Bulletin Carnegie 38 FRANIA

Taxonomy and Evolution of Species of the Genus Euchroa Brullé (Subgenus Dyschromus Chaudoir) of Central Mexico and the Island of Hispaniola (Coleoptera: Carabidae: Pterostichini: Euchroina)

Authors: Frania, Henry E., and Ball, George E. Source: Bulletin of Carnegie Museum of Natural History, 2007(38) : 1- 125 Published By: Carnegie Museum of Natural History URL: https://doi.org/10.2992/0145-9058(2007)38[1:TAEOSO]2.0.CO;2

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TAXONOMY AND EVOLUTION OF SPECIES OF THE GENUS EUCHROA BRULLÉ (SUBGENUS DYSCHROMUS CHAUDOIR) OF CENTRAL MEXICO AND THE ISLAND OF HISPANIOLA (COLEOPTERA: CARABIDAE: PTEROSTICHINI: EUCHROINA)

HENRY E. FRANIA and GEORGE E. BALL

NUMBER 38 PITTSBURGH, 2007

Provided are a key to, and descriptions of, adults of 35 species of Dyschromus Chaudoir, which is treated as a subgenus of Euchroa Brullé. Twenty-eight species are from central Mexico; seven are from the island of Hispaniola. Twenty-five species are new (type- localities in parentheses): Euchroa independencia n. sp. (DOMINICAN REPUBLIC, Independencia, 5.5 km. n.n.w. Angel Feliz), Euchroa pedernales n. sp. (DOMINICAN REPUBLIC, Pedernales, 1 km. s. Los Arroyos), Euchroa huautla n. sp. (MEXICO, Oaxaca, Rio Santiago, 11.7 km. e. Huautla de Jiminez), Euchroa cuiyachapa n. sp. (MEXICO, Veracruz, between Cuiyachapa and Ixtapa), Euchroa zongolica n. sp. (MEXICO, Veracruz, 5 km. s.w. San Andres de Tenejapa), Euchroa lasvigas n. sp. (MEXICO, Veracruz, 15.7 km. e. Las Vigas), Euchroa teotitlan n. sp. (MEXICO, Oaxaca, 24.5 km. e. Teotitlan Puerto de Soledad), Euchroa citlaltepetl n. sp. (MEXICO, Veracruz, Volcan Citlaltepetl, east slope), Euchroa perote n. sp. (MEXICO, Veracruz, 16 km. s. Las Vigas), Euchroa harrisoni n. sp. (MEXICO, Tamaulipas, 12.8 km. n.w. Gomez Farias), Euchroa soladevega n. sp. (MEXICO, Oaxaca, 14.9 km. n. Sola de Vega), Euchroa juchatengo n. sp. (MEXICO, Oaxaca, 32 km. s. Juchatengo), Euchroa nizavaguiti n. sp. (MEXICO, Oaxaca, n.w. of Santa Maria Nizavaguiti), Euchroa jalisco n. sp. (MEXICO, Jalisco, ca. 19.8 km. s. Tecalitlan), Euchroa tenancingo n. sp. (MEXICO, Mexico, Rio de Molino, near Valle de Bravo), Euchroa ixtapa n. sp. (MEXICO, Guerrero, crest of Sierra Madre del Sur, n.e. of Ixtapa), Euchroa atoyac n. sp. (MEXICO, Guerrero, 66.4 km. n.e. Atoyac de Alvarez), Euchroa puertogallo n. sp. (MEXICO, Guerrero, 32 km. n.w. Filo de Caballo), Euchroa filodecaballo n. sp. (MEXICO, Guerrero, 39.4 km. n.w. Filo de Caballo), Euchroa yucuyacua n. sp. (MEXICO, Oaxaca, Cerro Yucuyacua, summit e. of Nundaco), Euchroa santacatarina n. sp. (MEXICO, Oaxaca, 35.5 km. s. Juchatengo), Euchroa zempoaltepetl n. sp. (MEXICO, Oaxaca, 10.5 km. s. San Pedro y San Pablo Ayutla), Euchroa carbonera n. sp. (MEXICO, Oaxaca, 7.9 km. n.w. La Carbonera), Euchroa miahuatlan n. sp. (MEXICO, Oaxaca, 27.2 km. s. Miahuatlan), and Euchroa suchixtepec n. sp. (MEXICO, Oaxaca, 41.8 km. s. Suchixtepec). Larvae are also described of E. perote, E. harrisoni, E. soladevega, Euchroa dimidiata Chaudoir, E. atoyac, and E. filodecaballo.

Species of the subgenus Dyschromus live on the ground under cover in montane forests of tropical to temperate aspect, at elevations of 900 to 3400 meters. Adults are flightless, and potentially long-lived. Adults of most species are metallic colored dorsally.

All seven species from the island of Hispaniola are placed in the opaca group, and all 28 species from Mexico are placed in the nitidipennis group. The opaca group is divided into the opaca and cupripennis subgroups, but support for monophyly of the opaca subgroup is inconclusive.fThe nitidipennis group is divided into the nitidipennis, soladevega, and dimidiata subgroups. The nitidipennis subgroup contains all species of the subgenus Dyschromus in the mountains of eastern Mexico, north of the Rio Santo Domingo; the dimidiata subgroup comprises most of the other species in Mexico, except for three species from western Oaxaca that make up the soladevega subgroup. Body color and the condition of the elytral striae and microsculpture of adults, and some traits of the larvae, as well as certain ecological and distributional correlates, provide evidence that each of the three subgroups is monophyletic, but structure of the distal part of the median lobe of the aedeagus indicates that four or five members of the dimidiata subgroup from Oaxaca, including the only species from eastern Oaxaca, as a group, actually belong to the nitidipennis subgroup, as does one of the three species in the soladevega subgroup, while the other two species in the soladevega subgroup really belong to the dimidiata subgroup.

Just one species in Mexico is comparatively widespread, E. dimidiata, which inhabits dry oak woodlands that fringe the Rio Balsas Basin and Valley of Oaxaca. The other species are concentrated in two regions of Mexico: the mountains of eastern Mexico, north of the Rio Santo Domingo with 11 species; and the Sierra Madre del Sur and adjoining uplands in western Oaxaca and Guerrero with another 13 species. Most species are known only from a single mountain range. Some mountains have as many as five species, but sympatry is rare; instead most species are found at different elevations and/or in different types of forest. Outside of the above two areas, we have, besides E. dimidiata, one species each from the western and central parts of the Trans-Volcanic Sierra, and one from Cerro Zempoaltepetl in eastern Oaxaca. No species of the subgenus Dyschromus are known from the Sierra Madre Occidental or south of the Isthmus of Tehuantepec. All data indicates a long period of isolation and diversification of the nitidipennis subgroup in the humid forests of the mountains of eastern Mexico, north of the Rio Santo Domingo. Much of the diversification of the subgenus elsewhere in Mexico has been in drier forest formations, although some speciation in the dimidiata subgroup has taken place in the humid forests of the Sierra de Atoyac in Guerrero, and the Sierra de Miahuatlan in Oaxaca. Each of the principal mountain regions of Hispaniola has at least one, and at most two species of Dyschromus. Only certain of the more derived species of the cupripennis subgroup inhabit the northern part of the island.

The subgenus Dyschromus as a whole exhibits a relictual distribution and is an old (early Tertiary) South American component of the fauna of Mexico. Most of the humid forest elements of the subgenus in Mexico are relicts dating from the Miocene epoch. The most widespread lineages of Dyschromus, both in Mexico and on Hispaniola, are phyletically derived, and have a metallic green head and pronotum and brassy to coppery-colored elytra; and at least those in Mexico inhabit comparatively dry oak, or oak-pine forests. Whether there is an advantage for species of Dyschromus living in such conditions to have these colors is unknown.

INTRODUCTION Why are they all brachypterous? Are these beetles really restricted to central Mexico and the island of Hispaniola? In the first publication (1835) of the Baron Maximilien de How are Euchroa and Dyschromus related to one another? Chaudoir, the first taxon described was the genus To try and answer these questions, we resolved to Dyschromus (Frontispiece and Fig. 1), erected to accom- revise the group. Experience had shown us that individu- modate D. opacus, a new species of flightless pterosti- als of Dyschromus are usually difficult to find, so much so chine ground beetle, supposedly from the vicinity of Java, that collecting these beetles at a new site (or even at a but really from Haiti. Chaudoir chose those names pre- known locality) was a cause for celebration, and so it was sumably because the adult of this species is dull black in that only in 1997 did we examine some specimens bor- color (Fig. 2a). Over the next 100 years, nine more species rowed that year from the Laboratorio de Parasitologica were added to the genus (either directly or moved from the Vegetal, Universidad Autonoma del Estados de Morelos, genus Euchroa Brullé), all of them brachypterous, but only that enabled us to identify the female specimen that we had one as nondescript in color as D. opacus. At one extreme, collected in 1965 at Cuernavaca. Most other insights about four species were described with a bright metallic green Dyschromus (treated here as a subgenus of Euchroa) also head and pronotum, and coppery or brassy colored elytra came painfully slowly, and in our search for characters to (as in Frontispiece and Figs. 3c–d): Euchroa cupripennis establish a hypothesis of phylogeny, we eventually reared Chaudoir and Dyschromus perezi Darlington, both from larvae of some of the species, and Felix A.H. Sperling the island of Hispaniola, and Euchroa dimidiata Chaudoir sequenced a fragment of their mDNA (Sperling, Frania and Euchroa flohri Bates, from the Sierra Mixteca Alta and Ball, in prep.). Our knowledge of this group of beetles and the eastern part of the Trans-Volcanic Sierra of is still very incomplete, and in some ways all that we have Mexico, respectively. At the other extreme was a new been able to do is confirm the anomalous nature of the species from Mexico with a metallic bluish-purple head group. What we do now know about Dyschromus is the and pronotum and reddish-purple elytra (as in Fig. 2e): subject of this monograph. Euchroa nitidipennis Putzeys from the “Cordilière orientale d'Oaxaca.” Its geographical distribution proved to be another peculiarity of the genus; each of the ten species MATERIALS AND METHODS was known only from either the island of Hispaniola or central Mexico. As noted above, some of these species had Materials originally been described in the genus Euchroa Brullé, but This monograph is based on study of 822 adult specimens Euchroa, although always regarded as being closely relat- representing 35 species belonging to the subgenus ed to Dyschromus, came to include only a few species Dyschromus, more than half of which we collected our- from southeastern Brazil (Fig. 4). selves. We also reared and examined 189 first, second or One of us (Ball), first encountered living adults of the last instar larvae of six of the species. Additional adult subgenus Dyschromus in 1962, in oak-pine forest in the specimens were borrowed from the collections of various central part of the Trans-Volcanic Sierra of Mexico near individuals and institutions: Valle de Bravo. These individuals (one male and one female) had a metallic green head and pronotum and BMNH—Department of Entomology, The Natural History Museum, brassy colored elytra. In 1965, we (Ball), together with the Cromwell Road, London SW7 5BD, United Kingdom (N.E. Stork and late Donald R. Whitehead, collected a similarly colored M.J.D. Brendell); CASC—Department of Entomology, California female in oak-palmetto scrub further to the east at Academy of Sciences, Golden Gate Park, San Francisco, California Cuernavaca. Nine years later, the other of us (Frania) also 94118, USA (D.H. Kavanaugh); CMNC—Entomology Division, Canada Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4, had the privilege of seeing live adults of Dyschromus, this Canada (R.S. Anderson); CMNH—Section of Invertebrate Zoology, time in pine forest in the eastern part of the Trans-Volcanic Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Sierra, north of Tlaxco. That species had a metallic bluish- Pennsylvania 15213-4080, USA. (J.E. Rawlins, R.L. Davidson); purple head and pronotum, and reddish-purple elytra. By CNCI—Canadian National Collection of Insects, Eastern Cereal and Oilseed Research Centre, Ottawa, Ontario K1A 0C6, Canada (Y. then, we (Ball) and associates had collected from various Bousquet); CSIRO—Division of Entomology, GPO Box 1700, Canberra montane regions of Mexico north of Tehuantepec, speci- ACT 2601, Australia (B.P. Moore); CUIC—Department of Entomology, mens of a number of species of Dyschromus, all Comstock Hall, Cornell University, Ithaca, New York 14853-0999, USA brachypterous, and some most certainly undescribed. All (J.K. Liebherr); EMEC—Essig Museum of Entomology, University of of the specimens were found, like typical pterostichine California, 201 Wellman Hall, MC3112, Berkeley, California 94720- 3112, USA (K.W. Will); FFPC—Foster F. Purrington Collection, carabids, under cover on the ground. Department of Entomology, Ohio State University, Columbus, Ohio Having seen these beetles in the field, and found out 43210-1220, USA; FSCA—Florida State Collection of Arthropods, 1911 what was already known about them, some obvious ques- SW 34th Street, P.O. Box 147100, Gainesville, Florida 32601, USA tions came to mind. Why are many of the species so bril- (M.C. Thomas; P.E. Skelley; R.E. Woodruff); IEMM—Instituto Ecología, A.C., Departamento de Ecología y Comportamiento Animal, liantly colored, especially since they hide during the day? Apartado Postal 63, Xalapa, 91000 Veracruz, México (G. Halffter; E.D. Why do some of the species on Hispaniola and certain of Montés de Oca); IRSNB—Collections nationales belges d'insectes et those in Mexico exhibit the same combinations of colors? d'arachnides, Institut Royal des Sciences Naturelles de Belgique, 29, Rue

Fig. 1.—Head, pronotum, and base of elytra of subgenus Dyschromus, right side. A, E. tiburonica (Darlington); B, E. dimidiata Chaudoir. bli, basolateral impression; bp, basal puncture; br, basal ridge; dmg, dorsal mandibular grooves; ff, frontal furrow; ht, humeral tooth; msp, midlateral setal puncture; pbc, prebasal constriction; ptg, postocular transverse groove; s1, stria 1. Scale bar = 1.5 mm.

Vautier, B1040 Brussels, Belgique (K. Desender); MCZC—Department cleaned, and then returned to alcohol for storage. of Entomology, Museum of Comparative Zoology, Harvard University, For viewing at high magnifications, a few first and sec- 26 Oxford Street, Cambridge, Massachusetts 02138, USA (D.G. Furth); MHND—Departamento de Zoología, Museo Nacional de Historia ond instar larvae of each species were cleared in cold 10% Natural, Av. César Nicolas Penson, Plaza de la Cultura Juan Pablo KOH after puncturing the intersegmental membrane Duarte, Santo Domingo, República Dominica (C.A. Núñez); MNHB— between the head and thorax, and thorax and abdomen. The Museum für Naturkunde der Humboldt Universität zu Berlin, Bereich specimens were then washed in distilled water, followed by Zoologisches Museum, Invalidenstrasse 43, 1040 Berlin, Germany (F. 80% ETOH, and temporary glycerine mounts prepared as Hieke; M. Uhlig); MNHP—Entomologie, Muséum National d'Histoire Naturelle, Paris 75005, France (T. Deuve); MSUC—Department of described by Goulet (1977). Early instar larvae chosen for Entomology Collection, Michigan State University, East Lansing, illustration were cleared in Nesbitt’s fluid and mounted on Michigan 48824-1115, USA (F.W. Stehr); UASM—Strickland Museum, slides in Hoyer’s medium. Third instars were treated the Department of Biological Sciences, University of Alberta, Edmonton, same, except first the head and abdomen were detached, and Alberta T6G 2E9, Canada (G.E. Ball, D. Shpeley); SEMC—Snow Entomological Museum, University of Kansas, Lawrence, Kansas 66044, only temporary glycerine mounts were made. Exuviae were USA (J.S. Ashe); SLSC—S.L. Straneo Collection, Museo Civico di prepared for viewing in the same way as first and second Storia Naturale, Corso Venezia 55, 20121 Milano, Italy (C. Leonardi); instar larvae, but were not cleared. UAEM—Laboratorio de Parasitologica Vegetal, Universidad Autonoma del Estados de Morelos, Avenida Universidad 1001 Colonia Chamilpa, Illustrations.—Line drawings of major body parts of Cuernavaca 62210 Morelos, Mexico (A. Burgos Solorio); WIBF—West adults were made using a drawing tube mounted on a Indian Beetle Fauna Project, Montana State University, Bozeman, Montana 59717-3020, USA (M.A. Ivie). binocular dissecting microscope. Smaller adult structures, and features of larvae were illustrated using a drawing tube Holotypes of newly described taxa have been deposited in attached to a compound microscope. The color pastel the U.S. National Museum of Natural History, or returned paintings of the adult pronotum and adjoining portion of to the lending institution. the elytra were prepared by Patrice Stephens-Bourgeault, an artist and scientific illustrator at the Royal Ontario Museum. She viewed the specimens with a dissecting Rearing Techniques microscope, as well as the unaided eye, under both fluorescent and natural light, but added the warmer tones while Adults of the subgenus Dyschromus were obtained for rear- viewing the specimens under incandescent light. Most of ing during a collecting trip to Mexico in July and August, the SEM photomicrographs were taken digitally, using a 1992, during which time they were confined with some soil Hitachi Scanning Electron Microscope. in holding chambers made of cast blocks of Plaster of Paris that were first saturated with water, and then kept cool by Measurements and Statistics.—For a maximum of ten allowing the water to evaporate from the surface of the adult specimens of each sex of each species, various meas- block (Steel 1970). Back in Canada, the adults were kept in urements were made at up to 63× magnification using a plastic sandwich boxes containing soil and pieces of bark. dissecting microscope fitted with an ocular micrometer: Every two or three days the soil was examined for eggs or length of head (HL) in lateral view from apex of supraor- newly hatched larvae, which were transferred to small ver- bital ridge to posterior margin of eye; width of head (HW) sions of the Plaster of Paris holding chambers that we used including the eyes; length of pronotum along midline in the field. First instar larvae were fed adults and larvae of (PL); maximum width of pronotum (PWm); length of ely- Drosophila; second and third instars, and adults, were pro- tra (EL) from basal ridge to apex of elytra; maximum vided with pieces of larvae of the mealworm, Tenebrio width of elytra (EW); length and width of labial palpomere molitor. 3. For species known from at least five adults of each sex, the sample mean is given for each attribute, followed in Procedures for Study, Description,and parentheses by the range of values. If the sexes differed Illustration of Specimens statistically for an attribute (student’s t-test, p < 0.05), the mean and range were reported separately for each sex. If Preparation and Examination of Adults.—We used not, then only the mean and range for the total sample are standard methods to mount specimens, dissect out and pre- given, the minimum and maximum values followed by the pare the genitalia, etc. Adult microsculpture was studied at sex of the specimen with that measurement, unless a spec- 160 times magnification under diffuse light. Adult body imen of each sex was found to have that value. colors were viewed under fluorescent, incandescent, and Shape of the head and pronotum is indicated by the natural light. ratio of the width over the length of those parts of the body (HW/HL; PWm/PL), and elytral shape by the ratio Preparation of Larvae and Exuviae.—Larvae were pre- of the length to the width of the elytra (EL/EW). A meas- served by immersing them in just-boiled hot water until ure of the shape of labial palpomere 3 is provided by the their bodies became distended; they were then stored in ratio of the length to width of that palpomere. Ratios are 70% ETOH. Larval exuviae were placed directly in 70% reported in the same way as the measurement data, ETOH, and after a few minutes, were stretched and except the median value rather than the mean is given,

