A Review of Gallerucidia ChAudoiR (CARAbidAe: Lebiini: GALLeRuCidiinA): theiR oCCuRRenCes, wAys of Life, And AnotheR Genus ReCoRd foR CostA RiCA

Terry L. erwinhyper-diversity Group, department of entomology, MRC-187, national Museum of natural history, smithsonian institution,

P.o. box 37012, washington, d.C. 20013-7012

AbstRACt

the Lebiini subtribe Gallerucidiina, is composed of two genera; the new world genus Gallerucidia Chaudoir and the old world genus lebidia Morawitz. this paper provides diagnoses of these genera as well as diagnoses, redescriptions, illustrations, and distributional data of the five known species of Gallerucidia. Quite amazingly, the oriental species lebidia bioculata Morawitz has apparently been introduced to venezuela. identification keys are provided to the species of neotropical Gallerucidiina and to the eight subtribes of Lebiini known to occur in Costa Rica.

Key words: Carabidae, Costa Rica, Gallerucidia Chaudoir, lebidia Morawitz, Lebiini, neotropical Realm, oriental Realm

ResuMen

el grupo de Lebiini subtribu Gallerucidiina, está compuesto por dos géneros; el género del nuevo Mundo Gallerucidia Chaudoir y el del viejo Mundo lebidia Morawitz. en este trabajo, proveo diagnosis para estos géneros como también diagnosis, redescripciones, ilustraciones y datos de distribución de las cinco especies conocidas de Gallerucidia. sorprendentemente, la especie oriental lebidia bioculata Morawitz ha sido aparente-mente introducida en venezuela. Claves de identificación son provistas para las especies neotropicales de Gallerucidiina y para las ocho subtribus de Lebiini que se conocen presentes en Costa Rica.

PaLabras CLave: Carabidae, Costa Rica, dominio neotropical, dominio oriental, Gallerucidia Chaudoir, lebidia Morawitz, Lebiini

intRoduCtion

A n n A L s o f C A R n e G i e M u s e u M

voL 77, number 1, PP 135–146 20 JuLy 2008

the lebiine subtribe Gallerucidiina, is composed of two genera, the new world genus Gallerucidia Chaudoir, 1872, and the old world genus lebidia Morawitz, 1862. Many adult attributes support a Gallerucidiina hypothesis, including their general form with constricted neck, broad planular pronotum with broad and shallowly reflexed lat-eral margins, rounded elytral apex, subgranular elytral sur-face, swollen anterior process of metasternum, and medi-ally inflated male phallus. Chaudoir (1872) placed the two genera in his family-group taxon “Gallerucidiae,” here recognized as subtribe Gallerucidiina. bates (1883) pointed out the peculiarly swollen form of the anterior process of the metasternum in members of Gallerucidia (fig. 2d), which he also observed in members of cryptobatis eschscholtz and chelonodema Laporte de Castelnau, somewhat distantly related in our current classification of Lebiini. ball (in litt.) suggested that a phylogenetic analysis of members of chelonodema, Gallerucidiina, and Lebiina (in the historic sense) needs to be made and presumptive relationships re-evaluated. Although it is not within the scope of this paper, i suggest that future assessments of specific characters and their states will provide the basis for morphologically-based phylogenetic studies which will help infer the deep relationships within the Lebi-ini, and test whether or not there is a close relationship among cryptobatis eschscholtz, chelonodema Laporte de Castelnau, and the Gallerucidiina, which in my opin-ion does not seem likely. these attributes include the

distribution of the adhesive setae on male anterior tarsomeres, structure of the anterior process of the metaster-num, concavity of the prosternum, ultimate three umbilicate pores of the elytra, form of the apical margin of the elytra, form of the mandibular scrobe, female reproductive sys-tem, and male genitalia, particularly the folding patterns of the internal sac. of course, molecular data can also contrib-ute to this re-analysis and specimens need to be collected appropriately. Members of Gallerucidia are most likely arboreal and they are probably widely distributed in the neotropical re-gion. However, finding individuals is difficult. Series have been sampled only at black light and most of those seen for this study are females. better knowledge of species dis-tributions will be needed for a worthwhile monograph of Gallerucidia. the purpose of this paper is to provide summary infor-mation about species heretofore unknown from Costa Rica and to provide redescriptions of them, so that this informa-tion can be made available for the national biodiversity inventory Project of Costa Rica, tree of Life, and the en-cyclopedia of Life Projects. this is another in my series of papers with diagnoses of taxa in the beetle family Carabidae from Costa Rica (erwin 2000, 2002, 2004a, 2004b; erwin et al. 2004), ad-joining countries, and in other countries in the neotropics where these beetles live; past contributions can be found in erwin (1973a, 1973b, 1974, 1982, 1991, 1994).

136 ANNALS OF CARNEGIE MUSEUM VOL. 77

SPECIMENS AND METHODS

Methods and species concepts follow those previously described (E1win and Kavanaugh 1981; Kavanaugh and Erwin 1991). Measurements of length (ABL, SBL) and width (TW) follow those of Ball (1972) and Kavanaugh (1979): ABL (apparent body length), measured from apex of labmm to apex of longer elytron; SBL (standardized body length) equals the sum of the lengths of the head (measured from base of mandible to a point on midline at level of the posterior edge of compound eye), pronotum (measured from apical to basal margin along midline), and elytron length (measUl'ed from basal carina to apex of the longer elytron); and TW (total width), measured across left elytron at its widest point and doubled.

