with those of both Notopseudomorpha (Baehr) and Yasunimorpha Erwin & Geraci. This surprising large and colorful pseudomorphine came as a shock to the author, as all other species of the Tribe in the Western Hemisphere are quite dull fulvous, rufous, or black with no, or little, color contrast on the dorsal surface. Since little in the way of life information is available for the four specimens reported herein except seasonality of adult activity and lowland habitation, only speculation based on fogging samples in Perú and Ecuador can add much to this conclusion. "The pseudomorphines are a very interesting evolutionary off-shoot of the normal carabid morphotype in both form and function and are only just now beginning to be understood in North America. The fact that species of related genera in South America are living with arboreal ants will make learning about them far more difficult. Insecticidal fogging gets adults of these species, but only tearing apart arboreal Azteca ant nests while suspended in a tree will produce their larvae, and that is not for carabidologists faint of heart." Erwin and Amundson (in press).