and differences between the sexes were tested using the from geographically distant locations, we asked ourselves if Mann-Whitney U-test (Sokal and Rholf 1969). we thought we could distinguish between them if we had To indicate range of body size of a given species, the samples from the intervening area. If the answer was “no,” apparent body length was obtained by measuring with a we treated the samples as conspecific. ruler to the nearest 0.5 mm, the distance from the anterior The most widespread and commonly collected species, margin of the head, including the mandibles, to the apex of E. dimidiata, was especially difficult to characterize because the elytra of the largest and smallest individuals of that adults exhibit considerable geographic variation, including species (Darlington 1952). differences in the male genitalia, as well as external features. For some taxa a few additional ratios are given, i.e., To some extent we were able to circumvent the difficulty maximum width of pronotum over width of pronotum at because for morphologically divergent populations from two the base (PWm/PWb), width of elytra over width of head adjoining regions of Oaxaca (Sierra Juarez and Sierra (EW/HW), maximum width of pronotum over head width Mixteca), we had available amino acid sequence data for a (PWm/HW), and width of head over length of metaster- fragment of mDNA indicating that the amount of divergence num (HW/ML). in this feature was the same as between populations of well Specific Epithets.—In the course of our work, we used as known species of butterflies (Sperling, Frania and Ball, in informal designations the names of localities at which or prep.). However, this information, by itself, would have been near where specimens of species that seemed to be unde- of little use in evaluating the status of one of two males col- scribed had been collected. We became sufficiently com- lected from a geographically distant locality in the Trans- fortable with these informal designations that we offer Volcanic Sierra (Buenavista de Cuellar, Guerrero); although them here as formal names. Except for Euchroa harrisoni, the male in question differed but slightly in external features each new specific epithet is an eponym and a noun in from Oaxacan specimens, and not at all from the other male, apposition based on an appropriate geographical place the shape of the median lobe of the aedeagus of the first male name. was unlike that of any other specimen ascribed to this species (Fig. 40h). We still judged it to be a specimen of E. dimidiata because the median lobe of the other male was in all Recognition of Species respects typical of specimens from western Oaxaca (as in Delimiting certain species of the subgenus Dyschromus Fig. 40e). Had only the abberant male been available, we was not easy, in part because of the small number of spec- would not have hesitated to recognize it as a new species imens available upon which to base taxonomic decisions. because the median lobe of the aedeagus of another species, Six of the species recognized are each represented by 60 to Euchroa ixtapa, new species, is more similar in appearance 140 individuals from three to 20 localities; 22 species are to that of typical E. dimidiata (cf. Figs. 39c, 40f), even each known from four to 23 specimens from two to five though for the mDNA fragment mentioned above, E. ixtapa sites; while seven species are each represented only by one has a different amino acid sequence from that of E. dimidia- to five individuals from a single site. ta (Sperling, Frania and Ball, in prep.). It is still possible that In principle, we regard species generally as reproductive- the two males from Buenavista de Cuellar represent different ly isolated (Endler 1989: 628–632) holomorphological species, but more specimens and perhaps mDNA sequence forms that may be characterized clearly (if not easily) by one data are required to examine this alternative. or more structural features (including color, setation, body Based upon the situation in E. dimidiata, in some form, proportions, form of male aedeagus, etc.). Such differ- instances where just a specimen or two was available, it is ences must characterize population samples, though in our possible that we have recognized new species that are really study some “samples” consisted of single individuals. just distinctive populations of another species, e.g., the In practice, we accorded the rank of species to recogniz- Hispaniolan Euchroa pedernales, new species, may be con- able parapatric or sympatric forms on the assumption that the specific with Euchroa perezi (Darlington). Conversely, sam- differences observed were maintained by reproductive isola- ples from certain localities regarded as populations of more tion (intra-form genetic continuity, and inter-form genetic widespread species may actually be distinct species, e.g., discontinuity). By “sympatric” we mean two forms collected populations of E. dimidiata from the Sierra Madre del Sur in concurrently in precisely the same localities. Three such Guerrero, or from the central part of the Trans-Volcanic pairs are represented in the subgenus Dyschromus: Euchroa Sierra. Such decisions, while somewhat arbitrary, and thus a lasvigas, new species and E. nitidipennis Putzeys (5.7 km. matter of judgment, are an unavoidable part of the taxonom- WNW Xico, Veracruz, on Tonalaco Road, 13 July 1999); ic process. Euchroa santacatarina, new species and Euchroa juchatengo, new species (22.2 mi. S. Juchatengo, Oaxaca, 21–22. Geographic Regions VII. 1966); and E. dimidiata Chaudoir and Euchroa carbonera, new species (7.9 km. NW La Carbonera, Oaxaca, 11 Geographical distributions of species of the subgenus June, 1979). Dyschromus are described in terms of the principal moun- To deal with samples of structurally divergent individuals tain systems of Mexico and the island of Hispaniola that

they inhabit: for Mexico, the Sierra Madre Oriental, Trans- Bionomics Volcanic Sierra, Sierra Madre de Oaxaca, and Sierra Madre del Sur. The Trans-Volcanic Sierra of Mexico is Diet.—When the genital segments were dissected out of subdivided into three regions, eastern, central, and west- an adult specimen, the hindgut usually came out as well. ern, that are indicative of areas supporting nearly continu- The hindgut nearly always contained fragments of ous stands of oak-pine, pine, and fir forest. One problem is Lepidoptera larvae, the most recognizable being the cro- placement of the boundary between the eastern part of the chets and mouthparts. We infer from the habitat prefer- Trans-Volcanic Sierra and the Sierra Madre Oriental. Ball ences of the subgenus Dyschromus that these fragments (1970) took the Rio Moctezuma to be the boundary, while are the remains of ground-dwelling caterpillars. There Smith (1940) placed it further north at about 22º latitude. did not appear to be any fragments of other common lit- The only species of the subgenus Dyschromus found north ter and soil-dwelling arthropods such as larval Elateridae of the Rio Moctezuma is E. harrisoni, new species, which and Scarabaeidae, and millipedes, even though in captiv- also occurs south of this river in the Sierra Zacualtipan ity both larvae and adults of Dyschromus readily ate (Fig. 31b). Other species of beetles exhibit the same distri- pieces of mealworm larvae. Since we did not find a bution pattern (Frania 1992), and so the Sierra Zacualtipan Dyschromus larva in the field, we cannot comment on the is treated as part of the Sierra Madre Oriental. natural diet of larvae. The Rio Blanco (the “deep valley” referred to by Ball [1970:95]) at the town of Orizaba is used to mark the bound- Adult Longevity.—Indirect evidence indicates that in ary between the eastern part of the Trans-Volcanic Sierra and nature, some adults of the subgenus Dyschromus can live the Sierra Madre de Oaxaca (Fig. 31b). The Sierra Madre del for several years. First, some adults survived in captivity Sur is continuous through Oaxaca and Guerrero, but most for up to four years. Of 44 adult females representing species of Dyschromus within this area are known only from eight species collected for rearing purposes in July and small portions of the highest mountain ranges. For Oaxaca, August, 1992, and kept alive for as long as possible, the Binford’s (1989) maps show most of these mountain ranges percentage still alive in August of each subsequent year and adjoining ones, and his names are used for them. For was: 77% in 1993, 45% in 1994, and 7% in 1995. One Guerrero, we mostly refer just to the Sierra de Atoyac adult female of E. harrisoni lived in captivity for four (Whitehead and Ball 1997), because most species of the sub- years and three months. But can adults live this long in genus Dyschromus known from Guerrero are from the vicin- the field? We think so because about five percent of ity of that range of mountains. pinned museum specimens show signs of excessive wear: worn mandibles, numerous scratch marks on the head, pronotum and elytra, and parts of the integument of the NATURAL HISTORY OF THE SUBGENUS head, pronotum and elytra dull and black in color instead DYSCHROMUS CHAUDOIR of having the typical luster and color for that species. We consider that such individuals, when collected, to have Habitat Preferences been active as an adult for more than one year. [N.B.: The tarsi were examined for signs of differential wear Species of the subgenus Dyschromus are confined to mon- between the inner and outer tarsal claws (Davies 1987), tane forests, and have been found at elevations of about but no differences were observed between individuals]. 900 to 3400 m, but none are known to have an elevational We cannot assess whether some species live longer than range greater than 1100 m. Fourteen of the species in others, because individuals that had been in captivity for Mexico are found primarily in oak-pine forest of dry to nearly two months in the field tended not to live as long mesic to temperate aspect, four species are mainly in cloud as individuals that were captured only a short time before forest, eight are mostly in mesophytic or oak-pine forest of our return from Mexico. tropical aspect, one occurs in montane tropical evergreen forest, and one is largely restricted to arid tropical scrub Reproduction.—Just 13 out of a total of 44 adult and dry scrubby oak, oak-palmetto, and oak pine wood- females, representing six of eight species of the sub- lands, as defined by Leopold (1950) and Rzedowski genus Dyschromus laid eggs in captivity. Most began (1978). Adults live on the ground, where, during the day, laying eggs shortly after we returned from Mexico in we have found them in leaf litter, within rotten logs, or September of 1992 (i.e., E. harrisoni, Euchroa perote, under stones or other cover. On a few occasions we E. dimidiata, Euchroa atoyac, Euchroa filodecaballo); observed an individual running about at night on the forest but females of Euchroa soladevega did not lay any eggs floor, but it is our impression, based upon observation of until the next summer. The maximum period over which adults in captivity, that they spend a considerable amount an individual laid eggs was 44 days, and although no of time in the soil, as do larvae. This would account for individual laid eggs a second year, some of those that why we often had so much difficulty finding adults in the did lay eggs lived for one or two years afterward. One field, and did not find any larvae. female of E. dimidiata was already so worn in appearance when it was captured in August, 1992, that one of

us (Frania) recorded the fact at the time. This female away. Moreover, this species may be absent from parts of laid eggs a month later; these subsequently hatched, the region that appear to have suitable habitat (e.g. the while the female lived on for another six months. We Sierra Miahuatlan). Thus, even E. dimidiata seems to infer that females are able to lay eggs for several years. have a restricted ability to disperse. As to why females in captivity did not lay eggs a second year, only a small number of females actually did lay eggs, so perhaps the rearing conditions were deficient. ADULTS Captive males did not live as long as females, but Morphological Traits of Adults of the Subgenus some that had been confined for as much as two years Dyschromus Chaudoir were observed copulating with females; thus males probably also remain fertile for several years. Color.—A notable characteristic of adults of most species belonging to the subgenus Dyschromus is that the dorsum Duration of Immature Stages.—Eggs hatched in is metallic in color. The most attractive and conspicuous 6–10 days. The duration of the first larval instar species are bicolored, although certain species with the was 5–19 days, and of the second instar 8–22 days. same combination of colors are much darker in hue than In the one instance that we obtained a pupa (of others (cf. Figs. 2c–e). Each of the bicolored species E. soladevega), the duration of the third instar was 27 exhibits one of four basic color combinations: head and days. pronotum metallic bluish-purple, elytra reddish-purple Interspecific Competition.—Most species of the sub- (Figs. 2c–e); head and pronotum coppery, elytra reddish- genus Dyschromus appear to have allopatric or parap- purple (Fig. 2f); head and pronotum coppery, elytra atric distributions, or if their geographical ranges over- bronze-colored (Fig. 3f); head and pronotum metallic lap, then they occur at different altitudes. Only green, elytra coppery (Fig. 3c) or brassy (Fig. 3d). The ely- E. dimidiata and E. carbonera, E. nitidipennis and tra of some other species are colored the same as the head E. lasvigas, and E. juchatengo and E. santacatarina, and pronotum, the dorsum being entirely metallic bluish- have been collected at the same locality (see section on purple, or metallic green (Figs. 3a–b), or brassy, or cop- species recognition for details). Although the overlap in pery (Figs. 2b, 3e). Depending on the angle at which a body size between the first two species is minimal, the specimen is viewed, and the kind of lighting, portions of second pair overlap considerably, while the third pair of the body of some species that are metallic green in color species do not differ in body size. The nearest neighbor exhibit a bluish (Fig. 3d) or reddish-purple reflection (Fig. of E. juchatengo and E. santacatarina is E. soladevega; 3a), while parts of the body that are brassy-colored will the latter is by far the largest of the three species, but have a bluish or greenish cast (Fig. 3d). Although two many neighboring pairs of species do not differ in body species of Dyschromus appear to the unaided eye to be all size, and E. soladevega appears to live in drier forest black in color, under magnification, it is evident that the than E. juchatengo and E. santacatarina. Hence evi- elytra of Euchroa tiburonica has a slight reddish-purple, dence from body size for interspecific interactions is and that of Euchroa opaca a slight brassy sheen (Fig. 2a). inconclusive. The metallic colors are presumably structural in origin, but washing some specimens in a hot solution of dish- Dispersal Ability.—All members of both subgenera of washing detergent alters the color. For example, certain Euchroa seem to have have limited powers of dispersal. individuals of E. filodecaballo, new species that were The hind wings of adults of all known species are repre- coppery-colored in life turned a brassy color after being sented by small flaps that are at most just twice as long killed, washed, and pinned, as did a borrowed pinned spec- as the reduced metanotum, so that the individuals in imen that originally had a noticeable greenish hue. The question cannot fly. Although some flightless species of elytra of a pinned specimen of E. pedernales from Larimar ground beetles that occur in Mexico do have extensive mine, which was originally coppery in color with a distinct geographical distributions, e.g., Platynus stricticollis greenish tint, took on after washing an intense reddish hue. (Bates) (Liebherr 1988: Fig. 66), most species belonging So did a specimen of E. cupripennis, new species, but to the subgenus Dyschromus have very restricted distri- another retained its greenish tint. butions, each being confined to one or two mountain Microsculpture and Luster.—The microsculpture on ranges. Only E. dimidiata is comparatively widespread the dorsal surface of the body consists of sculpticells that (Fig. 42a), undoubtedly because it lives in the dry scrub- form a mesh pattern (Fig. 5a). Most of the variation in the by oak-palmetto and oak woodlands that form a relative- subgenus Dyschromus for micro-sculpture has to do with ly continuous fringe around the Rio Balsas Basin, Valley the elytra; that on the disc is invariably different in some of Tehuacan, and Valley of Oaxaca. Even so, populations respect from that at the sides and apex (cf. Figs. 5a–b, 5c; of E. dimidiata from different parts of this region are dis- 5d–e, 5f; 6a–b, 6c). Differences concern, in part, whether tinctive, and specimens from adjoining areas do not the surface of the sculpticells is flat (Figs. 5a–b), convex resemble each other more than those from areas further (Figs. 5d–e, 6a–b), or keeled (Figs. 7a–b). If the sculpti-

Fig. 2.—Paintings done in pastel of base of head, pronotum, and base of elytra of subgenus Dyschromus, left side and part of right side. A, E. opaca (Chaudoir); B, E. centralis (Darlington); C, E. sallei Chaudoir; D, E. perote n. sp.; E, E. teotitlan n. sp.; F, E. huautla n. sp.

Fig. 3.—Paintings done in pastel of base of head, pronotum, and base of elytra of subgenus Dyschromus, left side and part of right side. A, E. atoyac n. sp.; B, E. suchixtepec n. sp.; C, E. pedernales n. sp.; D, E. dimidiata Chaudoir; E, E. chrysophana Bates; F, E. yucuyacua n. sp.

cells are convex, then at low magnification, the surface of rounded (Figs. 1a, 12d), obtuse (Figs. 8a–b), acute and the elytra appears to be beaded (Fig. 15b). Although the slightly prominent (Figs. 1b, 12a, 12c, 13a–b), or acute and elytra of nearly all species of Dyschromus exhibit some prominent (Fig. 13c). The basolateral impressions (bli) are beading, in most of them it is confined to the sides short (Fig. 1b), moderately long (Fig. 1a), or long (Fig. (intervals 8–9) and apex; in many species only the apex 13a). A few species have a shallow posterior transverse is beaded (Fig. 15a). The sculpticells on different parts impression (pti) between the basolateral impressions (Fig. of the elytra and in different species can also differ in 8b). The apical third of the prosternal intercoxal process is shape and the degree to which they are impressed, being either margined all around, only laterally, or not margined. isodiametric to slightly transverse with respect to the longitudinal axis of the body (Figs. 5a, 5d), or elongate Pterothorax.—All members of the subgenus Dyschromus (Figs. 6a), and finely (Figs. 5a–b) to deeply impressed have somewhat vaulted elytra (Figs. 14a–b), markedly (Figs. 5d–e, 6a–b), or even effaced. The sculpticells reduced hind wings, and short metasternum. Species from invariably are flat and finely impressed if the elytra has the island of Hispaniola have the shoulders entirely rounded, a brilliant luster; they are convex and relatively deeply without indication of a humeral tooth (Fig. 1a). The shoul- impressed in many species in which the elytra has a dull ders of the elytra of species from Mexico are more promi- luster or is only slightly shiny. nent, and a humeral tooth is usually evident (Fig. 1b, ht). Consistent with the slightly vaulted elytra, the apical decliv- Head.—The frontal furrows are variously developed ity of the elytra is at least moderately steep (Fig. 14a), and in (Figs. 1, 8). At one extreme, the furrows extend posteriorly most individuals of a few species it is nearly perpendicular to the level of the anterior supraorbital setigerous punctures (Fig. 14b). and are shallow throughout (Fig. 1a, ff); at the other The basal punctures of the elytra (Fig. 1a, bp) are extreme, they extend almost to the postocular transverse absent from a few species. The striae are intact on the groove and are deeply and sharply impressed for nearly disc of the elytra, or at least stria 1 (Fig. 1b) is interrupt- their entire length (Fig. 1b). ed in one or more places. If two or more striae are inter- The contours of the submentum and adjoining portion of rupted, stria 1 is interrupted the most times, and the the transgular groove vary: at one extreme, the submentum outermost striae usually the fewest times, if at all. In (Fig. 9b, sm) is not prominent and the posterior face slopes some species where no individual was found to have evenly to the transgular groove (tgg), which is shallow; at the stria 1 interrupted more than 15 times, a few individuals other extreme, the submentum is very prominent and has a were observed to have all of the striae intact. This was concave posterior face, while the transgular groove posterior not so if any individual of a species had stria 1 interrupt- to the submentum is very deep (Figs. 9d, 10b). In most ed at least 20 times. Among species with individuals that species, the submentum is moderately prominent and slopes have all of the striae interrupted at least 20 times on the evenly to the transgular groove, which is moderately deep disc, one specimen of E. ixtapa is unusual in that striae (Figs. 9c, 10a). 2–6 are interrupted fewer times than stria 1 or stria 7 and 8. With increasing fragmentation, the striae become Mouthparts.—The dorsal mandibular grooves in most more shallowly impressed, those interrupted more than individuals are long (Fig. 1a, dmg), but are short in some 20 times being barely evident in some species. Among specimens of some species (Fig. 9a). Labial palpomere 3 is species in which all of the striae are intact on the disc, in broader than the more proximal ones, and in shape is either some the elytral intervals are somewhat to moderately securiform (Fig. 11b), broadly triangular (Figs. 11a, 11d), convex (Fig. 15a), in others the intervals are nearly flat narrowly triangular (Fig. 11c), or subtriangular (Fig. 11e); (Fig. 15b). The elytral intervals are invariably flat in it is wider in most males than females (cf. Figs. 11a–b, species with greatly interrupted striae. The mesepister- 11c–d), but in species where it is subtriangular in the male, num is either smooth or punctate. then it is also subtriangular in the female. The paramedian pits of the mentum (Fig. 10a, pmp) vary in size: in many Legs.—Males of many species of the subgenus species they are small, being about the same diameter as the Dyschromus have the inner face of the subapical part of the umbilicus at the base of the submental setae (Figs. 10a–b); middle tibia slightly to moderately expanded medially (Fig. in other species they are moderately large, i.e. distinctly 16a); females do not. larger in diameter than the umbilicus mentioned above, but Along the hind tibia are four rows of setae. The setae of not greater in diameter than the ventral tentorial pits; in a the dorsomedial, lateral, and ventrolateral rows are stout few species they are large, at least twice the diameter of the and spiniform (Fig. 16e, dmrs; Fig. 16b, lrs); those of the tentorial pits. medial row (mrs) are longer and more slender, and are referred to as flexile setae (Fig. 16b, fs). Each seta in the Prothorax.—The sides of the pronotum are entire dorsomedial and medial rows arises from a coarse punc- (oblique) posteriorly (Figs. 1a, 8a–b, 12a, 12d, 13d), or ture that is connected to the next puncture in the row by a slightly (Figs. 3b, 13b) or moderately (Figs. 1b, 12b, 13a) groove, zig-zag fashion (Fig. 16e, dmg; Fig. 16b, mg). The or markedly sinuate (Figs. 12c, 13c). The hind angles are medial groove (Fig. 16b) does not extend basally much