The habitus images of the adult beetles portray most of the character states refen·ed to in the keys provided. Male genitalia illustrations are standard for descriptive taxono­my of carabid beetles. Geographical data are presented for the species based on all known specimens available at the time of manuscript preparation. I also provide a composite map of cunent geo-references for all Costa Rican speci­mens herein studied (Fig. 4). Geo-references are given in either latitude-longitude or Lambert Azimuth Equal Area projections. Up-to-date distribution maps for both Costa Rican species of Gallerucidia will occur eventually in the Encyclopedia of Life, Smithsonian Institution website ( <http://www.eol.org/> ).

Common names for species are required in Spanish for the INBio website; hence, they are provided here in English and will be translated into Spanish when posted at the INBio web site. The Gallerucidia species list below, as well as anangement of descriptions that follow are ordered no1ih to south by species occunence. Geo-references have been dete1mined from infonnation given on the speci­men labels. At INBio, Lambert is customarily used, while I use latitude and longitude. I report here all infmmation available and use the customa1y addim "0" for minutes or seconds that are not available. Specimens illustrated are noted in figure captions by their ADP or CRI (Barcode) numbers which in tum link to their geo-reference in the descriptions.

A total of 101 specimens, including type specimens, was examined for this study thanks to the National Mu­seum of Natural Histmy, Washington, DC (NMNH); Museum National d'Histoire Naturelle, Paris (MNHP, Thierry Deuve); INBio, Santa Domingo, Costa Rica, (IN­Bio, Angel Solis); University of Alberta, Edmonton, A1-be1ta (UASM, Danny Shpeley); Califomia Academy of Sciences, San Francisco (CASC, David Kavanaugh); The NatUI'al History Museum, London (BMNH, Peter Ham­mond); and the Canadian National Collection (CNC, Yves Bousquet). Additional specimens were studied from the private collections of Jim Wappes (J EW C) and Robert Tumbow (RTC). All specimens will be retumed upon completion of the study or share-deposited at the NMNH.

SYSTEMATIC ZOOLOGY

Order Coleoptera Linne, 1758 Family Carabidae Latreille, 1802

Tribe Lebiini Bonelli, 1810

KEY TO THE COSTA RICAN SUB TRIBES OF THE TRIBE LEBIINI

(Modified from Ball 1975; Reichardt 1977; Ball and Hilchie 1983; Ball and Shpeley 1983; Erwin 1991; E1win et al. 2004)

Head ventrally without suborbital setigerous pores • ................................................................• 2

1' Head ventrally with at least one pair of suborbital setigerous pores. ........................................... 7

2 (1) Penultimate setigerous pore of elytral umbilicate series displaced laterally ................................. .. .. . . . . . Subtribe APENINA: A penes LeConte, 1851

2' Penultimate setigerous pore of elytral umbili­cate series not displaced laterally, OR displaced medially ..................................................... 3

3(2') Posterior tibial spurs markedly unequal, margins sen·ate, inner spur almost as long as tarsomere 1; neck markedly nanowed, head pedunculate ........ . ................. Subtribe NEMOTARSINA: Nemotarsus

LeConte, 1853

3' Posterior tibial spurs subequal, their margins smooth; neck not markedly nanowed, head not pedunculate ................................................. 4

4 (3 ') Mandible widened near base, scrobe wide, lateral margin markedly rounded ...... Subtribe AGRINA

4' Mandible not conspicuously widened near base, scrobe nan·owed, lateral margin not markedly rounded ...................................................... 5

5(4') Tarsomeres broad, tarsomere 4 with apex subtmn-cate, not bilobed ........................................... . Subtribe CYMINDINA: Pinacodera Schaum, 1857

5' Tarsomeres broad, tarsomere 4 bilobed, OR tar­someres slender and tarsomere 4 with apical mar-gin subtiuncate ............................................ 6

6(5') Tarsomeres stout, dilated, tarsomere 4 bilobed; penultimate setigerous pore of elytral umbilicate series displaced medially ...... Subti'ibe LEBIINA

6' Tarsomeres slender, tarsomere 4 with apical margin subtmncate; penultimate setigerous pore of elyn·al umbilicate se1'ies not displaced laterally, OR dis-placed medially ............... Subti'ibe DROMIINA

7(1 ') Labmm nan·ow, as long or longer than wide;

2008 ERWIN - REVIEW OF GALLERUCIDIA CHAUDOIR 137

7'

penultimate setigerous pore of umbilicate series displaced laterally ......... Subtribe PERICALINA

Labrum wide, wider than long; penultimate se­tigerous pore of umbilicate series not displaced lateran y ..................................................... 8

8(7') Elytron smooth, without visible stiae, elytral sur­face finely and densely punctulate; elytral apex obliquely rounded. Pronotum with sides curved, widest near base, nan-owed evenly anteriorly, apical margin much nan-ower than basal margin. Head sha1ply constricted posteriorly, pedunculate .......................... Subtribe GALLERUCIDIINA

8' Elytron striate; elytral apex truncate. Pronotum with sides sinuate posteriorly, widest at or anterior to middle. Head gradually constricted posteriorly

Subtribe METALLICINA: Euproctinus Leng and Mutchler, 1927

Subtribe Gallerucidiina

Gallerucidiina Chaudior, 1872:416. Type genus: Gallerncidia Chaudoir, 1872. Original designation.