beyond the basal seta in the medial row of setae, and in mary one. Several conditions of the primary haemolymph both sexes of most species is rather shallow, so that the channels are evident: three or more primary channels pres- setal punctures appear to be more discrete than in the other ent, largest one with origin at far right side of ostium, two rows. The lower edge of the medial groove forms a medi- or more progressively smaller channels originating at al ridge (Fig. 16c, mri). Males of many species of intervals to left of largest one (Fig. 18b); just two primary Dyschromus have at least the basal portion of the medial channels, a longer one at far right side of preapex, and a ridge represented by a row of subangulate to subrectangu- broader one to left of midline (Fig. 19c); only one primary lar tubercles, each of which is associated with one of the channel, at far right side of preapex (Fig. 34c); only one medial flexile setae (Figs. 16b, d–e). centrally located primary channel (Figs. 38b, 40c). The Most species of the subgenus Dyschromus have one last two conditions of the primary haemolymph channels row of medial flexile seta on the hind tibia (Fig. 16b, are not be confused with another in which nearly the entire mrfs), but E. tiburonica and E. opaca have two such rows preapex is occupied internally by a large haemolymph (Fig. 16f), with about ten setae in the outer row, and three sinus (Figs. 28e, 29c, 29f), which could be a developmen- to five setae in the inner row, all arising from a common tal abnormality. groove. Euchroa tiburonica and E. opaca also have two In most species the left wall of the periostial part of the rows of medial flexile setae on the middle tibia; so do median lobe (lw) is as extensive or slightly more so than the some other species of Dyschromus, but E. tiburonica and right wall, so that neither the right wall nor the ostium are E. opaca have more such setae than the others. visible in left lateral view (Fig. 19b); in a few species the left wall is not so extensive, so that both the right wall (rw) Abdomen.—Abdominal sterna V–VII each have a pro- and part of the ostium are visible in left lateral view (Fig. nounced basal sulcus, which does not extend to the lateral 34b). In most species the far right side of the periostial part margins of the abdomen (Fig. 14c, bs), except in Euchroa of the median lobe is produced ventrally, forming a perios- tenancingo, new species (Fig. 14d). The basal sulcus is tial ventrolateral bulge (pvb); depending on the species, the foveate laterally (Fig. 14c), and in some species the basal bulge is comparatively proximal (Fig. 19b) or distal sulcus of sternum VII has an additional median fovea, (Fig. 25b) in position, and is slightly (Figs. 25b, 26a) to while in a few species the entire sulcus is foveate (Fig. markedly prominent (Figs. 26b, 29b) in profile, and either 14d). In most individuals the foveae are relatively coarse, subangulate (Fig. 26c) or rounded (Figs. 26a–b) in cross- but are rather inconspicuous in at least some individuals of section. some species. In some species, a dorsomedial groove (dmg) extends the entire length of the central part of the median lobe (Fig. Median Lobe of Aedeagus.—The distal portion of the 34a); in certain other species it extends only a short dis- median lobe of the aedeagus is quite varied in form in the tance beyond the prebasal bend (Fig. 36f). subgenus Dyschromus compared to the subgenus Euchroa. Some of these traits of the median lobe appear only in To describe this variation, the median lobe is subdivided certain combinations in the subgenus Dyschromus, so that into four regions (Figs. 19a–b). The preapical part or five distinct types of median lobe are recognizable, one of “preapex” (p) extends from the apex of the median lobe to which is also characteristic of the subgenus Euchroa. the ostium (o). The periostial part (pr) is that portion of the Three of the other types are considerably more asymmet- median lobe which includes the ostium. The middle part rical than is usual in the subgenus Euchroa, while the (m) extends from the base of the ostium over the prebasal fourth type is more symmetric. The five types are bend (Fig. 19b, pb) to the basal bulb (bb), which is the last described in detail in Appendix 1, along with intermediate region. types, and the species that exhibit each type. The preapex is twisted downwards to the left (Fig. The internal sac is not so varied in structure as the 19b). That portion of the preapex approximately to the left median lobe. The apical portion of the sac is slender and of the midline of the median lobe is, in many species, thin- glabrous. The gonopore (gp) is located either dorsally or ner and not as darkly sclerotized as the area to the right on the right side of the sac, a short distance below the apex (Fig. 19c). In some species the area along the left margin (Figs. 19d, 24f, 41a). The main body of the sac is much is so thin that it is colorless (Fig. 32c); in other species, it wider than the apical portion, and in some species is con- is the middle third of the preapex that is extremely thin and stricted in the middle (Fig. 19d). The inner surface of the colorless (Fig. 24c). In certain species, the preapex is not sac has a vestiture of transverse scale-like microtrichia, the as extensive, or not developed at all, to the left of the mid- largest being moderately darkly sclerotized and arranged line of the median lobe (Figs. 33c, 34c). In many species a either in one broad central band (Fig. 24f), or a narrow preapical dorsolateral ridge (pdr) extends the length of the basal band (Fig. 19d), or both a basal and subapical band preapex on the far left side (Figs. 19a, 24a); this ridge is, (Fig. 41a); in the latter case, only portions of one or both in profile, flat (Figs. 19b, 47b) to markedly convex (Fig. bands may be evident. 34b). Up to three or more primary preapical haemolymph channels (phc) originate at the base of the preapex (Fig. Female Genitalia.—The female genitalia in the subgenus 18b); numerous secondary channels branch from each pri- Dyschromus vary in the form and length of the spermatheca:

short and nearly straight (Fig. 17a, sp), or long and recurved and lack the elytral “scutellar” stria (see Ball 1959, for the apically (Fig. 17b). correct interpretation of this trait). Most euchroines also exhibit the following: labial mentum with pair of parame- Sexual Dimorphism.—Besides all males having expand- dian pits; sterna V–VII each with transverse post-basal sul- ed front tarsi, males of most species belonging to the sub- cus (complete or not, punctate or not); ring around umbili- genus Dyschromus differ from females in that, as noted cal setigerous puncture, located near elytral humerus, above, labial palpomere 3 is broader (Figs. 11a–d), the inner asymmetrical and tuberculate; duct of spermathecal gland face of the middle tibia is expanded medially near the apex with tube-like extension (Fig. 17b, di); and bursa copulatrix (Fig. 16a), and at least the basal portion of the medial ridge with gooseneck shape (Shpeley and Araujo 1977, fig. 3). of the hind tibia is represented by a row of subangulate Euchroa, Bothynoproctus, and Microcephalus emerge tubercles (Figs. 16b, d–e). Males of a given species also as a group in Will’s (2000) cladogram, and along with tend to be smaller, with pronotum and elytra narrower than Lobobrachus, were included in the Euchroina by Straneo females (see, for example, description of E. perote), and the (1977) and Reichardt (1977). We refer to these four genera prebasal constriction at the sides of the elytra tends to be as the Euchroa-Microcephalus clade, which is further more pronounced (Fig. 1b). characterized by: color (dorsally, black to brilliant colors of purple, blue, green, and red, singly or in various combi- LIMITS AND RECOGNITION OF THE nations); labial mentum with median tooth broad at apex, SUBTRIBE EUCHROINA apical margin truncate or emarginate, paramedian pits small to large; abdominal sternal V–VII each with basal Since the creation of the group Euchroides by Chaudoir sulcus complete and deep, setigerous punctures of sternum (1874: 16) to include Euchroa Brullé (+ Dyschromus VII more or less foveate, and located medially or at least Chaudoir) and the Neotropical genus Microcephalus distinctly anteriad the posterior margin. Dejean, only Euchroa (with Dyschromus recognized as a distinct genus or not) has been consistently included in the SYSTEMATIC ACCOUNT Euchroina. Other genera that have been placed by one author or another in the subtribe are: the Neotropical ORDER COLEOPTERA Lobobrachus Sharp, Haplobothynus Tschitschérine, and FAMILY CARABIDAE Bothynoproctus Tschitschérine, and the Australian Setalis SUBFAMILY HARPALINAE Laporte de Castelnau (Tschitschérine 1898:46; Cisiki TRIBE PTEROSTICHINI 1929:522–524; Reichardt 1977:407–409; Straneo 1977: SUBTRIBE EUCHROINA 108–109, 115, and 1979: 347–349). Will (2000:54) in a cladistic analysis of pterostichine genera of the world markedly expanded the Euchroina, including therein, in Genus Euchroa Brullé, 1834 addition to the above genera (Will did not examine Euchroa Brullé, 1834: 335. TYPE SPECIES (by monotypy): Euchroa Lobobrachus or the subgenus Euchroa), 11 more genera nitidicollis Brullé, 1834: 336. Putzeys, 1844: 403. Lacordaire, from South America, some such as Abaris Dejean, with 1854: 319. Chaudoir, 1874: 15. Bates, 1882: 84. Tschitschérine, distributions extending to North America, two Australian 1898: 60. Csiki, 1929: 522. Straneo, 1938: 25. Jeannel, 1942: 735. genera included by Moore (1965) as part of his Setalis Reichardt, 1977: 407. Straneo, 1977: 115; 1979: 347. Shpeley and Araujo, 1997: 400. series, one genus from New Caledonia, one from North Dyschromus Chaudoir, 1835: 429. TYPE SPECIES (by monotypy): Africa and the Mediterranean Basin, and with some reser- Dyschromus opacus Chaudoir, 1835: 430. Lacordaire, 1854: 311. vation, the Holarctic genus Stomis Clairville. Chaudoir, 1871: 285; 1874: 15 (Euchroa). Bates, 1882: 84; 1891: 249. Tschitschérine, 1898: 58. Darlington, 1935:179; 1939:88. Reichardt, 1977: 407. Straneo, 1979: 349. Ball and Erwin, 1983: Recognition 491. As defined by Will (2000) the subtribe Euchroina is poly- Notes about Synonymy.—Chaudoir (1835: 430) based thetic (Sneath and Sokal 1973: 20–23), meaning that it is Dyschromus on the Hispaniolan species D. opacus, diagnosed by a combination of features all of which are not the type material of which he recorded incorrectly as necessarily exhibited by all of the included taxa (see Will having been collected in Java or nearby islands. 2000: 54, fig. 1.15. for a cladogram showing distribution of Subsequently, Chaudoir (1874: 15) synonymized the character states among the euchroines). Also, all of the names Euchroa and Dyschromus, an action followed by states in the diagnostic combination are exhibited by one Bates (1882: 84), indicating thereby the close relation- or more genera in other pterostichine assemblages. This ships of these two groups. Tschitschérine (1898: 58) also accounts in part for the historical instability in composition recognized the close affinity of Euchroa and of the Euchroina, the other reason being the limited geo- Dyschromus, but he concluded that they represented dif- graphical scope of some of the studies. ferent genera. Subsequent authors have accepted Euchroine adults lack the seta of the hind trochanter, Tschitschérine’s judgment. We believe that Euchroa and

Fig. 4.—Map of portion of western Hemisphere showing approximate geographical distributions of the subgenera of Euchroa.

Dyschromus are each monophyletic, but because of the Geographical Distribution.—The genus Euchroa is rep- absence of marked structural divergence between them, resented in the Neotropical Region in five separate areas we accept Chaudoir’s judgment that the two groups are (Fig. 4): South America (three widely separated centers, the congeneric, and in keeping with the ranking of other isolation of which is probably an artifact of inadequate col- groups of Neotropical pterostichines, we rank Euchroa lecting effort in the intervening areas); the central Mexican and Dyschromus as subgenera. highlands, in mainland Middle America north of the Isthmus of Tehuantepec; and the Caribbean island of Recognition.—Adults of this genus are euchroines of the Hispaniola, in the Greater Antilles. Euchroa-Microcephalus clade, with labial palpomere 2 bisetose, elytral interval 3 impunctate, basal sulci of Chorological Affinities.—See this topic under the sub- abdominal sterna V–VII foveate (crenulate) laterally, and generic treatments, below. abdominal sternum VII with posterior margin smooth, not beaded, without marginal sulcus, and with one pair of Phylogenetic Relationships.—This genus is a member of foveate setigerous punctures medially (both males and the clade that also includes Bothynoproctus, Lobobrachus females). and Microcephalus (see section above on limits and recognition of the Euchroina). Description.—Deferred until the subgenus Euchroa is revised. See treatment of subgenus Dyschromus for many details. Included Species.—The 41 species of Euchroa are arrayed in two subgenera, Euchroa (sensu stricto) and Dyschromus. Habitat.—Forest floor, in leaf litter, or under stones in pas- For details, see below. tures and disturbed areas adjacent to forest. Members of this genus occur in lowland broadleaf wet tropical forest, gallery forests in lowlands, and wet to dry tropical to tem- Subgenus Euchroa (sensu stricto) perate highland forests. The altitudinal range extends over about 3000 m, from 400 to 3400 m. Diagnosis.—Adults of this subgenus are distinguished from the subgenus Dyschromus by their comparatively

large size (apparent body length at least 13 mm), in combi- Diagnosis.—Compared to the subgenus Euchroa, most nation with the following: paramedian mental pits large; adults of the subgenus Dyschromus are considerably small- pronotum with two pairs of well-developed basolateral er in size, and in nearly all species the paramedian pits of impressions; lateral margins of the pronotum entire with the mentum are smaller in diameter, the lateral margins of hind angles rounded; elytral striae deep; elytral intervals the pronotum are at least slightly sinuate, the elytral striae convex, more so posterolaterally; microsculpture mesh pat- are shallower, and the elytral intervals are flatter. Of the tern on disc of elytra markedly transverse, sculpticells on two pairs of basolateral impressions present on the prono- all parts of elytra finely impressed and flat; basal sulcus of tum in the subgenus Euchroa, only the inner pair are well- abdominal sterna V–VII each with median fovea in addi- developed in the subgenus Dyschromus; the outer pair are tion to lateral foveae. rarely evident and then only as indistinct punctiform impressions. The microsculpture mesh pattern on the disc Description.—Deferred, pending a revision of the species in this subgenus. of the elytra also differs, the sculpticells being isodiametric Habitat.—Temperate Atlantic forest to Amazonian low- to slightly transverse or elongate in shape. Also the land tropical rainforest, on forest floor, in leaf litter. microsculpture at least at the sides and apex of the elytra is considerably more conspicuous than in the subgenus Geographical Distribution.—The range of Euchroa Euchroa because the sculpticells are at least moderately (sensu stricto) is confined to South America, in the deeply impressed, and in all but a few species, at least at the Amazon Basin of Peru in the west, to the Brazilian state of apex of the elytra, the surface is beaded, i.e., the sculpticells Rio de Janeiro. Bates (1882: 85) indicated a range for the are convex. While the basal sulcus of abdominal sternum group extending south to the Rio Plata, but this was prior VII in Dyschromus can have a median fovea in addition to to the recognition by Straneo (1938: 25–26) that the south- the usual lateral foveae, the basal sulcus of sterna V and VI ern species, Euchroa obscura Putzeys, belonged in Eumara is foveate only laterally. Tschitschérine rather than in Euchroa. A few females of E. (Dyschromus) perote are as large as the smallest specimens of the smallest member Chorological Affinities.—Among Euchroina, the range of of the subgenus Euchroa (an undescribed species from subgenus Euchroa overlaps part of the range of Espirito Santo and eastern Minas Gerais, Brazil, MNHP, Bothynoproctus, and most of the range of Microcephalus. BMNH, SLSC). In E. (Dyschromus) tiburonica and Probably it overlaps completely the range of Lobobrachus. E. (Dyschromus) opaca the paramedian pits of the mentum are as large in size as in the subgenus Euchroa. All indi- Phylogenetic Relationships.—This subgenus is the postu- viduals of Euchroa (Dyschromus) miahuatlan, new lated sister-group of the Mexican-Hispaniolan subgenus species, and some individuals of a few other species of the Dyschromus. subgenus Dyschromus approach the subgenus Euchroa in having the sides of the pronotum nearly entire posteriorly. Included Species.—The subgenus Euchroa includes six E. (Dyschromus) sallei and Euchroa (Dyschromus) huaut- species: Euchroa nitidicollis Brullé; Euchroa dives la, new species, have the elytral striae about as deeply Tschitschérine; Euchroa onkonegare Shpeley and Araujo; impressed and the elytral intervals about as convex as in and three new species, presently undescribed. the subgenus Euchroa. In Euchroa (Dyschromus) centralis, Euchroa (Dyschromus) jalisco, new species, and Subgenus Dyschromus Chaudoir, 1835 some individuals of a few other species, all of the sculpticells along the sides and at the apex of the elytra are flat, Synonymy.—See “notes about synonymy” in the generic treatment, as in the subgenus Euchroa. No two of these features are above. combined in all individuals of any one species of Dyschromus, so recognition is not difficult. Taxonomic History of the Species.—As already discussed (see Introduction), the first species of Dyschromus Description.—Body Size and Proportions. Apparent body length to be described was D. opacus from Haiti (Chaudoir 1835: 6.5–14.0 mm. Eyes moderately large and prominent: eye length about 430). The second, described in the genus Euchroa by 2.25× length of temples from posterior margin of eye to postocular transverse groove, width of head including eyes about 1.33–1.44× greater than Putzeys (1844: 403), was E. nitidipennis from Mexico. width of head between eyes (Figs. 1a–b, 8a–b). Pronotum moderately Next, Chaudoir in 1874 (p. 17–20), treating Euchroa and transverse: 1.23–1.56× wider than long, slightly to moderately constricted Dyschromus as congeneric, described E. cupripennis from posteriorly: maximum width 1.06–1.26× width at base (Figs. 1a–b, 8a–b, Haiti, and E. dimidiata and E.sallei from Mexico. Bates 12a–d, 13a–d). Elytra moderately broad, 1.31–1.69× longer than wide. Color and Luster. Dorsum black (Fig. 2a), brassy (Figs. 2b, 3e), metal- added two more species of Euchroa from Mexico: E. flohri, lic green (Figs. 3a–b) or metallic bluish-purple, or dorsum bicolored, with in 1882 (p. 85); and E.chrysophana in 1891 (p. 249). Most head and pronotum metallic bluish-purple or coppery colored, and elytra recently, Darlington, treating Dyschromus as generically reddish-purple (Figs. 2c–f), or head and pronotum metallic green and ely- separate from Euchroa, described D. tiburonicus from Haiti tra coppery (Fig. 3c) or brassy (Fig. 3d), or head and pronotum coppery and (1935: 179), and D. perezi and D. centralis from the elytra bronze (Fig. 3f). luster at least somewhat shiny, not iridescent. Microsculpture. Dorsum with microsculpture of sculpticells that form Dominican Republic (1939: 88). mesh pattern. Vertex of head with sculpticells isodiametric, finely

Fig. 5.—Scanning electron photomicrographs of microsculpture on elytra of subgenus Dyschromus. A and B, E. nitidipennis Putzeys; C, E. sallei Chaudoir; D, E, and F, E. dimidiata Chaudoir; A, D, disc, dorsal aspect; B, E, disc, lateral aspect; C, F, interval 9 near middle of elytra. ant, anteri- or; sc, sculpticell. Scale bar = 25 um.

Fig. 7.—Scanning electron photomicrographs of microsculpture on elytra of subgenus Dyschromus, E. opaca (Chaudoir). A–B, beginning of apical declivity, dorsal and lateral aspects. Scale bar = 25 um.

with most sculpticells slightly transverse, some isodiametric, finely impressed, effaced in spots in some species. Elytra with sculpticells on disc isodiametric to slightly transverse, finely to moderately deeply impressed, and flat (Figs. 5a–b), or moderately deeply impressed and slightly convex (Figs. 5d–e), or sculpticells elongate and markedly convex (Figs. 6a–b) or keeled (Figs. 7a–b), or sculpticells very finely impressed and effaced in spots, or elytral intervals on disc entirely smooth; outer half of interval 8 and interval 9, and apex of elytra with sculpticells at least moderately deeply impressed (Fig. 5c), outer half of interval 8 in a few species, and interval 9 and apex of elytra in most species with at least some patches of convex sculpticells (Figs. 5f, 6c); elytral surface beaded where sculpticells convex (Figs. 15a–b). Head. Frontal furrows variously developed, at one extreme extended posteriorly only to level of anterior supraorbital setigerous punctures and shallowly impressed throughout (Fig. 1a), at other extreme furrows extended posteriorly nearly to postocular transverse groove and deeply Fig. 6.—Scanning electron photomicrographs of microsculpture on elytra and sharply impressed for nearly entire length (Fig. 1b). Postocular trans- of subgenus Dyschromus, E. soladevega n. sp. A–B, disc, dorsal and lat- verse groove shallow but distinct dorsomedially (Fig. 1b) or obsolete dor- eral aspects; C, interval 9 near middle of elytra. ant, anterior. Scale bar somedially or nearly so (Fig. 1a). Submentum and transgular groove var- = 25 um. iously developed, at one extreme submentum not prominent, posterior face sloped evenly to transgular groove, with latter shallow posterad sub- impressed in most species, nearly effaced in some species, not more mentum (Fig. 9b), at other extreme, submentum very prominent, posteri- deeply impressed than on pronotum, except in E. tiburonica. Pronotum or face concave, and transgular groove very deep posterad submentum (Figs. 9d, 10b).