Lebidiina Jacobson, 1907:394. Type genus: Lebidia Morawitz, 1862. Original desgination.

Diagnosis.- (Figs. 1- 3) Head ventrally without suborbital setigerous pores; neck markedly constricted, pedunculate. Labrum wide, wider than long. Pronotum without anterior setigerous pore; pronotum with sides curved, widest near base, nanowed evenly anteriorly, apical margin much nar­rower than basal margin. Elytron smooth, without visible striae, elytral smface finely and densely punctulate; ely­tral apex obliquely rounded; penultimate setigerous pore of umbilicate series not displaced laterally. Anterior pro­cess of metasternum swollen. Claws pectinate. Male with medial v-shaped notch on abdominal sternum VII, female entire.

Notes.- Habu (1967) placed the genera Lebidia and Gallerucidia in the Subtribe Calleidina (= Agrina) based solely on the adhesive ventral vestiture of the fore and mid-tarsi of males. Habu did not mention the attributes of mouthparts observed by Chaudoir and referenced by Bates (although he did provide figures of dorsal and ven­tral aspects), nor the metathoracic attributes mentioned by Bates (1883). The mouthparts are much like those of the calleidines. In addition, Habu did not consider shared at­tributes that are clearly unique in members of these genera amongst Lebiini, those of rounded elytral apicies in adults (as opposed to truncate in all other lebiine members), the elytral sUl'face texture, f01m of the mesosternum and metasternum, and attributes of the male genitalia.

1'

KEY TO THE GENERA OF SUB TRIBE GALLERUCIDIINA

Pronotum broadly transverse, base equally as broad as width across middle; elytral fonn ovoid ............................ Gallerucidia Chaudoir, 1872

Pronotum broadly quadrate, base nan-ower than width across middle; elytral form rectangulate, slightly wider posterior of middle .................... .. ................ .................. .Lebidia Morawitz, 1862

Species of Lebidia Morawitz, 1862

Lebidia bioculata Morawitz, 1863:29. Burma, E. Siberia, Fom1osa, Japan, India, Indonesia, N . China, Venezuela (introduced).

Lebidia bioculata batesi Kano, 1929:523. Japan (this is merely a color variety, however it was described by Kano as a subspecies].

Lebidia dhankutani Kirschenhofer, 1994:101 L NepaL Lebidiaformosana Kano, 1929:524. Formosa. Lebidia octocelis Andrewes, 1924:244. Sikkim. Lebidia octoguttata Morawitz, 1862:323. China, E. Siberia, Formosa,

Japan, Korea.

Lebidia Morawitz, 1862

Lebidia Morawitz 1862:322. Type species: Lebidia octoguttata Morawitz, 1862 :323, by monotypy. Type locality: Hakodate, Hokkaido, Japan.

Diagnosis.-(cf Figs. 1A-1B). Head constricted behind eyes; frontal funow effaced. Pronotum wider than head, somewhat trapezoidal and broadly explanate; margins not beaded; anterior marginal setae absent, posterior setae present at hind angles. Elytron ovoid, smface densely and minutely punctulate; striae and dorsal setae absent; apex rounded. Male with medial notch in sternum VII, female sternum VII entire, both sexes with multiple setae on api­cal edge of sternum VII.

Geographic Distribution.- ASIA: Bm1na, China, E. Si­beria, Fonnosa, Indonesia, India, Japan, Korea. SOUTH AMERICA: Venezuela (introduced, possibly established).

Way of Life.- Adults of species in this genus are arboreal on tree foliage. They hibernate under tree bark. They are occasionally found at lights at night, suggesting they are nocturnal.

Notes.-For detailed descriptions and illustrations of the two known species occm1-ing in Japan, see Habu (1967). One of these, Lebidia bioculata, was apparently intro­duced into Venezuela [VENEZUELA. 6 females, GuARI­co, Hato Masaguaral, 15 km S Calabozo in gallery forest, lOOm, 08° 57'0N, 067°58'0W, June (M. Epstein and M. Deza)(NMNH: ADP109145, ADP109147, ADP109149, ADP109143, ADP109141 , ADP109139)] sometime be­fore 1989 when it was collected there by my colleague Mark Epstein, at black light. According to Habu (1967), the f01m with little or no dark patch within the elytral silvery spot (that variety described by Kano as Lebidia

138 annaLs of Carnegie museum voL. 77

fig. 1.—species of Gallerucidiina. A, lebidia bioculata Morawitz, male, dorsal aspect, specimen # AdP109145, (AbL = 8.92mm, venezuela, see text); B, lebidia octoguttata Morawitz, male, dorsal aspect, specimen # AdP109137, (AbL = 11.04mm, Japan: Gifu Prefecture); C, Gallerucidia erotyloides bates, male, dorsal aspect, specimen # AdP109157 (AbL = 7.39mm); D, Gallerucidia championi bates, male, dorsal aspect, specimen # AdP100381 (AbL = 6.86mm).

2008 ERWIN - REVIEW OF GALLERUCIDIA CHAUDOIR 139

bioculata batesi) is rare; he saw only two specimens, one from Hokkaido and the other from Manchuria. This rare f01m is the one occurring in Venezuela. Of considerable interest is that the collection site in Venezuela of this spe­cies is due north and not far from that of Cymbionotum (Procoscinia) negrei Perrault, 1994, another Old World lineage discovered only recently in South America (cf Ball and Shpeley 2005). In the case of Lebidia bioculata, it is clearly an established introduction, as I have compared both the descriptions in Habu (1967) and a series of 40 Lebidia spp. specimens in the NMNH and found no dif­ferences and little variation, including the male genitalia, from Habu's redescription ofL. bioculata. Two of the Ven­ezuela specimens are vety teneral, hence newly emerged and lending supp01t that the introduction survived at least one generation.

However, in the case of Cymbionotum negrei Penault, it seems that the New World lineage is wotthy of a new subgenus level (Ball and Shpeley 2005) and it has an adel­photaxon in Colombia, Cymbionotum fernandezi Ball and Shpeley; hence the histories of the Lebidia and Cymbiono­tum lineages in South America are quite different.

Iwata (1932) obsetved an adult L. octoguttata feeding upon the lruva of a species of Tortricidae (Lepidoptera). The Asian members of this subtribe are in need of revtston.

Species of Gallerucidia Chaudoir, 1872

Gallenlcidia erotyloidBS Bates, 1883:215, Costa Rica. El Salv-ador, Guatemala. Honduras, Mexico.

Gallerocidia championi Bates, 1883:216, Costa Rica. Guatemala. Mexico, Panama,

Gallerocidia dimidiata Chaudoir, 1872:429, Cuba. Gallerocidia basinotato Chaudoir, 1872:419, Brazil, Peru, Gallerocidia octonotata Chaudoir, 1872:418, Brazil,

Gallerucidia Chaudoir, 1872

Gallerucidia Chaudoir, 1872:417. Type species: Gallerucidia octonotata Chaudoir, 1872:418, first mentioned of three species, designated by Lorenz 1998:89. Type locality: "environs de Rio­Janeiro," Brazil.

Diagnosis.-(cf Figs. 1C- D, 2-3). Color testaceous, or mfous (Cuba), with dark elytral markings. Head with very large hemispheric eyes and constricted neck. Antennom­eres 1-4 glabrous except for apical ring setae, antennom­eres 5- 11 densely pubescent and with apical ring setae. Body convex and ovoid in outline; pronotum broadest at base: elytra rounded at apex. Male with white adhesive setae on anterior tarsomeres 1-3, middle tarsomeres 1- 3; bilobed tarsomere 4 on all legs of both male and female with modified white non-skid setae typical of many arbo­real carabids; females with non-skid setae also on tasom­eres 1-3 of middle and hind legs. Male with medial notch on abdominal stemum VII, female stemum VII entire; both sexes with two pairs of anal setae at apical margin of

stemum VII, plus some very short setae laterad of the pair also on the mru·gin.

Gi>ographic Dish'ibution.- Mexico south to southeastem Brazil, Peru, and on the island of Cuba.

Notes.- Motphologically, these beetles are extremely sim­ilar. They differ in color pattem, body form and little else. Except for one species, these beetles are apparently very difficult to collect, so few have tumed up in collections, particularly in South America. Adults of their adelphotax­on, Lebidia, are arboreal (Habu 1967) yet only two adults of G. octonotata Chaudoir were collected in Pent during my extensive fogging programs there, and none in Ecuador with even more extensive foggings. Ball and Whitehead (specimen label data) found adult G. erotyloides Bates in bromeliads and by beating vegetation in Mexico, and this species, throughout its extensive range, also flies to black light. It is likely that adults are arboreal, but nonnally dwell in the understoxy rather than the high canopy.

KEY TO THE SPECIES OF GALLERUCIDIA CHAUDOIR, 1872

1 Elytron pattemed, with spot(s) and/or vittae ...... 2

1' Elytron concolorous, without spots or vittae (Fig.

2(1)

2 '

3 (2)

3'

4(2')

4'

1 C) .................................. G. erotyloides Bates

Elytron testaceous or ntfous with a large black pre-apical spot (Figs. lD, 2A) ......................... 3

Elytron testaceous witl1 sh01t vittae and/or small spots (Figs. 2B-C). ........................................ .4

Elytron testaceous with large isolated spot, spots combined to fonn a somewhat heatt-shaped pattem; pronotum relatively wide (UW Ratio: 0.58 - 0.64) (Fig. lD) ................. G. championi Bates

Elytron mfous with apical half black; pronotum relatively nan·ow (L/W Ratio: 0.66) (Fig. 2A) ..... . •..........•••....................•.• G. dimidiata Chaudoir

Antennae testaceous, head black; pronotum relatively nru1·ow (L/W Ratio: 0.61 - 0.63); base of elytron broadly piceous or with two broad piceous vittae (Fig. 2B) .... G. basinotata Chaudoir

Antennomeres 5- 11 infuscated, head infuscated; pronotum relatively wide (1JW Ratio: 0.54 - 0.57); base of elytron with two parallel narrow elongate vittae (Fig. 2C) .............. G. octonotata Chaudoir

Gallerucidia erotyloides Bates, 1883 (Figs. 1C, 2D, 3A, 3C, 3D, 4)

Gallerocidia eroryloidBS Bates, 1883:215. Lectotype female labeled by author in BMNH. Type locality: GUATEMALA, San GerOnimo, !150m, 15 08' 0N, 090 !! 'OW. Seven Paralectotypes: five from Oaxaca, Mexico, two from San Geronimo, Guatemala.