Fig. 8.—Head and pronotum of subgenus Dyschromus, right side. A, E. cupripennis Chaudoir; B, E. perezi n. sp., holotype. msp, midlateral setal puncture; pti, posterior transverse impression. Scale bar = 1 mm.

Mouthparts. Dorsal mandibular grooves long in most species (Figs. stria 1 or scutellum (Figs. 1a–b), except shorter in E. lasvigas, new 1a–b), short in some or all individuals of a few species (Fig. 9a). Labial species; condition of elytral striae and intervals various, at one extreme palpomere 3 of males securiform (Fig. 11b), triangular (Fig. 11d), or sub- striae intact and deeply impressed for their entire length, and intervals triangular (Fig. 11e), of females triangular and narrower than in males moderately convex, at other extreme striae 1-6 and at least anterior two- Figs. 11a, 11c), or subtriangular as in males, palpomere 3 in males thirds of 7 very faintly impressed and interrupted more than 30 times, and 0.85–2.15 and females 1.26–2.61× longer than wide. Maxillary palpomere intervals flat, stria 8 and at least posterior third of 7 comparatively deeply 4 fusiform, 2.5–2.9× longer than wide; palpomere 3 1.7–2.1× longer than impressed. Hind wings very small, from about as long as, to about twice wide (Fig. 11f). Mentum with paramedian pits small (Figs. 10a–b) to length of, metanotum. Metasternum short. Mesepisternum smooth to large. coarsely punctate. Prothorax. Pronotum with sides evenly rounded anteriorly (Fig. 1b), Legs. Middle tibia with one or two rows of medial flexile setae (Fig. except slightly explanate anteriorly in E. tiburonica (Fig. 1a), sides nearly 16a). Hind tibia of most species with one row of medial flexile setae (Figs. flat posterolaterally, except entirely flat in Euchroa independencia, new 16b–e), a few species with two rows (Fig. 16f). species; lateral margins in most species at least slightly sinuate towards Abdomen. Sterna III–VII each with medial pair of foveate setigerous base (Fig. 13b), lateral margins entire in a few species (Fig. 1a); hind punctures, at about one-half distance to posterior margin of sternum (Fig. angles nearly rounded (Fig. 12d), obtuse (Fig. 1a), or acute (Fig. 1b); mid- 14c, fsp), sterna III–IV of E. tiburonica with few additional asetose foveae lateral setal puncture distinctly separated from marginal bead (Fig. 1a), basolaterally; sterna V–VII each with basal sulcus, not extended to lateral except in contact with marginal bead in E. perezi (Fig. 8b); median longi- margins of sternum (Fig. 14c), except E. tenancingo, n. sp. (Fig. 14d); basal tudinal impression extended posteriorly to point level with midpoint of sulcus foveate laterally, basal sulcus of sternum VII without median fovea basolateral impressions (Fig. 1a), except further in Euchroa puertogallo, (Fig. 14c), or with median fovea, or sulcus entirely foveate (Fig. 14d). new species, and some specimens of E. chrysophana and E. filodecabal- Male. Middle tibia with inner face expanded near apex (Fig. 16a) or lo, new species; inner pair of basolateral impressions well-developed, not. Hind tibia with medial ridge entire (Fig. 16c), or represented basally punctiform, or short (Fig. 1b), or moderately long (Fig. 13b), or long and (Fig. 16d) or for entire length by row of subangulate or subrectangular attaining posterior margin of pronotum (Fig. 12c), outer pair of basolater- tubercles (Figs. 16b, 16e). Aedeagus with preapical part of median lobe al impressions evident just in a few individuals and then only as faint twisted downward to left (Fig. 19b), variously shaped, uniformly thick punctiform impressions; transverse impression between inner basolateral throughout (Fig. 40c) or left half thinner and less darkly sclerotized than impressions present only in a few species (Fig. 8b). Prosternum with api- right half (Fig. 25f) or preapex only developed to right of midline of medi- cal third of intercoxal process margined or not. an lobe (Fig. 34c), preapical dorsolateral ridge present (Figs. 19a, 24c) or Pterothorax. Elytra slightly vaulted upward; shoulders entirely round- not, preapex with one or more primary haemolymph channels (Figs. 18b, ed (Fig. 1a), or with humeral angle prominent or dentiform (Fig. 1b); sides 19c); periostial part of median lobe with left wall more extensive than of elytra in males of most species slightly constricted anterad of middle right wall (Fig. 19b) or vice versa (Fig. 34b); periostial ventrolateral bulge (Fig. 1b), less or not at all in many females (Fig. 1a); apical declivity in present (Fig. 19b) or not (Fig. 20b), in cross-section rounded (Figs. 26a–b) most species moderately steep (Fig. 14a), in some or most individuals of or subangulate (Fig. 26c); middle part of median lobe with dorsomedial some species nearly perpendicular (Fig. 14b); basal ridge extending to groove present (Figs. 25d, 45a) or not (Fig. 18a). Internal sac with gono-

pore subapical (Fig. 19d); inner surface with vestiture of plate-like assemblages: the opaca group, from Hispaniola; and the microtrichia, largest microtrichia in one broad or one or two narrow bands nitidipennis group, from Mexico. Details are provided extended partially or entirely around sac (Figs. 19d, 24f, 41a), microtrichia oval to triangular in shape, except left side of basal band with microtrichia below. spinose in some populations of E. dimidiata (Figs. 24f, 41a–b). Female. Ovipositor and internal genitalia, including stylomeres, as in Euchroa onkonegare (Shpeley and Araujo 1997, figs. 2–3), except diver- CLASSIFICATION OF SPECIES OF THE ticulum (di) at base of duct of spermathecal gland long (Fig. 17b), sper- SUBGENUS DYSCHROMUS CHAUDOIR mathecal gland ovoid, and spermatheca short and nearly straight (Fig. 17a) or long and recurved (Fig. 17b), with wide, spiral ribbing. Opaca Group Remarks.—Another diagnostic trait for the subgenus Opaca Subgroup Dyschromus may be the subapical position of the gonopore 1. Euchroa tiburonica (Darlington) of the internal sac of the aedeagus. In other euchroines for 2. Euchroa opaca (Chaudoir) which we have been able to examine the internal sac, 3. Euchroa centralis (Darlington) including the undescribed species of the subgenus Euchroa referred to in the diagnosis of the subgenus Dyschromus, Cupripennis Subgroup the gonopore is at the apex of the sac. 4. Euchroa independencia n. sp. Besides the diagnostic characters, the subgenus 5. Euchroa cupripennis Chaudoir Dyschromus differs from the subgenus Euchroa in that it 6. Euchroa perezi (Darlington) exhibits a much greater range of interspecific variation for 7. Euchroa pedernales n. sp. many features, such as body color, form of the submentum, condition of the elytral striae, elytral intervals and elytral Nitidipennis Group microsculpture, structure of the median lobe of the aedeagus, etc. As there are five times as many species known of the Nitidipennis Subgroup subgenus Dyschromus, this is to be expected, but the gener- 8. Euchroa nitidipennis Putzeys alization holds for the seven species on the island of 9. Euchroa sallei Chaudoir Hispaniola, even though they comprise a monophyletic 10. Euchroa huautla n. sp. assemblage (opaca group). 11. Euchroa cuiyachapa n. sp. 12. Euchroa zongolica n. sp. Habitat.—Species of the subgenus Dyschromus live on the 13. Euchroa lasvigas n. sp. ground under cover in dry to wet montane forests of tropi- 14. Euchroa flohri Bates cal to temperate aspect at elevations of 600 to 3400 m. 15. Euchroa teotitlan n. sp. 16. Euchroa citlaltepetl n. sp. Geographical Distribution.—This subgenus is known in 17. Euchroa perote n. sp. Mexico only from north of the Isthmus of Tehuantepec, 18. Euchroa harrisoni n. sp. where it occurs in the Sierra Madre del Sur in Oaxaca and Guerrero, the complex of mountains in eastern Oaxaca, the Soladevega Subgroup Trans-Volcanic Sierra, and the Sierra Madre Oriental, north 19. Euchroa soladevega n. sp. to southern Tamaulipas. In the West Indies, Dyschromus 20. Euchroa juchatengo n. sp. is known only from the Greater Antillean island of 21. Euchroa nizavaguiti n. sp. Hispaniola, where it is represented in the mountains of both Haiti and the Dominican Republic. Dimidiata Subgroup Chorological Affinities.—The subgenus Dyschromus is 22. Euchroa jalisco n. sp. the only group of New World euchroines that is not repre- 23. Euchroa tenancingo n. sp. sented in South America. Although a wide geographical 24. Euchroa ixtapa n. sp. gap separates the range of Dyschromus from that of its sis- 25. Euchroa dimidiata Chaudoir ter group, the subgenus Euchroa (Fig. 4), nevertheless, the 26. Euchroa atoyac n. sp. range of Dyschromus is overlapped by the ranges of three 27. Euchroa puertogallo n. sp. other euchroine genera: Ophryogaster Chaudoir, Abaris 28. Euchroa filodecaballo n. sp. Dejean, and Pseudabarys Chaudoir. However, most mem- 29. Euchroa chrysophana Bates bers of these groups live at altitude lower than those occu- 30. Euchroa yucuyacua n. sp. pied by Dyschromus, and the adults are smaller. Thus, it 31. Euchroa santacatarina n. sp. seems unlikely that any members of these groups interact in 32. Euchroa zempoaltepetl n. sp. any significant way with species of Dyschromus. 33. Euchroa carbonera n. sp. 34. Euchroa miahuatlan n. sp. Taxonomic Composition and Classification.—This sub- 35. Euchroa suchixtepec n. sp. genus includes 35 species arranged in two monophyletic

KEY TO IDENTIFICATION OF ADULTS OF and posterad of basolateral impressions. Median THE SUBGENUS DYSCHROMUS CHAUDOIR lobe of aedeagus with periostial ventrolateral bulge not evident or barely so (Fig. 22b)...... The key to adults of species of the subgenus Dyschromus is ...... 4. E. independencia n. sp., p. 29 based largely on external traits, and so applies to both 5'. Elytra coppery in color, with distinct reddish hue (as males and females. It emphasizes body color and other eas- in Fig. 3c), or with bluish or greenish reflection. ily observable features, but certain species are quite similar Labial palpomere 3 broader, in males broadly trian- in external appearance, and so the aedeagus should be gular (as in Fig. 11a) to securiform (as in Fig. 11b), examined if a male is available. A case in point is that in females narrowly triangular (as in Fig. 11c) to among the “Biologia” material identified by experts of the broadly triangular (as in Figs. 11a, 11d). Pronotum day as E. nitidipennis, are specimens of E. cuiyachapa, with surface raised laterad and posterad of basolat- E. citlaltepetl, and E. perote. eral impressions. Median lobe of aedeagus with pro- 1. Submentum not prominent and transgular groove nounced periostial ventrolateral bulge (Fig. 24b). 6 shallow posterad submentum (Fig. 9b). Elytra with shoulders rounded (Fig. 1a). Haiti and Dominican 6(5'). Head rather small, ratio PWm/HW: 1.68–1.69 (Fig. Republic...... 2 8a). Aedeagus with preapical part of median lobe comparatively short, apex subtruncate (Figs. 1'. Submentum at least slightly prominent and transgu- 22d–e). Mt. Basil, northern Haiti...... lar groove at least moderately deeply impressed ...... 5. E. cupripennis Chaudoir, p. 31 posterad submentum (Figs. 9c–d). Elytra with shoulders relatively prominent, humeral angle sharp 6'. Head of usual size, ratio PWm/HW: 1.57–1.62 (Fig. or dentiform (Fig. 1b). Mexico...... 8 8b). Aedeagus with preapical part of median lobe relatively long, right side terminated in a more or 2(1). Dorsum all black or brassy in color (Figs. 2a–b); if less distinct lobe (Figs. 22f–g, 24a–f). Western dorsum distinctly brassy in color, then outer part of Dominican Republic and southern Haiti...... 7 elytral interval 8 and all of interval 9 with mesh microsculpture markedly elongate...... 3 7(6'). Legs reddish brown, somewhat infuscate. Pronotum with midlateral setal puncture in contact with later- 2'. Dorsum bicolored, head and pronotum metallic al marginal bead, latter notched at point of contact green, elytra coppery (Fig. 3c) or brassy (as in Fig. (Fig. 8b). Prosternal intercoxal process with apex 3d); if dorsum appearing uniformly brassy in color, not margined, or just indistinctly margined laterally. then sides of elytra with mesh microsculpture isodi- Aedeagus with preapex of median lobe terminating ametric...... 5 in a weakly developed lobe (Figs. 22f–g). Cordillera Septentrional and C. Central, Dominican Republic. 3(2). Labial palpomere 3 broadly triangular in females, ...... 6. E. perezi (Darlington), p. 32 securiform in males (as in Figs. 11a–b). Elytra with all striae moderately impressed and entire on disc; 7'. Legs nearly black. Pronotum with distinct gap elytral intervals slightly convex...... between midlateral setal puncture and lateral mar- ...... 1. E. tiburonica (Darlington), p. 24 ginal bead, latter entire at level of puncture (Fig. 8a). Prosternal intercoxal process with apex distinctly 3'. Labial palpomere 3 subtriangular in both sexes (Fig. margined at least laterally. Aedeagus with preapex 11e). Elytra with striae 1–6 very shallowly of median lobe terminating in a comparatively well- impressed and interrupted numerous times on disc; developed lobe (Figs. 24a–f). Sierra Baoruco and elytral intervals flat...... 4 Massif de La Selle, southern Dominican Republic and Haiti...... 7. E. pedernales n. sp., p. 33 4(3'). Dorsum dull black, elytra at most with slight brassy sheen (Fig. 2a). Elytra with microsculpture on disc 8(1'). Dorsum entirely metallic bluish-purple...... 9 consisting mostly of elongate sculpticells (as in Figs. 6a–b)...... 2. E. opaca (Chaudoir), p. 25 8'. If head and pronotum metallic bluish-purple, then elytra reddish-purple (Figs. 2c–e)...... 12 4'. Dorsum brassy (Fig. 2b). Elytra with microsculpture on disc consisting of both isodiametric and slightly 9(8). Mandibles with dorsal grooves short (Fig. 9a). transverse sculpticells (as in Figs. 5d–e)...... Submentum very prominent and concave posteri- ...... 3. E. centralis (Darlington), p. 27 orly (Figs. 9d, 10b). Elytral intervals 8 and 9 with surface at most slightly beaded in spots, all or near- 5(2'). Elytra brassy in color (as in Fig. 3d). Labial ly all sculpticells flat. Mesepisternum smooth. palpomere 3 narrower, in males subtriangular (as in Sierra Madre Oriental...... Fig. 11e) to narrowly triangular (as in Fig. 11c), in ...... 18. E. harrisoni n. sp., p. 57 females subtriangular. Pronotum nearly flat laterad

9'. Mandibles with dorsal grooves long (as Figs. 14'. Dorsum with colors at least somewhat brighter 1a–b). Submentum slightly to moderately promi- than above (Figs. 2d–e). Submentum and adjoin- nent, posterior face sloping evenly to transgular ing portion of transgular groove variously devel- groove (as Figs. 9c, 10a). Elytral intervals 8 and 9 oped, but if in doubt about color (i.e. some espe- with entire surface distinctly beaded, sculpticells cially dark specimens of E. perote and E. citlalte- markedly convex. Mesepisternum coarsely punct- petl), then submentum very prominent and con- ate. Sierra de Miahuatlan and vicinity, Oaxaca. 10 cave posteriorly, and transgular groove very deep (Figs. 9d, 10b). Elytral intervals not more than 10(9'). Apparent body length at least 13 mm. Elytra with slightly convex on disc...... 15 microsculpture on disc of elongate sculpticells that are about as deeply impressed as those on intervals 15(14'). Dorsum with colors relatively subdued (Fig. 2d). 8 and 9 (Figs. 6a–c)...... Submentum very prominent and with posterior ...... 19. E. soladevega n. sp., p. 59 face concave, adjoining portion of transgular 10'. Apparent body length not more than 11 mm. Elytra groove very deep (Figs. 9d, 10b)...... 16 with microsculpture on disc of isodiametric sculp- 15'. Dorsum with colors comparatively bright (Fig. 2e). ticells that are considerably more finely impressed Submentum not so prominent as above, posterior than those on intervals 8 and 9...... 11 face evenly sloped, adjoining portion of transgular groove not so deep as above (Fig. 10a)...... 17 11(10'). Submentum moderately prominent (nearly as prominent as in Fig. 9d). Pronotum with hind 16(15). Pronotum comparatively narrow, ratio PWm/HW: angles slightly prominent (as in Fig. 12a). 1.53–1.57. Aedeagus with preapex moderately Aedeagus with preapical part of median lobe mod- asymmetrical in outline, area to left of midline of erately long (Fig. 36d)...... median lobe evident but considerably thinner than ...... 20. E. juchatengo n. sp., p. 61 area to right (Figs. 33a, 33c). Volcan Citlaltepetl. . 11'. Submentum slightly prominent (as in Fig. 9c)...... 16. E. citlalteptel n. sp., p. 55 Pronotum with hind angles obtuse (Fig. 12d). 16'. Pronotum comparatively broad, PWm/HW: 1.5– Aedeagus with preapical part of median lobe long 1.68. Aedeagus with preapex markedly asymmetri- (Fig. 36f)...... 21. E. nizavaguiti n. sp., p. 61 cal in outline, not developed to left of midline of median lobe, stout throughout (Figs. 34a, 34c). 12(8'). Head and pronotum metallic bluish-purple or cop- Cofre de Perote and vicinity...... pery, elytra reddish-purple (Figs. 2c–f). Elytral stri- ...... 17. E. perote n. sp., p. 55 ae deeply impressed, entire...... 13 12'. If head and pronotum coppery, then elytra bronze 17(15'). Pronotum and elytra comparatively narrow (Fig. in color (Fig. 3f), or dorsum entirely metallic green 12c), ratio PWm/Hw: 1.42–1.48, Ew/Hw (Figs. 3a–b) or brassy (Fig. 3e), or coppery, or head 1.71–1.75. Elytral intervals on disc with sculpti- and pronotum metallic green and elytra brassy cells uniformly and moderately finely impressed. (Fig. 3d). Elytral striae shallowly impressed in Median lobe of aedeagus with preapex stout and most individuals, and at least stria 1 in most indi- darkly sclerotized throughout (Fig. 29f)...... viduals interrupted at least twice on disc...... 21 ...... 13. E. lasvigas n. sp., p. 50 17'. Pronotum and elytra of most specimens compara- 13(12). Head and pronotum coppery (Fig. 2f). Prosternal tively broad, ratio PWm/Hw at least 1.48 and intercoxal process not margined. Elytral intervals Ew/Hw at least 1.75, if less (some specimens of E. moderately convex (as in Fig. 15a)...... cuiyachapa) then elytral intervals on disc with ...... 10. E. huautla n. sp., p. 45 sculpticells very finely impressed and in spots 13'. Head and pronotum metallic bluish-purple (Figs. entirely effaced. Median lobe of aedeagus with at 2c–e). Elytral intervals flat to moderately convex least left margin of preapex thinner and less darkly (Figs. 15a–b), if moderately convex then apex of sclerotized than rest of preapex (Figs. 25c, 25f, prosternal intercoxal process margined all around. 28c, 29c, 30c)...... 18 ...... 14 18(17'). Head with transgular groove moderately deep pos- 14(13'). Dorsum with colors very dark in hue (Fig. 2c). terior to submentum (Fig. 9c). Pronotum compara- Submentum slightly prominent, adjoining portion tively broad, 1.32–1.45x wider than long; basolat- of transgular groove moderately deep (as in Fig. eral impressions short (Fig. 12b). Elytra compara- 9c). Elytral intervals moderately convex (Fig. 15a). tively broad, 1.30–1.53x longer than wide. Male ...... 9. E. sallei Chaudoir, p. 40 with labial palpomere 3 securiform (as in Fig.

11b). Median lobe of aedeagus with preapex mod- 21(12'). Head and pronotum coppery in color, with or with- erately long and widest near tip (Fig. 25a)...... out greenish reflection, elytra bronze (Fig. 3f)...... 8. E. nitidipennis Putzeys, p. 37 Elytral striae barely evident and interrupted 18'. Head with transgular groove deep posterior to sub- numerous times, or not evident...... mentum (Fig. 9d). Pronotum in most specimens ...... 30. E. yucuyacua n. sp., p. 78 comparatively narrow, 1.28–1.38× wider than long; 21'. If dorsum distinctly bicolored, then head and basolateral impressions usually at least moderately pronotum metallic green and elytra brassy (Fig. long (as in Fig. 13b). Elytra in most specimens 3d). Elytral striae distinct, continuous or interrupt- comparatively narrow, 1.51–1.66× longer than ed few to numerous times...... 22 wide. Male with labial palpomere 3 broadly triangular in shape (as in Fig. 11d). Median lobe of 22(21'). Dorsum light metallic green, with coppery or red- aedeagus with preapex long and widest towards dish-purple reflections (Fig. 3b). Elytral intervals base (Figs. 28a, 29a, 32a)...... 19 smooth on disc. . . 35. E. suchixtepec n. sp., p. 83 22'. If dorsum uniformly metallic green, then shade 19(18'). Colors brilliant. Prosternal intercoxal process not much darker than above (Fig. 3a). Elytral intervals margined. Elytral intervals on disc with sculpti- with distinct mesh pattern on disc...... 23 cells finely impressed, effaced in spots; intervals nearly flat. Median lobe of aedeagus with preapex 23(22'). Dorsum uniformly coppery or brassy, with or with- slightly offset in dorsal aspect (Figs. 28a, 28d). . . out green reflections (Fig. 3e). Head with postocu- ...... 11. E. cuiyachapa n. sp., p. 46 lar transverse groove almost obsolete medially. 24 19'. Colors more subdued (Fig. 2e). Prosternal inter- 23'. Dorsum uniformly metallic green (Fig. 3a) OR coxal process with apical third at least shallowly head and pronotum metallic green, and elytra margined all around. Elytral intervals on disc with brassy (Fig. 3d). Head with postocular transverse sculpticells uniformly and moderately finely to groove nearly as deep medially as laterally (Fig. finely impressed; intervals somewhat convex. 1b)...... 27 Median lobe of aedeagus with preapex centered along midline in dorsal aspect (Figs. 29a, 32a). . . 24(23). Elytral striae 1-6 or 1-7 each interrupted in several ...... 20 places on disc. Elytral interval 9 and outer half of 8 with surface distinctly beaded, sculpticells 20(19'). Apparent body length, 9.5 mm. Pronotum com- markedly convex. Aedeagus with preapex not paratively narrow with lateral margins markedly developed to left of midline of median lobe (Figs. sinuate posteriorly, ratio PWm/PL: 1.31, 47d, 47f)...... 31. E. santacatarina n. sp., p. 78 PWm/PWb: 1.20. Elytra comparatively narrow, EW/HW: 1.76. Median lobe of aedeagus with 24'. Elytra with not more than striae 1-5 interrupted on most of preapex stout and darkly sclerotized, nar- disc. Elytral intervals 8-9 with surface beading row strip along left margin of preapex very thin confined to apical third of interval, anterior two and nearly colorless (Fig. 29c); left wall of perios- thirds of each interval with sculpticells flat. tial part of median lobe of moderate height (Fig. Aedeagus with preapex well developed to left of 29b); preapical dorsolateral ridge and periostial midline of median lobe (Figs. 44a, 45a)...... 25 ventrolateral bulge both markedly convex in profile (Fig. 29b)...... 25(24'). Pronotum with lateral margins moderately sinuate ...... 12. E. zongolica n. sp., p. 48 posteriorly (as in Fig. 12b), ratio PWm/PWb: 1.15–1.18. Elytral stria 1 interrupted not more than 20'. Apparent body length, 10.5–12.5 mm. Pronotum 6 times on disc, usually 2 or 3 times; other striae in of most individuals comparatively broad and with most individuals entire, stria 2 and/or 3 interrupted lateral margins not so distinctly sinuate posterior- not more than once on disc. Aedeagus with middle ly, ratio PWm/PL: 1.28–1.38, PWm/PWb: part of median lobe distal to prebasal bend of usual 1.12–1.20. Elytra comparatively broad, EW/HW: thickness in lateral aspect (Figs. 44b, 44e)...... 1.79–1.98. Median lobe of aedeagus with right ...... 27. E. puertogallo n. sp., p. 73 half of preapex stout and darkly sclerotized, left half less so (Figs. 32c, 32f); left wall of periostial 25'. Pronotum with lateral margins slightly sinuate or part of median lobe markedly high (Figs. 32b, oblique posteriorly (as in Figs. 13b, 13d), ratio 32e); preapical dorsolateral ridge and periostial PWm/PWb: 1.03–1.16. Elytral stria 1 in most indi- ventrolateral bulge both somewhat flat in profile viduals interrupted from 10 to 30 times on disc, (Figs. 32b, 32e)...... stria 2 from 5 to 26 times, and stria 3 from 1 to 10 ...... 15. E. teotitlan n. sp., p. 52 times on disc, striae on one elytron in some individuals interrupted fewer times. Aedeagus with

middle part of median lobe distal to prebasal bend 31(30'). Pronotum with lateral margins markedly sinuate stout in lateral aspect (Figs. 45b, 45e)...... 26 posteriorly; hind angles very prominent (Fig. 13c). Elytra without basal punctures...... 26(25'). Pronotum with lateral margins slightly sinuate ...... 33. E. carbonera n. sp., p. 81 posteriorly; hind angles slightly prominent (as in 31'. Pronotum with lateral margins slightly to moder- Fig. 13b). Median lobe of aedeagus with preapex ately sinuate posteriorly; hind angles obtuse to of uniform thickness (Figs. 45a, 45c)...... slightly prominent (Figs. 1b, 13b). Elytra with ...... 28. E. filodecaballo n. sp., p. 75 basal punctures present...... 32 26'. Pronotum with lateral margins oblique to slightly sinuate posteriorly; hind angles obtuse (as in Fig. 32(31'). Elytral stria 1 in most individuals interrupted at 13d). Median lobe of aedeagus with far right side least 20 times on disc, considerably less in a few of preapex thicker than area to left, and with individuals, but if so then stria 7 interrupted at least preapical dorsolateral ridge (Figs. 45d, 45f). . . . . 5 times on disc. Apical declivity of elytra nearly ...... 29. E. chrysophana Bates, p. 77 perpendicular (Fig. 14b), or as below. Median lobe of aedeagus of extreme symmetric type 4 (Figs. 27(23'). Head, pronotum and elytra all colored same shade 39a–b, 40a–h)...... 33 of dark metallic green, with or without purple 32'. Elytra stria 1 entire or not interrupted more than reflections (Fig. 3a)...... 28 seven times on disc, stria 7 entire on disc. Apical 27'. Head and pronotum metallic green, elytra brassy, declivity of elytra comparatively gentle (as in Fig. with or without greenish or bluish reflections (Fig. 14a). Median lobe of aedeagus of intermediate 3d)...... 29 asymmetric type 2 (Figs. 30a–c, 48a–c)...... 34

28(27). Pronotum with basolateral impressions long (Fig. 33(32). Submentum moderately prominent and sloped 13a). Mesepisternum with a few indistinct punc- evenly to transgular groove (as in Fig. 9c). Elytra tures. Median lobe of aedeagus with preapex long, with stria 1-7 in most individuals each interrupted more or less angular in outline (Figs. 38a–b). about 25 times on disc, in a few individuals con- Trans-Volcanic Sierra, Jalisco...... siderably less but if so then stria 7 interrupted at ...... 22. E. jalisco n. sp., p. 63 least five times on disc. Median lobe of aedeagus 28'. Pronotum with basolateral impressions at most slender (Figs. 39c-d). . . 24. E. ixtapa n. sp., p. 66 moderately long (as in Fig. 13b). Mesepisternum 33'. Submentum in most individuals very prominent with many distinct punctures. Median lobe of and concave posteriorly (Figs. 9d, 10b), or as aedeagus with preapex moderately long, nearly above in some individuals. Elytra with stria 6 and rounded in outline (Figs. 43a, 43c–e). Sierra 7 entire on disc, or 6 interrupted less than ten times Madre del Sur, Guerrero...... and 7 not more than once on disc. Median lobe of ...... 26. E. atoyac n. sp., p. 71 aedeagus moderately stout (Figs. 40a–h)...... 25. E. dimidiata Chaudoir, p. 68 29(27'). Abdominal sterna V–VII with basal sulcus extended to lateral margins of abdomen; basal sulcus of 34(32'). Elytra with all striae entire; surface beading on sternum VII entirely foveate (Fig. 14d)...... interval 9 confined to posterior third of interval, ...... 23. E. tenancingo n. sp., p. 65 anterior two-thirds of interval with sculpticells flat. 29'. Abdominal sterna V–VII with basal sulcus ending Prosternal intercoxal process not margined. distinctly short of lateral margins of abdomen; Aedeagus with left wall of periostial part of medi- basal sulcus of sternum VII without median fovea an lobe more extensive than right wall, ostium not (Fig. 14c), or distinct gap between median fovea visible in left lateral aspect (Fig. 30b). Cofre de and lateral foveae...... 30 Perote, Veracruz...... 14. E. flohri Bates, p. 50 34'. Elytra with stria 1 interrupted as many as seven 30(29'). Pronotum with lateral margins oblique posteriorly; times on disc, if entire on disc, then stria 1 or 2 hind angles rounded or obtuse (Fig. 13d). Elytra interrupted several times near apex of elytra; without basal punctures...... patches of surface beading present along entire ...... 34. E. miahuatlan n. sp., p. 82 length of interval 9. Prosternal intercoxal process 30'. Pronotum with lateral margins at least slightly sin- with apical third margined laterally or all around. uate posteriorly; hind angles obtuse to prominent Aedeagus with right wall of periostial part of (Figs. 1b, 13b, 13c). Elytra with or without basal median lobe more extensive than left wall, ostium punctures, if absent then hind angles of pronotum visible in left lateral view (Fig. 48b). Cerro very prominent...... 31 Zempoaltepetl, Oaxaca...... 32. E. zempoaltepetl n. sp., p. 80

TAXONOMIC TREATMENT OF ADULTS Chorological Affinities.—The range of the opaca group OF THE SUBGENUS does not overlap with that of its sister group, the nitidipen- DYSCHROMUS CHAUDOIR nis group, since the latter is confined to central Mexico. Opaca Group Taxonomic Composition.—The opaca group contains seven species in two subgroups. Diagnosis.—The opaca group is distinguished from the nitidipennis group by the gentle contours of the submentum and transgular groove, rounded shoulders of the elytra, and Opaca Subgroup the microtrichia of the internal sac of the median lobe of the aedeagus, which are larger, and arranged in single narrow Diagnosis.—The uniformly dark color of the body in com- band around the base of, or in a broad band around the mid- bination with the elongate mesh microsculpture along the dle of the sac. The frontal furrows on the head of most sides of the elytra distinguishes the opaca subgroup from species are either shallow throughout, or the deeply and the cupripennis subgroup. sharply impressed portion does not extend posteriorly to Description.—Color. Dorsum all black (Fig. 2a) or brassy (Fig. 2b). the level of the anterior supraorbital setigerous punctures, Appendages reddish-brown. as it does in all but one member of the nitidipennis group. Microsculpture. Elytra with sculpticells on disc somewhat elongate, moderately deeply impressed, and markedly convex or keeled, or sculp- Description.—Chaetotaxy. Hind coxae with posterior seta absent from ticells on disc isodiametric to slightly transverse, finely impressed, and many individuals. Femur of middle and hind legs with inner seta of pair slightly convex; at sides and apex of elytra sculpticells slightly to located along posteroventral margin absent in many individuals. markedly elongate, and flat, convex, or keeled. Microsculpture. Elytra with surface of disc at least slightly beaded, Head. Frontal furrows shallow, sharply impressed for not more than sculpticells at least slightly convex. half their length, if at all. Head. Frontal furrows variously developed, at one extreme shallow Prothorax. Pronotum without transverse impression between basolat- throughout (Fig. 1a), at other extreme deeply and sharply impressed for eral impressions. Prosternal intercoxal process with entire apical third nearly entire length and with deeply and sharply impressed portion distinctly margined. extended posteriorly to level of anterior supraorbital setigerous punctures Pterothorax. Elytra on disc with striae moderately impressed and (Fig. 8b). Postocular transverse groove obsolete dorsomedially or nearly entire, and intervals moderately convex, or disc of elytra with striae 1-6 so (Fig. 1a). Submentum not prominent, posterior face sloped gently and and at least anterior two-thirds of 7 very faint and interrupted numerous evenly to transgular groove; latter very shallow posterad submentum times, and intervals flat. Mesepisternum smooth to coarsely punctate. (Fig. 9b). Legs. Hind tibia with one or two rows of medial flexile setae. Mouthparts. Mandibles with dorsal grooves long (Fig. 1a). Mentum Abdomen. Sternum VII with all of basal sulcus foveate, or with one with paramedian pits moderate to large in size. median fovea in addition to lateral foveae. Prothorax. Pronotum with midlateral setal puncture in contact with or Male. Median lobe of aedeagus with right third of preapex slightly separated by one-half to two times its diameter from lateral marginal bead thicker and somewhat more darkly sclerotized than rest of preapex; (Figs. 8a–b); basolateral impressions short to moderately long, not extend- preapical dorsolateral ridge not evident (Fig. 18a) or very slightly devel- ed to hind margin of pronotum; transverse impression present between oped (Fig. 19a); two to four primary preapical haemolymph channels basolateral impressions in some individuals of a few species (Fig. 8b). present (Figs. 18b, 19c). Internal sac with large microtrichia in narrow Pterothorax. Elytra with shoulders rounded; basal punctures present band at base of sac (Fig. 19d), microtrichia oval in outline, those on left (Fig. 1a). side transverse. Abdomen. Sternum VII with basal sulcus with median fovea as well Female. Spermathecal gland short and straight, or long and recurved. as lateral foveae, or entire sulcus foveate (as in Fig. 14d). Male. Median lobe of aedeagus with preapex short (Fig. 18a) to mod- Geographical Distribution.—Members of this subgroup erately long (Fig. 24a), broad, right third at least somewhat thicker and are known from southern Haiti (Massifs de La Hotte and de more darkly sclerotized than rest of preapex, if preapical dorsolateral ridge present (Fig. 19a), then flat in profile (Figs. 19b, 24b); periostial La Selle) and the central part of the Dominican Republic part of median lobe with left wall at least as extensive as and entirely con- (Cordillera Central) (Fig. 21). cealing right wall in left lateral view (Fig. 19b); periostial ventrolateral bulge not evident (Fig. 20b) or comparatively basal in position (Fig. 19b) Phylogenetic Relationships.—Although there is some and subangulate in cross-section (as in Fig. 26c); middle part of median evidence for monophyly of the opaca subgroup, other evi- lobe without dorsomedial groove. Internal sac with largest microtrichia in dence indicates that one or more of the species are more broad central band (Fig. 24f), or narrow basal band (Fig. 19d). Female. Spermatheca short and nearly straight (Fig. 17a), or long and closely related to the cupripennis group than they are to recurved (as in Fig. 17b). other members of the opaca subgroup. See phylogenetic analyses below for details. Habitat.—The opaca group appears to be restricted to the vicinity of uplands above 900 m. elevation (Figs. 21, 23). Chorological Affinities.—The geographical range of the opaca subgroup overlaps that of the cupripennis subgroup Geographical Distribution.—The opaca group is con- (cf. Figs. 21, 23). fined to Haiti and the Dominican Republic. Taxonomic Composition.—The opaca subgroup contains Phylogenetic Relationships.—The nitidipennis group is E. tiburonica (Darlington), E. opaca (Chaudoir), and the sister group of the opaca group. E. centralis (Darlington).

Fig. 9.—A, right mandible of E. (Dyschromus) nitidipennis Putzeys, dorsal aspect; B–D, head of subgenus Dyschromus, lower portion, right lateral aspect; B, E. perezi n. sp.; C, E. nitidipennis; D, E. dimidiata Chaudoir, 64.4 km northwest of Ciudad Oaxaca. Only drawing of E. dimidiata to scale. sm, submentum; tgg, transgular groove. Scale bar = 0.8 mm.

1. Euchroa (Dyschromus) tiburonica (Darlington) Apparent body length 11.0M–13.0F mm. Length (n=1 M,2F): head Figs. 1a, 18a–b, 21 0.95F–1.01F, pronotum 2.44M–2.72F, elytra 6.10M–6.90F mm; width: head 2.26M–2.50F, pronotum 3.37M–3.81F, elytra 2.98M–3.30F mm. Body Proportions. Head 2.33M–2.48F× wider than long. Pronotum Dyschromus tiburonicus Darlington 1935:179. Type material. HOLO- F F TYPE male, labeled: “NE foothills/ La Hotte/ 2-4000 ft./ Oct 1.38–1.40 × wider than long. Elytra 1.45–1.50 × longer than wide. 10–24”; “Haiti/ 1934/ Darlington”; “22021/ M.C.Z./ Holotype/ Color. Dorsum black, elytra with slight reddish-purple tint. tiburonicus D.” [red paper] (MCZC). PARATYPES, two female Microsculpture. Head between eyes with sculpticells more deeply specimens labeled same as holotype, except “M.C.Z/ Paratype/ impressed than on pronotum. Elytra with sculpticells on disc, interval 8, tiburonicus,” 1 (MCZC), 1 (BMNH). and apex moderately deeply impressed, slightly elongate, and markedly convex or keeled, interval 9 with sculpticells more isodiametric and Type Locality.—Monte Macaya, Massif de la Hotte, Department of Sud, slightly convex. Haiti. Mouthparts. Labial palpomere 3 securiform in male and broadly triangular in females, in male 1.02 and females 1.26–1.33× longer than wide. Mentum with paramedian pits large. Diagnosis.—This all-black species is distinguished from Prothorax. Pronotum with sides somewhat explanate anterolaterally the other species of the subgenus Dyschromus on the island (Fig. 1a); lateral margins oblique to slightly sinuate posteriorly; hind of Hispaniola, including the only other black one, by the angles obtuse; basolateral impressions moderately long, narrowly entire and moderately impressed elytral striae, and some- impressed, shallow; midlateral setal puncture separated by about twice its what convex elytral intervals. Traits unique to E. tiburoni- diameter from lateral marginal bead. Pterothorax. Elytral striae moderately impressed, entire on disc, stri- ca are that the sculpticells on the head between the eyes are ae 1 and 2 interrupted several times apically; elytral intervals moderately comparatively deeply impressed, the sides of the pronotum convex. Mesepisternum with coarse punctures. are explanate, and there are some foveae present at the base Legs. Hind tibia with two rows of medial flexile setae, with origin of abdominal sterna III and IV. from common groove, outer row with about 10, inner with 3 to 5 setae (as in Fig. 16f). Description.—Body Size. Individuals moderate to relatively large in size. Abdomen. Sterna III–IV with a few foveae at anterior margin.

Fig. 10.—Scanning electron photomicrographs of portion of venter of head of subgenus Dyschromus. A, E. teotitlan n. sp.; B, E. citlaltepetl n. sp. pfs, posterior face submentum; pmp, paramedian pits of mentum; ttg, transgular groove. Scale bar = 0.384 mm.