140 annaLs of Carnegie museum voL. 77

fig. 2.— species of Gallerucidia. A, Gallerucidia dimidiata Chaudoir, female holotype, dorsal aspect, specimen # AdP109135 (AbL = 6.22mm); B, Gallerucidia basinotata Chaudoir, male, dorsal aspect, specimen # AdP108835 (AbL = 6.78mm); C, Gallerucidia octonotata Chaudoir, female holo-type, dorsal aspect, specimen # AdP108793 (AbL = 7.31mm); D, Gallerucidia erotyloides bates, male, mesothorax and metathorax, oblique ventral aspect, specimen # AdP108816.

A

2008 erwin – review of Gallerucidia Chaudoir 141

Common name.—Mimetic pleasing carabid beetle.

Diagnosis.—(fig. 1C) Completely testaceous; pronotum relatively wide (L/w Ratio: 0.54 – 0.56).

Description.—(figs. 3A, C–d). size medium: AbL = 7.51–7.73mm, sbL = 6.70–7.20mm, tw = 3.80–3.96mm. color.—see diagnosis. luster.—shiny. Microsculpture.—entire dorsal surface with finely engraved isodia-metric meshes. Head.—short; much narrower across eyes than pronotum; frontal fur-rows absent; eyes very large, hemispheric, facets small; frons flat, vertex very slightly convex; ultimate labial palpomere normal; antennae long, reaching well posterior of humerus. Prothorax.—Pronotum transverse, much narrower anteriorly than base, moderately convex; side margins broadly explanate, well rounded from anterior to hind angles; hind angles obtuse with setigerous pore; base slightly lobed posteromedially. Pterothorax.—normal for Lebiini, fully winged. legs.—normal for Lebiini. abdomen.—sterna normal for Lebiini, mostly glabrous, except nor-mal paired ambulatory setae on sterna 4–6; males and females with two pairs medium length setae on sternum 7. Male Genitalia.—Phallus (fig. 3A) robust, slightly swollen at middle, with ostium one-third length of phallus and left pleuropic; phallus apex narrow, elongate and slightly hooked; endophallus microtrichiate, without defined fields. Parameres: left rectangulate, relatively broad; right small, narrow, and bifid at apex. Female reproductive Tract.— bursa copulatrix simple, elongate with scattered microtrichia on the anterior two-thirds (fig. 3C2); spermatheca markedly elongate, annulate throughout; spermathecal duct short, sper-mathecal gland elongate and filiform, base located at base of spermatheca, both on a short stalk attached to dorsal side of bursa copulatrix (fig. 3C2); stalk with small ventral lobe (fig. 3C1). basal gonocoxite glabrous, robust; apical gonocoxite short, broadly rounded at apex, armed with a ventral pad of ensiform setae (fig. 3d).

Dispersal Potential.—these beetles are fully winged and likely are good dispersers. the specimen from honduras was collected at black light and those from México from arboreal bromeliads.

Way of Life.—Arboreal, at mid altitudes, often in brome-liads or on understory vegetation. bates (1883) remarked that these beetles resemble adults in the genus Omoiotelus(erotylidae).

Other Specimens Examined.—COSTA RICA. 1 female, guanaCasTe, P.n. guanaCasTe, sw slope volcán Cacao, 1000–1400m, Ln323300,375700, May (M. Reyes)(inbio: CRi000413627); 1 female, estación Pitilla, 9 km s santa Cecilia, 700m, 10 59’33”n, 085 25’46”w, Ln330200,380200, March (P. Rios, C. Moraga, and R. blanco)(inbio: CRi000211397); 1 female, finca Jenny, 30 km n Liberia, 200m, Ln316200,364400, May (e. Araya)(inbio: CRi001183554). EL SALVADOR. 1 female, monTe CrisTo, 23 km n Metapan, 2300m, May (h. and A. howden)(uAsM: AdP054705). GUATEMALA. 1 female, QuiChe, 10 km se La Primavera, 1500m, 10 59’33”n, 085 25’46’ w, April (J.M. Campbell)(CnC: AdP057486); 1 female, saCaTePeQuez, Paramos, 1524m, 10 59’33”n, 085 25’46” w, June (J.M. and b.A. Campbell)(CnC: AdP059085). HONDURAS. 1 male, Comayagua, Rancho Chiquito, Km 62, 600m, 14 04’0n, 087 36’0w, May (f.s. blanton, A.b. broce and R.e. woodruff)(uAsM: AdP043586); 1 male, franCisCo morazán, Lizapa, 10 km s Zamorano, 900m, 14 01’0n, 086 22’0w, June (h. and A. howden)(uAsM: AdP109185); 1 female, Zamorano, 30 km. ese tegucigalpa, 830m, 14 04’0n, 086 22’0w, June (h. and A. howden)(uAsM: AdP109183); 1 female, 6 km se