Sternum VII with entire basal sulcus coarsely foveate. Notes about Type Material.—Two specimens, a male and Male. Middle tibia with inner face slightly expanded near apex. Hind female, in the Oberthür-Chaudoir collection (MNHP), are tibia with medial ridge entire (as in Fig. 16f). Aedeagus with generalized asymmetric type median lobe; preapex short and nearly symmetrical in associated with the following handwritten label that was outline (Figs. 18a–b), no indication of preapical dorsolateral ridge, three pinned originally in the bottom of an insect box: “opaca/ or four primary preapical haemolymph channels present, longest with Chaud/ Haiti/ Cte. Mannerheim.” The male is labeled: origin at extreme right side of ostium (Fig. 18b); periostial ventrolateral “Gen/ Dyschromus/ Chaud” [handwritten]; “Ex Musaeo/ bulge not evident (as in Fig. 20b). Internal sac without constrictions. Chaudoir” [red print]. The female has only the single label Female. Spermathecal gland long, apical portion recurved (Fig. 17b). “Ex Musaeo/ Chaudoir” [red print]. Because of the Material Examined.—Three specimens. Holotype male, and two Chaudoir labels, it seems clear that both specimens were paratype females (MCZC, BMNH). part of the Chaudoir collection before it was purchased and added to by René Oberthür. Collecting Notes.—The type series of E. tiburonica was The species D. opacus was based on a single specimen, found at elevations of about 600 to 1200 m, “under stones a male (Chaudoir, 1835: 430). The female evidently was and logs in damp woods and coffee plantations” obtained subsequently, for Chaudoir (1874: 19) remarked (Darlington 1935). that he now had both sexes, each represented by a single specimen. He noted also that he had been mistaken in Geographical Distribution.—This species is known only believing that the male was from Java or nearby islands; from the the foothills and slopes of Monte Macaya in the rather, it must have been from Haiti (like the female). Massif de la Hotte of extreme southwestern Haiti (Fig. 21). We draw two conclusions: first, the information on the Phylogenetic Relationships.—Euchroa tiburonica could box label probably refers to the female, which must have have a sister group relationship with E. opaca, or it occu- been received after the species was described, and thus can- pies, by itself, the basal position within the opaca sub- not be type; second, the male, with the label bearing its group, or the entire opaca group. See section on phylogeny generic identification, is the specimen referred to in the below. original description and accordingly, we accept it as the holotype of D. opacus. Chorological Affinities.—This is the only species of Dyschromus known from the Massif de la Hotte. Its closest Type Locality.—As noted above, Chaudoir (1835:430) relatives, E. opaca and E. centralis, occur further to the east mistakenly recorded the holotype as having been collected in the Massif de La Selle (Haiti), and the Cordillera Central in the vicinity of Java. The specimen was undoubtedly (Dominican Republic), respectively (Fig. 21). from Haiti (Chaudoir 1874), presumably from somewhere on the Massif de La Selle (Department of Sud Est), which we designate as the type area. 2. Euchroa (Dyschromus) opaca (Chaudoir) Figs. 2a, 7a–b, 11e, 16f, 19a–d, 21 Diagnosis.—Among the three uniformly dark-colored species of the subgenus Dyschromus on the island of Dyschromus opacus Chaudoir 1835:430. Type material. HOLOTYPE Hispaniola, E. opaca is larger than E. centralis, and the dor- male (MNHP). For details, see “Notes about type material” below. sum is black rather than brassy in color; also, the Darlington, 1935:180; 1939:89. Euchroa opaca Chaudoir: Chaudoir, 1874:16–20; Tschitschérine, microsculpture on the disc of the elytra consists mostly of 1898:60. elongate rather than isodiametric sculpticells. It is readily

Fig. 11.—A–E, left labial palpomere 3 of subgenus Dyschromus, ventral aspect; A–B, E. sallei Chaudoir, female and male, respectively; C–D, E. harrisoni n. sp., female and male; E, E. opaca (Chaudoir), male; F, left maxillary palpomere 3 and 4 (mp4) of E. harrisoni, female, ventral aspect. Scale bar = 0.5 mm.

distinguished from the other all-black species, E. tiburoni- 11e). Mentum with paramedian pits large. ca, by the greatly interrupted and shallowly impressed ely- Prothorax. Pronotum with lateral margins entire or slightly or moderately sinuate posteriorly; hind angles obtuse; basolateral impressions tral striae and flat elytral intervals. variable, at one extreme moderately long and narrowly and shallowly impressed (as in Fig. 1a), at other extreme short, deeply impressed and Description.—Body Size. Individuals moderate in size. Apparent body nearly punctiform (as in Fig. 2b); midlateral setal puncture separated by length 10.0M–11.5F mm. Length (n=6M,4F): head 0.89(0.83–0.95M), M M M M its diameter from lateral marginal bead. pronotum 2.35(2.20 –2.52 ), elytra 5.59(5.05 –5.85 ) mm; width: Pterothorax. Elytral striae 1-6 and at least anterior two-thirds of 7 head 2.24(2.06M–2.36M), pronotum 3.25(2.88M–3.47F), elytra 3.98 M F very slightly impressed and interrupted numerous times on disc; elytral (3.55 –4.30 ) mm. intervals flat. Mesepisternum impunctate. Body Proportions. Head in males 2.46(2.41–2.48) and females Legs. Hind tibia with two rows of medial flexile setae, with origin 2.55(2.5–2.71)× wider than long. Pronotum 1.39(1.31M–1.45F)× wider M from common groove, outer row with about 10, inner with 3 to 5 setae than long. Elytra 1.41(1.31–1.52 )× longer than wide. (Fig. 16f). Color. Dorsum black, elytra with faint brassy sheen (Fig. 2a). Abdomen. Basal sulcus of sternum VII with small median fovea, as Microsculpture. Head between eyes with sculpticells about as finely well as coarse lateral foveae. impressed as on pronotum. Elytra with sculpticells deeply impressed and Male. Middle tibia with inner face slightly expanded near apex. Hind markedly convex or keeled (Figs. 7a–b), sculpticells on disc slightly tibia with medial ridge represented at very base by a few slightly elevat- elongate, outer half of interval 8, interval 9, and apex of elytra with sculp- ed tubercles (Fig. 16f). Aedeagus with generalized asymmetric type ticells markedly elongate. median lobe; preapex short and subangulate in outline at apex (Figs. 19a, Mouthparts. Labial palpomere 3 subtriangular, in males 19c), preapical dorsolateral ridge slightly developed (Fig. 19a), two pri- 2.02(1.96–2.12) and females 2.13(1.93–2.23)× longer than wide (Fig.

mary preapical haemolymph channels present, longest with origin at slightly transverse, slightly convex; outer part of interval 8, interval 9, extreme right side of ostium, broadest with origin to left of midline (Fig. and apex of elytra with sculpticells markedly elongate, flat. 19c); periostial ventrolateral bulge somewhat prominent in profile (Fig. Mouthparts. Labial palpomere 3 subtriangular, in males 2.0–2.09 and 19b). Internal sac constricted at middle (Fig. 19d). female 2.0× longer than wide. Mentum with paramedian pits moderate in Female. Spermathecal gland long, apical portion recurved (as in Fig. size. 17b). Prothorax. Pronotum with lateral margins slightly sinuate posteriorly; hind angles obtuse; basolateral impressions short, deeply impressed Material Examined.—Twelve specimens. HAITI: [La] Visite & and nearly punctiform (Fig. 2b); midlateral setal puncture separated by vic[inity,] La Selle Range, 5–7000 ft., Sept. 16–23 1934, Darlington, 4M, one-half its diameter from lateral marginal bead. 2F (MCZC), 2M (BMNH); same as previous label, except Morne Pterothorax. Elytral striae 1-6 and at least anterior two-thirds of 7 Tranchant, Mt. La Selle, 6000 ft., XI-11 1934, 2M, 2F (MCZC). very slightly impressed and interrupted numerous times on disc; intervals flat. Mesepisternum impunctate. Collecting Notes.—Euchroa opaca has been collected at Legs. Hind tibia with one row of medial flexile setae. elevations of about 1500 to 2100 m. Abdomen. Sternum VII with entire basal sulcus coarsely foveate. Male. Middle tibia with inner face not expanded near apex. Hind tibia with medial ridge entire. Aedeagus with generalized asymmetric type Geographical Distribution.—This species is known only median lobe; preapex very short and slightly asymmetrical in outline from the Massif de La Selle in southern Haiti (Fig. 21). (Figs. 20a, 20c), no indication of preapical dorsolateral ridge (Fig. 20a), three or four primary preapical haemolymph channels present, longest Phylogenetic Relationships.—Euchroa opaca could have with origin at extreme right side of ostium (Fig. 20c); periostial ventro- a sister group relationship with either E. tiburonica or lateral bulge not evident (Fig. 20b). Internal sac with bulge on right side at middle. E. centralis, or it may occupy by itself, the second most Female. Spermathecal gland short, straight. basal position within the entire opaca group. For details, see phylogenetic analyses, below. Material Examined.—Holotype, and one male (BMNH) and one female (MCZC) paratype from type locality, and male paratype from Valle Chorological Affinities.—Euchroa opaca is the only Nuevo (MCZC), and one additional female specimen. LA VEGA. 8 km. member of the subgenus Dyschromus known from the S. Constanza, Berlese[,] cloud forest litter, 02.IX.1997, P.W. Kovarik (WIBF). Massif de La Selle. It has not yet been found in the Sierra Baoruco, the eastward continuation of the Massif de La Collecting Notes.—The holotype and all but one of the Selle in the Dominican Republic. It appears to occupy the paratypes were found in accumulations of dead leaves, in middle part of the geographical range of the opaca sub- depressions on the ground near an arroyo in damp forest, at group (Fig. 21). an elevation of about 900 m; the other paratype was under a log in a small patch of cloud forest, at about 1200 m ele- 3. Euchroa (Dyschromus) centralis (Darlington) vation (Darlington 1939). Figs. 2b, 20a–c, 21 Geographical Distribution.—Euchroa centralis is known Dyschromus centralis Darlington 1939:88. Type material. HOLOTYPE only from the vicinity of the Loma Vieja in the Cordillera male, labeled: “Constanza/ Aug. '38, Dom. Rep/ 3–4,000 ft./ Central of the Dominican Republic (Fig. 21). Darlington”; “23511/ M.C.Z./ Holotype/ centralis D.” [red paper] (MCZC). PARATYPES, same labels as holotype, except “M.C.Z./ Phylogenetic Relationships.—This species could be most M F Paratype/ centralis,” 3 , 1 (MCZC, BMNH). “Valle Nuevo/ SE closely related to E. opaca, or it has a sister group relation- Constanza/ Aug.'38, Dom. Rep/ c[a]. 7,000 ft., Darl.”; “M.C.Z./ Paratype/ centralis” [red paper], 1M (MCZC). ship with the entire cupripennis subgroup. See section below on phylogeny for details. Type Locality.—Constanza, Province of La Vega, Dominican Republic. Chorological Affinities.—Euchroa centralis is the only Diagnosis.—Euchroa centralis is the smallest species of species of the opaca subgroup known from the Cordillera the subgenus Dyschromus on the island of Hispaniola. It is Central. The other two members of this subgroup live in the further distinguished from the others by its uniform brassy mountains along the south coast of Haiti (Fig. 21). color, and by the microsculpture on the disc of the elytra, which consists of finely impressed, isodiametric to slightly transverse sculpticells. Cupripennis Subgroup

Description.—Body Size. Individuals relatively small. Apparent body Diagnosis.—The metallic green head and pronotum, and length 6.5M–9.5F mm. Length (n=3M, 1F): head 0.71M–0.79F, pronotum brassy or coppery-colored elytra in combination with the 1.81M–2.10F, elytra 4.20M–5.00F mm; width: head 1.65M–2.00F, prono- isodiametric mesh microsculpture along the sides of the tum 2.46M–3.06F, elytra 2.90M–3.55F mm. elytra distinguish the cupripennis subgroup from the opaca Body Proportions. Head 2.31M–2.53F× wider than long. Pronotum 1.36M–1.45F× wider than long. Elytra 1.41F–1.48M× longer than wide. subgroup. Color. Dorsum brassy (Fig. 2b). Microsculpture. Head between eyes with sculpticells more finely Description.—Color. Dorsum of head and pronotum metallic green, impressed than on pronotum, nearly effaced in spots. Elytra with sculpti- with or without bluish or brassy cast, elytra coppery (Fig. 3c) or brassy cells moderately finely impressed, sculpticells on disc isodiametric to (as in Fig. 3d), with greenish reflection or not. Antennomeres 1–3, labial and maxillary palps, and legs reddish brown, slightly infuscate to

Fig. 12.—Pronotum of subgenus Dyschromus, right side. A, E. independencia n. sp.; B, E. nitidipennis Putzeys, holotype; C, E. lasvigas n. sp.; D, E. nizavaguiti n. sp. Scale bar = 1 mm.

nearly black, with metallic green reflection. very shallowly impressed and interrupted numerous times on disc; inter- Microsculpture. Elytra with sculpticells deeply impressed or moder- vals flat. Mesepisternum with few shallow punctures. ately so, most sculpticells on disc and all at sides isodiametric, some on Legs. Hind tibia with one row of medial flexile setae. disc slightly elongate, sculpticells on disc moderately convex, sculpti- Abdomen. Sternum VII with basal sulcus with coarse median fovea, cells at sides and apex of elytra markedly convex. in addition to lateral foveae. Head. Frontal furrows deeply and sharply impressed for nearly entire Male. Median lobe of aedeagus with right third slightly to consider- length, deeply and sharply impressed portion extended posteriorly to ably thicker and more darkly sclerotized than rest of preapex (Figs. 22c, level of anterior supraorbital setigerous punctures (Figs. 8b) or not (Fig. 22g), part of middle third of preapex much thinner than rest of preapex in 8a). some species (Figs. 22e, 22g), preapical dorsolateral ridge slightly to Prothorax. Pronotum with transverse impression between basolater- markedly developed (Figs. 22d, 22f), two primary preapical haemolymph al impressions present or not. Prosternal intercoxal process with apical channels present, longest with origin at extreme right side of ostium, third distinctly margined or not. broadest with origin to left of midline (Fig. 22e). Internal sac with large Pterothorax. Elytral striae 1-6 and at least anterior two-thirds of 7 microtrichia arranged in broad central band extended entirely around sac,

microtrichia mostly subtriangular in outline, at least some on left side pronotum by the brassy rather than coppery-colored elytra, more elongate (Fig. 24f). by the pronotum which is unusually flat posterad and laterad of the basolateral impressions, and by labial palpomere Geographical Distribution.—The cupripennis subgroup 3, which is narrower than in the other species. Certain spec- is known from all of the major mountain systems in the imens that have a pronounced brassy cast to the color of the western part of the Dominican Republic, and also from head and pronotum (see below) could be confused with northern and southern Haiti, but not from the central part E. centralis (opaca subgoup), but are readily separable by (Fig. 23). the pronotal traits, and also by elytral microsculpture (see Phylogenetic Relationships.—Most evidence indicates key). The median lobe of the aedeagus most resembles that that the cupripennis subgroup is monophyletic, and the of E. cupripennis, but the preapex is not unusually thin api- opaca subgroup, or some portion of that subgroup is the comedially, and the periostial ventrolateral bulge is not evi- sister group. For details, see phylogenetic analyses, below. dent or barely so. Description.—Body Size. Individuals small to moderate in size. Apparent Chorological Affinities.—The geographical range of body length 9.5–11.5F mm. Length (n=10M,10F): head 0.87 the cupripennis subgroup overlaps with that of the (0.77F–0.95F), pronotum 2.35 (2.13F–2.63F), elytra 5.81 (5.19F–6.69F) opaca subgroup (cf. Figs. 21, 23). mm; width: head 2.09 (1.81F–2.44F), pronotum 3.37 (3.03F–3.69F), elytra in males 3.90 (3.69–4.06) and females 4.17 (3.66–4.56) mm. Taxonomic Composition.—Four species are included Body Proportions. Head 2.39 (2.26M–2.66F)× wider than long. Pronotum 1.44 (1.38M–1.50F)× wider than long. Elytra 1.46 in this subgroup: E. independencia n. sp.; E. cupripen- (1.31F–1.50M)× longer than wide. nis Chaudoir; E. perezi (Darlington); and E. pedernales Color. Dorsum of head and pronotum clear metallic green (as in Fig. n. sp. 3d), or with pronounced bluish or brassy cast, elytra brassy and with green reflection or not. Appendages somewhat infuscate. Microsculpture. Head between eyes with sculpticells as well- 4. Euchroa (Dyschromus) independencia, new species developed as on pronotum. Figs. 12a, 22a–c, 23 Head. Frontal furrows deeply and sharply impressed for about half distance to anterior supraorbital setigerous punctures (as in Fig. 8a). Type Material.—Seventy-eight specimens, labeled as follows. HOLO- Mouthparts. Labial palpomere 3 narrowly triangular to subtriangular TYPE male, “DOMINICAN REPUBLIC: Independencia, Sierra/ de in males, subtriangular in females, in males 1.80 (1.67–1.96) and females Neiba near crest,/ 5.5 km NNW Angel Feliz/ 18-41N, 71-47W. 1750m”; 2.10 (1.92–2.52)× longer than wide. Mentum with paramedian pits of “21–22 July 1992/ J. Rawlins, S. Thompson/ C. Young, R. Davidson/ moderate size. Dense cloud forest,” (CMNH). PARATYPES, same labels as holotype, Prothorax. Pronotum unusually flat laterad and posterad of basolater- 1M, 1F (CMNH); the same, except “...Sierra/ de Neiba just south/ of crest, al impressions; lateral margins slightly sinuate and only slightly conver- 5 km NNW/ Angel Feliz, 1780 m,” “18-41N, 71-47W/ 13–15 October gent posteriorly (Fig. 12a); hind angles slightly prominent; basolateral 1991/ J. Rawlins, R. Davidson/ C. Young, S. Thompson/ cloud forest,” 1F impressions short, narrowly and faintly or more broadly and deeply (CMNH); the same, except “...Sierra de Neiba/ south slope near summit,/ impressed; no transverse impression between basolateral impressions; 4.0 km N Angel Feliz,/ 18-40-21N, 71-46-05W”; “1825m, 1–2 Apr 2004, midlateral setal puncture separated by one-half its diameter from lateral J./ Rawlins, C. Young, R./ Davidson, broadleaf cloud/ forest without pine, marginal bead. Prosternal intercoxal process with entire apical third dis- hand/ collected. Sample 34243,” 2M, 4F (CMNH). “DOMINICAN tinctly margined all around. REPUBLIC: San/ Juan. Sierra de Neiba,/ Sabana del Silencio,/ 10.0 km Pterothorax. Elytral striae 1-7 shallowly impressed on disc but evi- SSW El Cercado/ 18-39-07N, 71-33-21W,/ 2009 m, 20 June 2003”; “J. dent without magnification. Rawlins, C. Nunez, R./ Davidson, C. Young, P. Acevedo, M de la Cruz/ Male. Middle tibia with inner face slightly expanded near apex. Hind cloud forest along Dan-/ thonia savannah. hand/ collected. Sample tibia with medial ridge entire. Aedeagus with generalized asymmetric type 33242,” 13M, 4F (CMNH); the same, except “...Neiba,/ 9.4 km SSW El median lobe; preapex very short and slightly asymmetrical in outline Cercado,/ 18-39-18N, 71-32-51W,/ 1973 m, 22 June 2003”; “...meadow (Figs. 22a, 22c), right third of preapex slightly thicker and somewhat more near mature pine/ forest, hand collected, Sample 32242,” 10M, 6F darkly sclerotized than rest of preapex (Fig. 22c), preapical dorsolateral (CMNH); the same, except “...Neiba,/ Sabana del Silencio,/ 10.0 km SSW ridge slightly developed (Fig. 22a); periostial ventrolateral bulge not evi- El Cercado/ 18-39-07N, 71-33-21W,/ 2009 m, 16–17 Nov 2004”; “J. dent or barely so (Fig. 22b). Internal sac with bulge on right side at base Rawlins, V. Verdecia,/ C. Young, C. Nunez, W. A./ Zanol, cloud forest and left side at middle. along/ Danthonia savannah. hand/ collected. Sample 33245,” 19M, 5F (CMNH); the same, except “...Neiba,/ 9.4 km SSW El Cercado,/ 18-39- Material Examined.—The type series. 18N, 71-32-51W,/ 1973 m, 18–19 Nov 2004”; “...meadow near mature/ pine forest, hand/ collected , Sample 32245,” 6M, 4F (CMNH). Remarks.—Specimens from the vicinity of El Cercado “DOMINICAN REPUBLIC: Elias/ Pina. Sierra de Neiba,/ 9.1 km WSW vary in appearance because the green color of the prono- Hondo Valle,/ 18-41-38N, 71-46-56W,/ 1856m, 26 November 2004”; “J. tum, or head and pronotum often has a brassy cast; in 12 Rawlins, C. Young,/ C. Nunez, V. Verdecia,/ W. Zanol, wet montane/ for- out of the 67 specimens it is so pronounced that at first est with pine, hand/ collected, Sample 31245,” 1F (CMNH). sight these individuals appear to be uniformly brassy in Type Locality.—5.5 km NNW of Angel Feliz, Province of color dorsally. Otherwise, El Cercado specimens do not Independencia, Dominican Republic. differ in any way from specimens collected near Angel Feliz. Diagnosis.—Euchroa independencia is distinguishable from the other three species of the subgenus Dyschromus Collecting Notes.—Specimens of E. independencia have on the island of Hispaniola with a metallic green head and been collected in cloud forest, wet montane forest with

Fig. 13.—Pronotum of subgenus Dyschromus, right side; A, E. jalisco n. sp.; B, E. zempoaltepetl n. sp.; C, E. carbonera n. sp.; D, E. miahuatlan n. sp. Scale bar = 1 mm.

pine, and adjoining meadows, at elevations of about 1750 Phylogenetic Relationships.—Most evidence indicates to 2010 m. that E. independencia occupies the basal position in the cupripennis subgroup. See section below on phylogeny for Geographical Distribution.—E. independencia is known details. from several localities in the western and central parts of the Sierra de Neiba, all in the Dominican Republic Chorological Affinities.—E. independencia is the only (Fig. 23). member of the subgenus Dyschromus presently known from the Sierra de Neiba. Other species belonging to the Bionomics.—Some of the specimens collected in the vicin- cupripennis subgroup occur in the mountains to the north ity of El Cercado on June 20 or 22, 2003, were teneral. and south (Fig. 23).