Zamorano, in thorn scrub, 850m, 14 02’0n, 086 20’0w, May (h. and A. howden)(uAsM: AdP109171); 2 females, Zamorano, 431m, 14 04’0n, 086 22’0 w, July (R. turnbow)(RtC: AdP109183, AdP108832), 2 females, May (RtC: AdP108830, AdP108828), 1 female, 1 male, June (RtC: AdP108826, female, AdP108824, male), 1 male, october (RtC: AdP108822); 1 male, 1 female, yoro, Parque nacional Pico Pijol, ca. 1000m, 15 05’0n, 087 28’0w, June, (R. turnbow)(RtC: AdP109181, male, AdP109169, female); 1 male, intibuca, 7.0 km w san Juan, 1687m, 14 19’0n, 088 10’0w, october (R. turnbow)(RtC: AdP108820); 3 females, el Paraiso, 35 km w danli, 686m, 14 25’0n, 087 35’0w, May (R. turnbow)(RtC: AdP108818, AdP108816, AdP108814). MÉXICO. 1 female, ChiaPas, hwy 190, 52.3 km e Comitan, 670m, roadside; sweeping bushes, 16 15’0n, 092 11’0w, september (G.e. ball and d.R. whitehead)(uAsM: AdP109167); 1 male, nuevo Leon, hwy 60, 23.8 km w Linares, 730m, 24 49’0n, 099 35’0w, october (G.e. ball and d.R. whitehead)(uAsM: AdP109165); oaxaCa, 1 male, hwy 131, 70 km s oaxaca, Rio de la y, Km 20, w of jct., 2150m, 16 30’0n, 097 0w, June (h. and A. howden)(uAsM: AdP109179); 1 male, 1 female, 24.8 km s. Juchatengo, 1707m, 16 15’0n, 097 17’0w, ex. bromeliads, March (G.e. ball and d.R. whitehead)(uAsM: AdP109177, male, AdP109163, female); 1 sex unknown, 33 km nw oaxaca City, 1676m, 17 15’0n, 096 54’0w, dry oak savanna, August (R.s. Anderson and w. Maddison) (uAsM: AdP109161); 1 male, sinaLoa, hwy 40, 40.4 mi. n. Concordia, 1524m, 23 25’0n, 105 45’0w, in bromeliads, January (d.R. whitehead)(uAsM: AdP109159); 1 female, sonora, sierra Alamos, 2.7 km. s, 1.9 km w Alamos, 518m, 27 02’0n, 108 55’0w, thorn forest, uv light, July (s. McCleve)(uAsM: AdP109175); 1 male, 1 female, méxiCo, tejupilco, temescaltepec, 1294m, 18 15’0n, 100 12’0w, July (h. e. hinton)(CAs: AdP100391, male, AdP100389, female); 2 males, 2 females, veraCruz, hwy 143, 18.8 km ne huatusco, el Mirador, 1850m, 19 16’0n, 096 50’0w, ravine, coffee finca, ex. bromeliads, december (G.e. and K.e. ball)(uAsM: AdP109151, AdP109153, males, AdP109155, AdP109157, females).

Geographic Distribution.—Known from Costa Rica (fig. 4), and el salvador, Guatemala, honduras, México.

Notes.—bates (1883:215) regarded the labels on bMnh specimens collected by höge as coming from Jalapa to be in error; bates believed that the specimens emanated from oaxaca.

Gallerucidia championi bates, 1883(figs. 1d, 4)

Gallerucidia championi bates, 1883:216. Lectotype female labeled by author in bMnh. type locality: GuAteMALA, Aceituno, 1500m, 14 37’0n, 09 28’0w.

Common name.—Champion’s pleasing carabid beetle.

Diagnosis.—(fig. 1d) Completely testaceous except for black heart-shaped spot on the posterior half of the elytra; pronotum relatively wide (L/w Ratio: 0.58–0.64).

Description.—(fig. 1d). size medium: AbL = 7.12–7.43mm, sbL = 6.69–6.88mm, tw = 3.52–3.84mm. color.—see diagnosis. luster.—shiny. Microsculpture.—entire dorsal surface with finely engraved isodia-metric meshes. Head.—short; much narrower across eyes than pronotum; frontal furrows absent; eyes very large, hemispheric, facets small; frons flat, ver-tex very slightly convex; ultimate labial palpomere slightly securiform; antennae long, reaching well posterior of humerus. Prothorax.—Pronotum transverse, much narrower anteriorly than at

142 annaLs of Carnegie museum voL. 77

fig. 3.—Genitalia of Gallerucidia. A, phallus, dorsal, ventral, left lateral aspects of Gallerucidia erotyloides bates, specimen # AdP043586; B, phallus, dorsal, ventral, left lateral aspects of Gallerucidia basinotata Chaudoir, specimen # AdP108835; C, female reproductive system of Gallerucidia erotyloides bates, specimen # AdP109171; C1, right oblique aspect of spermathecal duct stalk (stk); C2, dorsal aspect of bursa copulatrix (bc) and spermathecal ap-paratus (co, common oviduct; sp1, spermatheca; bc, bursa copulatrix; gc, gonocoxites; sg, spermathecal gland); D, female gonocoxites, ventral aspect, of Gallerucidia erotyloides bates, specimen # AdP109171 (gc1, basal gonocoxite; gc2, apical gonocoxite; ves, ventral ensiform setal pad).

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2008 erwin – review of Gallerucidia Chaudoir 143

base, moderately convex; side margins broadly explanate, well rounded from anterior to hind angles, hind angles; obtuse with setigerous pore; base slightly lobed posteromedially. Pterothorax.—normal for Lebiini, fully winged. legs.—normal for Lebiini. abdomen.—sterna normal for Lebiini, mostly glabrous, except normal paired ambulatory setae on sterna 4–6; females with two pairs medium length setae on sternum 7. Male.—unknown. Female reproductive Tract.—not studied.

Dispersal potential.—these beetles are fully winged and likely are good dispersers.

Way of life.—these beetles are likely arboreal as other species in the genus. Members of this species occur at alti-tudes between 600 and 1400 meters.

Other Specimens Examined.—COSTA RICA. 1 female, CarTago, iiCA experimental station – CAtie, 600m, 09 53’0n, 083 39’0w, Ln574947,208307, May (ekis)(nMnh: AdP 046994); 1 female, Monumento nacional Guayabo, A.C.A.C. Amistad, 1100m, Ln570000,217400, July (G. fonseca, #3126)(inbio: CRi001889573). HONDURAS. 1 female, oLanCho, Parque nacíonal La Muralla, 880m, 15 41’0n, 086 47’0w, June (R. turnbow)(RtC: AdP108794); 1 female, ComayaQue, 2.8 km nne Los Planes, 1799m, 14 04’0n, 088 01’0w, May (R. turnbow)(RtC: AdP108796). MÉXICO. 1 female, san Luis PoTosí, Las Abritas area on hwy 80, 742m, 22 30’0n, 099 24’0w, June (J. wappes)(JewC: AdP100379). PANAMÁ. 1 female, ChiriQui, Las Lagunas, 4 km w hato del volcan, 1360m, 08 05’0n, 081 37’0w May (h. and A. howden)(uAsM: AdP085298); 1 female, 1400m May (G. ekis)(nMnh: AdP022458); 1 female, 20.5 km nw volcan, 1378m, 09 00’0n, 082 35’0w, May (R. turnbow)(RtC: AdP108834).

Geographic Distribution.—Known from Costa Rica (fig. 4), and Guatemala, México, and Panamá.

Gallerucidia dimidiata Chaudoir, 1872(fig. 2A)

Gallerucidia dimidiata Chaudoir, 1872:420. holotype female in MnhP. type area: CubA AdP109135.

Common name.—Cuban pleasing carabid beetle.

Diagnosis.—(fig. 2A) Completely rufotestaceous except apical half of elytron which is black; pronotum relatively wide (L/w Ratio: 0.66).

Description.—(fig. 2A). size medium: AbL = 6.22mm, sbL = 5.73mm, tw = 3.4mm. color.—see diagnosis. luster.—shiny, slight iridescence in black portion of elytra. Microsculpture.—entire dorsal surface with finely engraved isodia-metric meshes, those of the elytra slightly stretched transversely. Head.—short; much narrower across eyes than pronotum; frontal fur-rows absent; eyes very large, hemispheric, facets small; labrum slightly emarginated; frons flat; vertex very slightly convex; ultimate labial palpomere slightly securiform; antennae long, reaching well posterior of humerus. Prothorax.—Pronotum transverse, much narrower anteriorly than at base, moderately convex; side margins broadly explanate, well rounded from anterior to hind angles; hind angles obtuse with setigerous pore; base slightly lobed posteromedially. Pterothorax.—normal for Lebiini, fully winged. legs.—normal for Lebiini.

abdomen.—sterna normal for Lebiini, mostly glabrous, except normal paired ambulatory setae on sterna 4–6; females with two pairs medium length setae on sternum 7. Male genitalia: unknown. Female reproductive Tract.—not studied.

Dispersal potential.—these beetles are fully winged and likely are good dispersers.

Way of Life.—unknown. if this species is like others in the genus, it likely occurs at mid altitudes and such locali-ties have been closed to collecting for decades, hence no new specimens are available.

Other Specimens Examined.—none.

Geographic distribution.—Known only from the holo-type from Cuba (no details on locality or way of life are given on the labels).

Gallerucidia basinotata Chaudoir, 1872(figs. 2b, 3b)

Gallerucidia basinotata Chaudoir, 1872:419. holotype male in MnhP. type locality: brazil, ega (haut-Amazone), now called tefé, 43m, 03 22’0n, 064 42’0w.

Common name.—basally-spotted pleasing carabid beetle.

Diagnosis.—(fig. 2b). Antennae testaceous, head black, base of elytron broadly piceous or with two broad, short piceous vittae; pronotum relatively narrow (L/w Ratio: 0.61–0.63).

Description.—(figs. 2b, 3b). size medium: AbL = 6.70–7.20mm, sbL = 6.31–6.88mm, tw = 3.12–3.70mm.color.—see diagnosis, otherwise completely testaceous. luster.—very shiny. Microsculpture.—entire dorsal surface with finely engraved, shallow isodiametric meshes; those of the elytra largely effaced. Head.—short; much narrower across eyes than pronotum; frontal fur-rows absent; eyes very large, hemispheric, facets small; labrum slightly rounded; frons flat; vertex very slightly convex; ultimate labial palpomere slightly securiform; antennae long, reaching well posterior of humerus. Prothorax.—Pronotum transverse, much narrower anteriorly than at base, moderately convex; side margins broadly explanate, well rounded from anterior to hind angles; hind angles obtuse with setigerous pore; base slightly lobed posteromedially. Pterothorax.—normal for Lebiini, fully winged. legs.—normal for Lebiini. abdomen.—sterna normal for Lebiini, mostly glabrous, except normal paired ambulatory setae on sterna 4–6; females with two pairs medium length setae on sternum 7. Male genitalia.—Phallus (fig. 3b) markedly robust, swollen at middle, with ostium one-third length of phallus and left pleuropic; phal-lus apex narrow, short and markeldy hooked; endophallus microtricheate, without defined fields. Parameres: left rectangulate, moderately broad; right small, narrow, not bifid at apex. Female reproductive Tract.—not studied, see Gallerucidia eroty-loides bates.

Dispersal Potential.—these beetles are fully winged and likely are good dispersers, as are adults of most aboreal species.