Fig. 14.—Subgenus Dyschromus, posterior portion of body. A, E. carbonera n. sp.; B–C, E. dimidiata Chaudoir; D, E. tenancingo n. sp., from type- locality. A–B, right lateral aspect. C–D, ventral aspect. apd, apical declivity; bs, basal sulcus; fsp, foveate setose puncture; st VII, sternum 7. Scale bar = 2.0 mm.

5. Euchroa (Dyschromus) cupripennis Chaudoir Diagnosis.—Euchroa cupripennis is distinguishable from Figs. 8a, 22d–e, 23 the other two species of the subgenus Dyschromus on the island of Hispaniola with a metallic green head and pronotum and coppery colored elytra, E. perezi and E. peder- Euchroa cupripennis Chaudoir 1874:19. Type material. HOLOTYPE female, labeled: “Ex Musaeo/ Chaudoir” [red print], in the nales, by the comparatively small head (see key). It is also Oberthür-Chaudoir collection (MNHP). The holotype is associat- separable from E. perezi by the pronotal midlateral setal ed with the following handwritten label that was pinned originally puncture which is distinctly separated from the lateral mar- in the bottom of an insect box: “cupripennis/ Chaud/ Haiti/ A. ginal bead, and from E. pedernales by its lighter colored Deyrolle.” Tschitschérine, 1898:60. Dyschromus cupripennis (Chaudoir): Darlington, 1935:180; 1939:88. legs. The median lobe of the aedeagus most resembles that of E. perezi, but the preapex is shorter. Notes about Type Material.—Chaudoir (1874:19) recorded in association with the original description of this species, a single female, from Description.—Body Size. Individuals moderate in size. Apparent body Haiti, received from the famous Parisian insect dealer, A. Deyrolle. length 10.5F–11.0F mm. Length (n=1M,2F): head 0.85M–0.93F, pronotum 2.44M–2.56F, elytra 5.60–6.00F mm; width: head 2.02M–2.26F, pronotum Type Locality.—The type area, as indicated in the original description is 3.41M–3.81F, elytra 3.90M–4.45F mm. “Haiti.” We restrict the type locality to Mount Basil, south of Ennery, Body Proportions. Head 2.32M–2.43F× wider than long. Pronotum Department of Artibonite. 1.40M–1.49F× wider than long. Elytra 1.35F–1.44M× longer than wide.

Color. Dorsum of head and pronotum metallic green with bluish coppery colored elytra, E. cupripennis and E. pedernales, reflection, elytra bright coppery in color and with pronounced greenish to by the pronotal midlateral setal puncture, which impinges reddish tint. Appendages somewhat infuscate. Microsculpture. Head between eyes with sculpticells somewhat upon the lateral marginal bead, and by the prosternal inter- effaced. coxal process, which at most is only distinctly margined lat- Head. Frontal furrows with deeply and sharply impressed portion erally. The median lobe of the aedeagus most resembles that extended to level of anterior supraorbital setigerous punctures (as in Fig. of E. pedernales, but the right side of the preapex does not 8b) or not (Fig. 8a). terminate in a lobe. Mouthparts. Labial palpomere 3 broadly triangular in male, broadly to narrowly triangular in females, in male 1.24 and females 1.38–1.75× Description.—Based upon the type series. longer than wide. Mentum with paramedian pits of moderate size. Body Size. Individuals moderate in size. Apparent body length Prothorax. Pronotum with lateral margins entire to slightly sinuate 11.0 mm. Length (n=1M,2F): head 0.95F–1.03M, pronotum and moderately convergent posteriorly (Fig. 8a); hind angles obtuse; 2.40F–2.48M, elytra 6.10F–6.20 mm; width: head 2.26F–2.34M, basolateral impressions short, narrowly and deeply impressed to nearly pronotum 3.59F–3.75M, elytra 4.30–4.35F mm. punctiform; shallow transverse impression between basolateral impres- Body Proportions. Head 2.27M–2.42F× wider than long. sions; midlateral setal puncture separated by one-half its diameter from Pronotum 1.50F–1.51M× wider than long. Elytra 1.42F–1.44M× lateral marginal bead. Prosternal intercoxal process with entire apical longer than wide. third distinctly margined. Color. Dorsum of head and pronotum metallic green with bluish Pterothorax. Elytral striae 1-7 very shallowly impressed on disc, but reflection, elytra bright coppery with slight greenish tint. Appendages evident without magnification. somewhat infuscate. Male. Middle tibia with inner face not expanded near apex. Hind tibia Microsculpture. Head between eyes with sculpticells somewhat with basal half of medial ridge represented by a row of slightly developed effaced. subangulate tubercles. Aedeagus with intermediate asymmetric type 1 Head. Frontal furrows with deeply and sharply impressed portion median lobe; preapex short, right third of preapex slightly thicker and extended to level of anterior supraorbital setigerous punctures (Fig. somewhat more heavily sclerotized than rest of preapex and terminating 8b). in a slight lobe (Figs. 22d–e), middle third of preapex along distal mar- Mouthparts. Labial palpomere 3 securiform in male, narrowly tri- gin very thin compared to rest of preapex and colorless (Fig. 22e), preapi- angular in females, in male 1.08 and females 1.53–1.87× longer than cal dorsolateral ridge slightly developed (Fig. 22d); periostial ventrolat- wide. Mentum with paramedian pits of moderate (n=1) to large (n=1) eral bulge prominent in profile (as in Fig. 24b). Internal sac with bulge on in size. left side at base. Prothorax. Pronotum with lateral margins evenly and very markedly curved inward posteriorly (Fig. 8b); hind angles obtuse; Material Examined.—Three specimens. HAITI: Mt. Basil, N Haiti, basolateral impressions short and narrowly and deeply impressed, to 4700 ft., Sept 9 1934, Darlington (1M, 1F, MCZC; 1F, BMNH). nearly punctiform; shallow transverse impression between basolateral impressions; midlateral setal puncture impinging upon lateral mar- Collecting Notes.—Euchroa cupripennis has been collect- ginal bead. Prosternal intercoxal process not margined. ed at about 1400 m. elevation. Pterothorax. Elytral striae 1-7 very shallowly impressed on disc, not or barely evident without magnification. Geographical Distribution.—This species is known only Male. Middle tibia with inner face not expanded near apex. Hind from Mount Basil in northern Haiti (Fig. 23). tibia with basal half of medial ridge represented by row of slightly prominent subangulate tubercles. Aedeagus with extreme asymmetric type 1 median lobe; preapex moderately long (Figs. 22f–g), right third Phylogenetic Relationships.—Euchroa perezi and E. ped- of preapex considerably thicker and more darkly sclerotized than left ernales together comprise the sister group of E. cupripennis. two thirds and terminated in distinct lobe (Figs. 22f–g), much of middle third of preapex very thin compared to rest of preapex and color- Chorological Affinities.—E. cupripennis is the only less (Fig. 22g), preapical dorsolateral ridge distinctly developed (Fig. species of the subgenus Dyschromus known from northern 22f) and flat in profile (as in Fig. 24b); periostial ventrolateral bulge prominent in profile (as in Fig. 24b). Internal sac with bulge on left Haiti. It is isolated by considerable distances from its clos- side at base. est relatives, E. perezi and E. pedernales (Fig. 23). Material Examined.—Holotype, and two paratypes (1, MCZC; 1, BMNH), and one additional female specimen. LA VEGA: P[arque] 6. Euchroa (Dyschromus) perezi (Darlington) N[acional]. A[rmando]. Bermudez, [La] Cienaga, 19.VII-2.VIII.95, 1010 Figs. 8b, 9b, 22f–g, 23 m, trop. evgrn. for., FIT S. + J. Peck, 95-33, (CMNC). Remarks.—The specimen from La Cienaga differs in Dyschromus perezi Darlington 1939:88. Holotype male, labeled “Mt. several respects from the others. Most notably, the lateral Diego de/ Ocampo, Dom. Rep/ 3–4,000 ft., July '38/ Darlington"; margins of the pronotum are very slightly sinuate posteri- “23510/ M.C.Z./ Holotype/ perezi D.” [red label], (MCZC). PARATYPES, three females, labeled same as holotype, except orly (as in Fig. 3c); this is in contrast to specimens of the “M.C.Z./ Paratype/ perezi,” (MCZC). type series which have the lateral margins more evenly and markedly curved inwards posteriorly than in any Type Locality.—Mount Diego de Ocampo, Province of Santiago, other member of the subgenus Dyschromus. In addition, Dominican Republic. the La Cienaga specimen is larger than the others: appar- Diagnosis.—Euchroa perezi is separable from the other two ent body length 12.0 cm; length of pronotum 2.56 and ely- species of the subgenus Dyschromus on the island of tra 6.63 mm, width of head 2.47, pronotum 3.91, and ely- Hispaniola with a metallic green head and pronotum and tra 4.69 mm. Also, the prosternal process is slightly margined laterally.

Fig. 15.—Scanning electron photomicrographs of posterolateral portion of elytra of subgenus Dyschromus. A, E. sallei Chaudoir; B, E. soladevega n. sp. i9, interval 9. s5 and s6, striae 5 and 6. Scale bar = 0.5 mm.

Collecting Notes.—The type series of E. perezi was “taken PARATYPES, “DOMINICAN REPUBLIC:/ Pedernales. 5 km NE/ Los under trash and leaves in a tiny banana grove at about 3,500 Arroyos, 1680 m./ 18-15N, 71-45W”; “20 October 1991/ J. Rawlins, R. Davidson/ C. Young, S. Thompson/ cloud forest,” 1F (CMNH). ft. altitude” (Darlington 1939). “DOM:REP.: Pedernales/ 37 km N Pedernales,/ Los Arroyos, 1400m/ cloud forest sweeps/ 2.XI.91/ Masner & Peck, 91–360,” 1F (CMNC). Geographical Distribution.—The type series of E. perezi “REPUBLICA DOMINICANA:/ Pedernales, P.N. Sierra de/ Bahoruo, is from Mt. Diego de Ocampo in the Cordillera Las Abejas/ (18o9'8"N,71o37'334"E),/ 1316 m/ 2 September 2001, coll. Septentrional, the most northern mountain system in the K.A/ ”; “Photographed/ 2002-2003/ CBSD”; “28649 (MHND),” 1M Dominican Republic (Fig. 23). The additional specimen is (MHND); “Monteada Nueva/ Barahona, PROV./ BARAHONA, R. D./ 14-VI-1980/ Col. Inchaustegui,” and with “M.N.H.N.” printed sideways from the Cordillera Central; given that it is conspecific along right border of label; “18904,” 1M (MHND); the same, except with the others, E. perezi is the only Hispaniolan species “18907,” 1F (MNHD). “DOMINICAN REP.: Prov./ Barahona, nr. of the subgenus Dyschromus that is known to occur in Filipinas/ Larimar Mine: 20-26-/ VI-1992: R.E. Woodruff,” 1F (FSCA). more than one mountain system. “DOMINICAN REPUBLIC:/ Barahona. Eastern Sierra/ Bahoruco, Reserva Cachote/ 12.8 km NE Paraiso,/ 18-05-54N, 71-11-21W,/ 1230 m, 19-21 May 2004,” C. Young, C. Nunez,/ J. Rawlins, J. Fetzner,/ cloud Phylogenetic Relationships.—The sister species of E. forest with tree/ ferns, hand collected,/ Sample 44244”; “Carnegie perezi is E. pedernales. Museum Specimen Number CMNH-367,031,” 1M (CMNH). “Haiti: Kenskoff: Belot-[Le]Montcel/ N 18o27'11.3" W 72o21'06.4" 20-21-X- Chorological Affinities.—Euchroa perezi is the only 2006/ 1566m col. S. Crews & E. Fernandez/ SCC06 076 under rocks, on species of the subgenus Dyschromus known from the buildings, walls at,” 1F (EMEC). Cordillera Septentrional, and the only member of the Type Locality.—1 km south of Los Arroyos, Province of Pedernales, cupripennis subgroup that has been found in the Cordillera Dominican Republic. Central. There is a gap between the ranges of E. perezi and its closest relative, E. pedernales, which lives further south Diagnosis.—Euchroa pedernales is separable from the in the Sierra de Baoruco (Fig. 23). Neither species has other two species of the subgenus Dyschromus on the ever been found in the intervening uplands (Sierra de island of Hispaniola with a metallic green head and Neiba) despite intensive collecting there in recent years, pronotum and coppery colored elytra, E. cupripennis and which did yield numerous specimens of another member E. perezi, by its darker-colored appendages, and by the of the cupripennis subgroup, E. independencia. right side of the preapical part of the median lobe of the aedeagus, which terminates in a distinct lobe. Also, the head is larger than in E. cupripennis, and the pronotal 7. Euchroa (Dyschromus) pedernales, new species midlateral setal puncture is distinctly separated from the Figs. 3c, 17a, 23, 24a–f lateral marginal bead, unlike in E. perezi. Type Material.—Nine specimens, labeled as follows. HOLOTYPE Description.—Body Size. Individuals moderate to large in size. Apparent male, “DOMINICAN REPUBLIC:/ Pedernales. 1 km S/ Los Arroyos, body length 11.5–12.5F mm. Length (n=4M,5F): head 0.89F–1.03, prono- 1125 m./ 18-14N, 71-45W”; “18 October 1991/ R. Davidson, C. Young,/ tum 2.56F–2.74F, elytra 6.28F–6.85F mm; width: head 2.28F–2.52F, S. Thompson, J. Rawlins/ Second growth forest,” (CMNH).

Fig. 16.—Left tibia of males of subgenus Dyschromus, distal portion, dorsal aspect. A, middle tibia, E. dimidiata Chaudoir, 64.4 km n.w. of Oaxaca; B–F, hind tibia; B, E. dimidiata, as above; C, E. centralis (Darlington); D, E. nitidipennis Putzeys; E, E. chrysophana Bates; F, E. opaca (Chaudoir). dmg, dorsomedial groove; dmrs, dorsomedial row of setae; lrs, lateral row of setae; mg, medial groove; mri, medial ridge; mrfs, medial row of flexile setae. Only drawings of E. dimidiata are to scale. Scale bar = 0.5 mm.

pronotum 3.66F–4.07F, elytra 4.38F–4.85F mm. Chorological Affinities.—Euchroa pedernales is the only Body Proportions. Head 2.31M–2.59F× wider than long. Pronotum member of the cupripennis subgroup that is known from 1.42M–1.51F× wider than long. Elytra 1.38 M–1.49F× longer than wide. Color. Dorsum of head and pronotum metallic green, elytra coppery the Sierra Baoruco and Massif de La Selle. A gap is evi- with very pronounced reddish hue, with or without greenish reflection (Fig. dent between the geographical ranges of this species and its 3c). Appendages markedly infuscate, almost black. closest relative, E. perezi (see description of that species). Microsculpture. Head between eyes with sculpticells somewhat effaced. Head. Frontal furrows with deeply and sharply impressed portion extended to level of anterior supraorbital setigerous punctures (as Fig. 8b). Nitidipennis Group Mouthparts. Labial palpomere 3 securiform in males and broadly to narrowly triangular in females, in males 0.92–1.11 and females Diagnosis.—The nitidipennis group is distinguishable 1.32–1.70× longer than wide. Mentum with paramedian pits large. Prothorax. Pronotum with lateral margins oblique or very slightly from the opaca group by the more pronounced contours sinuate posteriorly, moderately convergent (Fig. 3c); hind angles obtuse; of the submentum and transgular groove, and the shoul- basolateral impressions short and narrowly and deeply impressed, to ders of the elytra which are prominent and have a more nearly punctiform; shallow to deep transverse impression between baso- or less sharp or dentiform humeral angle. The frontal fur- lateral impressions; midlateral setal puncture separated by one-half its diameter from lateral marginal bead. Prosternal intercoxal process with rows on the head are deeply and sharply impressed for apical third weakly to distinctly margined all around or only laterally. nearly their entire length, and typically, the deeply and Pterothorax. Elytral striae 1-7 very shallowly impressed on disc, not sharply impressed portion extends posteriorly at least to evident without magnification. the level of the anterior supraorbital setigerous punc- Male. Hind tibia with basal half of medial ridge represented by row tures, but is shorter in E. yucuyacua, as in most members of prominent subangulate tubercles. Aedeagus with extreme asymmetric type 1 median lobe; preapex moderately long (Figs. 24a, 24d), right third of the opaca group. Another typical feature is that the of preapex considerably thicker and more darkly sclerotized than rest of internal sac of the median lobe of the aedeagus has one preapex and terminated in a prominent lobe (Figs. 24c, 24e), much of narrow apical, and in most species also a narrow basal middle third of preapex very thin compared to rest of preapex and nearly band of large microtrichia extended at least partially colorless (Figs. 24c, 24e), preapical dorsolateral ridge distinctly developed and flat in profile (Figs. 24a–b, 24d); periostial ventrolateral bulge around the sac, but males of E. nitidipennis have just a moderately prominent in profile (Fig. 24b). Internal sac with bulge on left single, broad, central band of large microtrichia extended side at base; arrangement and shape of large microtrichia typical for around the sac, as do opaca group males. member of cupripennis subgroup (Fig. 24f). Description.—Chaetotaxy. Hind coxae with posterior seta present. Material Examined.—Type series only. Femur of middle and hind legs usually with anteroventral pair of setae both present. Remarks.—The male of E. pedernales from Los Arroyos, Microsculpture. Elytra in most species with sculpticells on disc flat, Pedernales Province, differs from males from Barahona in some species surface of disc beaded, sculpticells convex. Head. Frontal furrows deeply and sharply impressed for nearly entire Province in that the preapical dorsolateral ridge of the length, in most species deeply and sharply impressed portion extended median lobe of the aedeagus is narrower (cf. Figs. 24a, posteriorly at least to level of anterior supraorbital setigerous punctures (as 24d), and more of the middle third of the preapex is very in Fig. 8b), or beyond (Fig. 1b), shorter in E. yucuyacua (as in Fig. 8a). thin and colorless (cf. Figs. 24c, 24e). It remains to be Postocular transverse groove in most species shallow but distinct dorsomedially (Fig. 1b), obsolete or nearly so in a few species. Submentum determined whether the differences seen in the male from slightly (Fig. 9c) to very prominent (Fig. 9d), in most species evenly Los Arroyos are constant because the preapex is broken off sloped to transgular groove (Fig. 10a), posterior face concave if submen- and missing in the only other male at hand from Pedernales tum very prominent (Fig. 10b); transgular groove moderately to very deep Province (Las Abejas). The female from Belot-Le Montcel, posterad submentum (Figs. 9c–d). Haiti, is considerably smaller than the other specimens, but Mouthparts. Mandibles with dorsal grooves long in most individuals (Fig. 1b), short in some individuals of a few species (Fig. 9a). Submentum does not differ from them in any other way. with paramedian pits small in most species, of moderate size in a few species. Collecting Notes.—Some specimens of E. pedernales Prothorax. Pronotum with midlateral setal puncture separated by one- from the Sierra de Baoruco were taken on the south face in half its diameter from lateral marginal bead (Fig. 1b); no transverse canyon heads with riparian woody growth, e.g., Las Abejas impression between basolateral impressions. Pterothorax. Elytra with shoulders prominent, humeral angle more or (J.S. Rawlins, pers. com.). Others were collected in second less sharp, dentiform or not (Fig. 1b); basal punctures present or not. growth forest and cloud forest (Los Arroyos). The known Abdomen. Sternum VII with basal sulcus in most species not foveate altitudinal range for this species is 1125–1680 m. medially (Fig. 14c) or just with single faint fovea, entire sulcus foveate in E. tenancingo (Fig. 14d). Geographical Distribution.—Euchroa pedernales is Male. Aedeagus with preapical part of median lobe long or moderate- known from the Sierra de Baoruco in the extreme south- ly so, broad and of uniform thickness and more or less evenly sclerotized throughout (Fig. 38b), or thicker and more darkly sclerotized to right of western part of the Dominican Republic, and also the midline (Fig. 25c), or only developed to right of midline (Fig. 34c), if Massif de La Selle, the westward extension of the Sierra de preapical dorsolateral ridge present, then in profile flat (Fig. 45e) or con- Baoruco in Haiti (Fig. 23). vex (Figs. 25b, 34b); periostial part of median lobe with left wall at least as extensive as and entirely concealing right wall (Fig. 25b) or right wall Phylogenetic Relationships.—The sister species of more extensive than left wall and visible in left lateral view (Fig. 34b); periostial ventrolateral bulge evident and apical in position (Figs. 25b, E. pedernales is E. perezi. 34b); middle part of median lobe with or without dorsal groove (Figs. 25a,

Fig. 17.—Spermatheca and associated glands of females of subgenus Dyschromus, dorsal aspect. A, E. pedernales n. sp.; B, E. dimidiata Chaudoir. bc, bursa copulatrix; di, diverticulum; sgd, duct of spermathecal gland; sgl, spermathecal gland; sp, spermatheca. Scale bar = 0.4 mm.