144 annaLs of Carnegie museum voL. 77

Way of Life.—on the western side of the Amazon for-est these beetles are arboreal in lowland rainforest and ac-tive in the dry season. the two specimens from Perú were fogged from the high canopy, one from two intertwined trees, Matisia sp. and Hirtella sp., and the other from Pouteria sp.

Other Specimens Examined.—A second male specimen collected by bates at the type locality. PeRÚ. madre de dios, Pakitza, 320m, bioLAt Zone 2, 12 07’0s, 070 58’0w, 1 female, february, (t.L. erwin and b.d. farrell)(nMnh: AdP100375), 1 female, september (nMnh: AdP100377).

Geographic Distribution.—Known from the western Amazon basin.

Notes.—Chaudoir based his description on a single male from bates’ collecting efforts at ega, brazil. bates collect-ed a second specimen at ega, apparently later, that Chau-doir did not see at the time of his description.

Gallerucidia octonotata Chaudoir, 1872(fig. 2C)

Gallerucidia octonotata Chaudoir, 1872:418. holotype female in MnhP. type locality: brazil, “environs de Rio-Janeiro” AdP108793.

Common name.—eight-spotted pleasing carabid beetle.

Diagnosis.—(fig. 2C) head lightly infuscated; antenno-meres 5–11 infuscated; ventrally, mouth parts testaceous

fig. 4.—Geographical distribution map of the species of Gallerucidia in Costa Rica: Gallerucidia championi bates (open circle, ), Gallerucidia ero-tyloides bates (star, ).

2008 erwin – review of Gallerucidia Chaudoir 145

including gula; elytron with two basal narrow vittae and two subapical spots; mesosternum and metasternum infus-cated laterally; pronotum relatively wide (L/w Ratio: 0.54 –0.57).

Description.—(fig. 2C). size medium: AbL = 7.07–7.32mm, sbL = 6.79–6.80mm, tw = 3.66–3.74mm. color.—see diagnosis, otherwise completely testaceous. luster.—shiny. Microsculpture.—entire dorsal surface with finely engraved isodia-metric meshes. Head.—short; much narrower across eyes than pronotum; frontal furrows absent; eyes very large, hemispheric, facets small; frons flat; ver-tex very slightly convex; ultimate labial palpomere slightly securiform; antennae long, reaching well posterior of humerus. Prothorax.—Pronotum transverse, much narrower anteriorly than at base, moderately convex; side margins broadly explanate, well rounded from anterior to hind angles; hind angles obtuse with setigerous pore; base slightly lobed posteromedially. Pterothorax.—normal for Lebiini, fully winged. legs.—normal for Lebiini. abdomen.—sterna normal for Lebiini, mostly glabrous, except normal paired ambulatory setae on sterna 4–6; females with two pairs medium length setae on sternum 7. Male genitalia.—unknown. FemalereproductiveTract.—not studied.

Dispersal Potential.—these beetles are fully winged and likely are good dispersers.

Way of Life.—the two known specimens from mid-altitude forests in the south Atlantic area of brazil (Mata Atlântica) lived in a habitat that has been severely reduced or converted over the past 500 years to nearly 4% of what it was. whether this beetle species persists in the remain-ing fragments is questionable.

Other Specimens Examined.—bRAZiL. – 1 female, rio de Janeiro, Petropolis, 803m, 22º48’0s, 043º14’0w (sahlberg) (MnhP).

Geographic Distribution.—southeastern brazil.

disCussion

further studies on this interesting lineage of Carabidae must await more specimens from more places. the known species seem to be arboreal, but not in the high canopy and probably not in the middle canopy either, or they would have turned up in greater numbers in my decades-long fog-ging programs in the western Amazon basin. the other lebiine lineages with the peculiar form of the metaster-num are also found in the understory rather than the high canopy (i.e., members of cryptobatis and chelonodema); hence a common morphological feature shared among un-related lineages may be a clue to a common way of life. Like members of Gallerucidia, those of cryptobatis and chelonodema are rarely found in the fogging samples. Like many “rare” lineages of lebiines, members of Gal-lerucidia do come to lights, as evidenced by series from México and Costa Rica. ball and his colleagues have taken

a good number in bromeliads in Mexico; so in bromeliads is where collecting needs to be done in the Mata Atlântica of brazil and at middle altitude on the eastern slopes of the Andes. Curiously, all species of Gallerucidia are known mostly from females, not usual in carabid beetle species. A secondary project might be determining whether leb-idia bioculata has become established in venezuela and tracing the history of how it became introduced there.

ACKnowLedGMents

first and foremost, deeply appreciated thanks go to George e. ball for my apprenticeship into the world of carabid beetles and his continuing role as my mentor and amigo; to him this paper is dedicated. And thanks also to his colleague and fellow carabidologist, danny shpeley, whose help with the Méxican collection (uAsM) contributed significantly to this paper. special thanks also go to vichai Malikul who prepared the male genitalia illustrations, and young sohn who prepared those of the female; to warren e. steiner who provided the extended focus images and Microptics images for this paper, as well as organization of the materials for illustration; to stephen McJonathan of Gt vision for the loan of equipment and software necessary to make the extended focus images; and to the curators mentioned above under Methods for loan of specimens. thanks also to valeria Aschero who prepared the spanish translation of the Abstract, and to two anonymous reviewers who really improved many aspects of the presentation. the smithsonian’s national Museum of natural history provided funding for the project.

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