25d). Internal sac in all but one species with large microtrichia arranged in Taxonomic Composition.—The nitidipennis group com- two narrow transverse bands separated by a central band of much smaller prises all 28 species of the subgenus Dyschromus in microtrichia (Fig. 41a), bands not always complete, basal or apical band in some species not evident, E. nitidipennis with large microtrichia Mexico. The species are distributed among three subgroups. arranged in just one broad central band (as in Fig. 24f). Female. Spermatheca long and recurved (Fig. 17b). Nitidipennis Subgroup Geographical Distribution.—The nitidipennis group is known only from, and apparently confined to, central Diagnosis.—The nitidipennis subgroup is characterized by Mexico. Members of this group occur at least locally the elytral striae, which are uninterrupted and comparative- throughout the southern part of the Sierra Madre Oriental, ly deeply impressed for their entire length. Nine of the 11 the western, central and eastern parts of the Trans-Volcanic species in this subgroup are further distinguished by the Sierra, and most upland areas of Oaxaca and Guerrero (Fig. combination of a metallic bluish-purple or coppery-colored 54). head and pronotum, and reddish-purple elytra. Euchroa harrisoni is uniformly metallic bluish-purple in color dorsally, Phylogenetic Relationships.—The opaca group is the sis- as are all members of the soladevega subgroup, while E. ter group of the nitidipennis group. flohri has a metallic green head and pronotum and brassy- colored elytra, a color combination also found in several Chorological Affinities.—The geographical ranges of the species belonging to the dimidiata subgroup. Geographical nitidipennis and opaca groups are disjunct, since the former distribution, as well as character differences given in the is confined to central Mexico, and the latter to the island of key, serve to distinguish these two species from similarly Hispaniola (Fig. 4). colored members of the other two subgroups.

Description.—Color. Dorsum of head and pronotum metallic bluish- purple or coppery, elytra reddish-purple (Figs. 2c–f); or dorsum uniformly metallic bluish-purple; or head and pronotum metallic green, and elytra brassy (Frontispiece). Microsculpture. Elytra with most sculpticells on disc isodiametric, some slightly transverse, in most specimens sculpticells on disc and intervals 8 and 9 flat, in some individuals of E. perote and E. harrisoni portions of disc and interval 9, as well as apex of elytra slightly beaded, sculpticells slightly convex. Head. Postocular transverse groove shallow but distinct dorsomedially. Mouthparts. Mentum with paramedian pits small. Prothorax. Pronotum with sides slightly to moderately sinuate posteriorly; basolateral impressions short, moderately long or long. Pterothorax. Elytra with basal punctures present; striae relatively deeply impressed, and entire from near base to apex of elytra; intervals nearly flat to moderately convex. Abdomen. Sterna V–VII with basal sulcus foveate only laterally, foveae coarse in most individuals. Male. Middle tibia with inner face distinctly expanded near apex, except indistinctly so in E. harrisoni. Median lobe of aedeagus with preapex at least moderately long, area to left of midline somewhat to considerably thinner and less darkly sclerotized than area to right (Figs. 25c, 25f, 29c, 30c), or preapex only developed to right of midline (Fig. 34c), preapical dorsolateral ridge distinctly developed and convex in profile (Figs. 25b, 29b, 34b); periostial ventrolateral bulge at least moderately prominent in profile (Figs. 25b, 29b, 34b), rounded in cross-section (Figs. 26a–b). Internal sac without constrictions.

Geographical Distribution.—The nitidipennis subgroup is known only from eastern Mexico, north of the Rio Santo Domingo (Figs. 27a–b, 31a–b). Fig. 18.—Median lobe of aedeagus of E. (Dyschromus) tiburonica (Darlington), dorsal aspect. phc, primary haemolymph channels. Scale Phylogenetic Relationships.—We consider the nitidipen- bar = 0.8 mm. nis subgroup to be the sister group of the other two subgroups of Dyschromus in Mexico. See phylogenetic analy- J. Putzeys” [purple label] (IRSNB). Chaudoir, 1874:19; Bates, sis below for details. 1882:85; Tschitschérine, 1898:60.

Chorological Affinities.—Neither of the other two sub- Type Locality.—Putzeys noted in the original description groups in Mexico are known to occur within the geograph- of E. nitidipennis (1844:403) that the holotype is from the ical range of the nitidipennis subgroup, although there is a “Cordilière orientale d'Oaxaca,” Mexico. In the absence of specimen of E. dimidiata (dimidiata subgroup) with ques- additional specimens from that region, the state of Oaxaca tionable locality data indicating that it is from the eastern is considered the type area. part of the Trans-Volcanic Sierra in Veracruz (see description of that species for details). Diagnosis.—Among species of the subgenus Dyschromus with a relatively bright metallic bluish-purple head and Taxonomic Composition.—Eleven species are included pronotum, and reddish-purple elytra, E. nitidipennis is dis- in this subgroup: E. nitidipennis Putzeys; E. sallei tinguished by the comparatively shallow transgular groove, Chaudoir; E. huautla n. sp; E. cuiyachapa n. sp.; E. zon- securiform male labial palpomere 3, the rather broad golica n. sp.; E. lasvigas n. sp.; E. flohri Bates; E. teotitlan pronotum and elytra, and the short pronotal basolateral n. sp.; E. citlaltepetl n. sp.; E. perote n. sp.; and E. harrisoni impressions. The form of the median lobe of the aedeagus n. sp. A few species that we have placed in the dimidiata is distinctive with the preapical part widest near the apex subgroup may also belong to this subgroup. For details, see rather than at the base. phylogenetic analyses, below. Description.—Body Size. Individuals relatively small to moderate in size; apparent body length 9.5–11.5 mm. Length (n=11M,10F): head 0.90 8. Euchroa (Dyschromus) nitidipennis Putzeys (0.79F–1.03F), pronotum in males 2.45 (2.24–2.66) and females 2.49 (2.18–2.82), elytra in males 5.60 (4.90–6.00) and females 5.84 (5.3–6.2) Figs. 5a–b, 9a, 9c, 12b, 16d, 25a–c, 26a, 27a mm; width: head in males 2.07 (1.91–2.24) and females 2.09 (1.91–2.32), pronotum in males 3.24 (3.02–3.57) and females 3.48 (2.96–3.97), elytra Euchroa nitidipennis Putzeys 1844:403. Type material. HOLOTYPE in males 3.79 (3.35–4.10) and females 4.02 (3.7–4.5) mm. female, labeled: “Euchroa nitidipennis/ Mex. Putz.” [white label]; Body Proportions. Head 2.30 (2.17M–2.43F)× wider than long. “E. nitidipennis Ptz/ Mex. f” [green label]; “Soc. Ent. belg/ Coll. Pronotum in males 1.35 (1.32–1.41) and females 1.41 (1.35–1.45)× wider Putzeys” [white label]; “Coll. R. I. Sc. N. B./ Mexique/ ex. coll. than long. Elytra 1.46 (1.41M–1.53)× longer than wide.

Fig. 19.—Median lobe of aedeagus of E. (Dyschromus) opaca (Chaudoir). A and C, dorsal aspect; B, left lateral aspect; D, right lateral aspect, internal sac everted. bb, basal bulb; blm, basal band of large microtrichia; gp, gonopore; is, internal sac; lwp, left wall of periostial part of median lobe; m, middle part of median lobe; o, ostium; p, preapex; pdr, preapical dorsolateral ridge; phc, primary haemolymph channels; pr, periostial part of median lobe; pvb, periostial ventrolateral bulge; rsp, right side of preapex; rwp, right wall of periostial part of median lobe. Scale bar = 0.8 mm.

Color. Dorsum of head and pronotum bright metallic bluish-purple, Mouthparts. Dorsal mandibular grooves short (Fig. 9a). Labial elytra bright reddish-purple (as in Fig. 2e). palpomere 3 in males securiform and females narrowly triangular, in Microsculpture. Elytra with sculpticells on disc finely impressed, flat males 0.93 (0.81–1.13) and females 1.79 (1.68–2.0)× longer than wide. (Figs. 5a–b). Prothorax. Pronotum with sides slightly to moderately sinuate poste- Head. Submentum slightly prominent, posterior face sloped evenly to riorly; hind angles obtuse to slightly prominent; basolateral impressions transgular groove; latter moderately deep posterad submentum (Fig. 9c). short (Fig. 12b). Prosternal intercoxal process with entire apical third in

Fig. 20.—Median lobe of aedeagus of E. (Dyschromus) centralis (Darlington). A and C, dorsal aspect; B, left lateral aspect. Scale bar = 0.8 mm.

most specimens not margined or indistinctly margined, distinctly margin- patch, tree (Liquidambar) covered area, in a maize plantation, pasture ed in 4 out of 58 specimens examined for this trait. edge, September, October, November 1999, February, March, April, May, Pterothorax. Elytral intervals somewhat convex. Mesepisternum with July, August, September, October, 2000, E. Montes de Oca & Q. Santiago, few to many small punctures or impunctate. 72 M & F (IEMM); Rancho Viejo, ca. 10 km. s.w. Xalapa, 1,430 m., mesic Abdomen. Sterna V–VII with foveae in basal sulcus coarse. forest, broad valley, Liquidambar, creek margin and leaf litter, Male. Middle tibia with inner face distinctly expanded near apex. 02.IX.2000, E. Montes de Oca, J.R. Spence, G.E. Ball, 1M (IEMM); Las Hind tibia with basal portion of medial ridge represented by few slightly Vigas, Mexico, Hoege, B.C.A. Col. I. 1., Euchroa flohri Bates, 1M, 1F developed subangulate tubercles (Fig. 16d). Aedeagus with an intermedi- (BMNH), 1M (UASM), 1F (MNHB); Coscoantla, Municipo ate asymmetric type 2 median lobe of usual proportions (Figs. 25a–b); Coscomatepec, 1,250m., 26-IX-94, Liquidamber/ Quercus, Purrington & preapex moderately long, widest near apex (Fig. 25a), entire area to left of Drake, pasture, 1F (FFPC). LOCALITY UNKNOWN: Mexico, 1M, 2F midline somewhat thinner and somewhat less darkly sclerotized than area (MNHB). to right (Fig. 25c), two large primary haemolymph channels present, longest with origin at extreme right side of ostium, broadest arising to left Remarks.—The female holotype is one of the largest spec- of midline (Fig. 25c); left wall of periostial part of median lobe more imens on hand of those that we are attributing to this extensive than, and entirely concealing right wall in lateral view, periostial ventrolateral bulge moderately prominent in profile (Fig. 25b); middle species (apparent body length 11.5 mm), and it has a wider part of median lobe without dorsomedial groove (Fig. 25a). Internal sac pronotum (3.85 mm) than most of the other specimens, and with large microtrichia arranged in one very broad band extended around a narrower head (HW/ ML:1.15) than the others (HW/ ML: sac (as in Fig. 24f). 1.20 [1.16F–1.30M]). The comparatively narrow head com- Material Examined.—Holotype, and 91 additional specimens. VER- bined with the broad pronotum (PWm/ PL:1.44), gives the ACRUZ: km. 2.5, Carr[aterra]. Vieja a Coatepec, P[ar]que Ecol[ogia] holotype a distinctive appearance. Also, it is just one of a Clavigero, 23.VII.[19]89, L. Arellano, Bosque Mesofilo abierto—mucha few individuals with the entire apical third of the prosternal hojarazca trampa copro humano, 1F (IEMM); 2.5 km. s. Jalapa, 26 May intercoxal process distinctly margined. Based upon the M 1991, 1,370 m., J.S. Ashe, #9, ex. sifted leaf litter, 1 (SEMC); Xalapa, original description, the holotype presumably was collect- Inst[ituto]. Ecologia, 1,450 m., mesophytic forest, litter or bromeliads, 10.VII.1999, 99-12, E. Montes de Oca, J.R. Spence & G.E. Ball, 2M ed in the Sierra Madre de Oaxaca, and so is from further (UASM); the same except, leaf litter, stones, vegetation, 23.VII.1999, 99- south than the rest of the specimens. Without a male we can 26, 2M, 1F (UASM); the same except, Cofre de Perote, 4.5 km. w.n.w. only surmise that the distinctive features of the female Xico on Tonalaco Rd., acacia pasture, 1,500 m., 13.VII.1999, 99-13, 1M holotype are owing to geographic variation. Local variation (UASM); the same except, 5.7 km. w.n.w. Xico on Tonalaco Rd., liquidambar-pasture-bromeliads, 1,640 m., 13.VII.1999, 99-14, 1M, 1F is apparent, at least for body size, because most individuals (UASM); Xico (towards El Cofre), variously labeled as follows: 1,310, from the vicinity of Xico are decidedly larger than else- 1,330, 1,390, 1,550 m. a[bove] s[ea] l[evel], in pitfall trap, in a cow-dung where in Veracruz, including specimens from the grounds baited trap, king grass plantation, small coffee plantation patch, tree cover of the Ecological Institute outside of Jalapa, which is only

Fig. 21.—Map of the island of Hispaniola, showing geographical distributions of species of the subgenus Dyschromus belonging to the opaca subgroup. a few kilometers away from and at about the same eleva- in the vicinity of Volcan Citlaltepetl, has been found at an tion as the nearest sites around Xico. even lower elevation and in forest of still more tropical aspect than E. nitidipennis. On the Cofre de Perote, Collecting Notes.—Specimens of this species have been E. nitidipennis and E. lasvigas have both been found at collected in mesophytic forest of tropical aspect at eleva- some of the same sites, but E. nitidipennis may not occur as tions of about 1250 to 1650 m, and also in disturbed areas high up on the mountain as E. lasvigas. at these elevations, e.g., “king grass plantation, small coffee plantation patch, tree cover patch, maize plantation, pasture edge.” 9. Euchroa (Dyschromus) sallei Chaudoir Figs. 2c, 5c, 11a–b, 15a, 25d–f, 27a Bionomics.—A teneral male was collected at the Ecological Institute near Jalapa on July 10, 1999. Euchroa sallei Chaudoir 1874:20. Type material. Three specimens, in the Oberthür-Chaudoir collection (MNHP), associated with the following handwritten label that was pinned originally in the bot- Geographical Distribution.—According to the original tom of an insect box: “sallei/ Chaud/ Mexique.” LECTOTYPE description, the type specimen of E. nitidipennis was col- here selected (first specimen in the series) male, and two male lected in the Sierra Madre de Oaxaca. All of the other spec- PARALECTOTYPES, each labeled “Ex Musaeo/ Chaudoir [red imens that we have attributed to this species, and for which print]. Bates, 1882:85. Tschitschérine, 1898:60. we have locality data, are from the eastern part of the Notes about Type Material.—Chaudoir (1874:20) wrote Trans-Volcanic Sierra in Veracruz (Fig. 27a). that this species was represented by three individuals (of Phylogenetic Relationships.—Euchroa nitidipennis occu- two sexes), sent by M. Sallé. We believe that the three spec- pies the basal or second most basal position within the imens noted above were those that Chaudoir had before nitidipennis subgroup, depending upon the placement of him when he described the species, though all are males. E. sallei and its sister species E. huautla. See phylogenetic Selection of the first specimen as lectotype is arbitrary. analyses below for details. Type Locality.—In conjunction with the original descrip- Chorological Affinities.—Euchroa nitidipennis lives at tion of E. sallei, Chaudoir (1874:20) wrote that the speci- lower elevations and in forest of more tropical aspect than mens were also sent by M. Sallé, but probably from a dif- any other species of the subgenus Dyschromus in its range, ferent locality. The comparison was with Chaudoir’s com- except for E. lasvigas and E. sallei. The last-named species, ments for the preceding species, E. nitidipennis Putzeys, of which he had received two specimens from Sallé that had

Fig. 22.—Median lobe of aedeagus of subgenus Dyschromus: B, left lateral aspect, other figures in dorsal aspect; A–C, E. independencia n. sp.; D–E, E. cupripennis Chaudoir; F–G, E. perezi n. sp. m3p, middle third of preapex; pdr, preapical dorsolateral ridge; r3p, right third of preapex. Scale bar = 0.8 mm.

Fig. 23.—Map of the island of Hispaniola, showing geographical distributions of species of the subgenus Dyschromus belonging to the cupripennis subgroup. 1, Barahona localities, 2, Pedernales localities.

been collected in Mexico. In the absence of more detailed ple, elytra very dark reddish-purple (Fig. 2c). information about where in Mexico the type series was Microsculpture. Elytra with sculpticells on disc finely impressed, flat. Head. Submentum slightly prominent, posterior face sloped evenly to found, we restrict the type locality to the State of Veracruz, transgular groove; latter moderately deep posterad submentum (as in Fig. and more precisely the town of Fortin de las Flores, just 9c). outside of which this species has been collected several Mouthparts. Mandibles with dorsal grooves short. Labial palpomere 3 times in the last 30 years. securiform in males, triangular in females, in males 0.85–1.0 and females 1.33–1.81× longer than wide (Figs. 11a–b). Diagnosis.—The dark hue of the head and pronotum, and Prothorax. Pronotum with lateral margins slightly to moderately sinuate posteriorly; hind angles slightly prominent; basolateral impressions especially of the elytra, distinguishes E. sallei from the short to moderately long. Prosternal intercoxal process with entire apical seven other species of the subgenus Dyschromus that have third distinctly margined. a metallic bluish-purple head and pronotum and reddish- Pterothorax. Elytral intervals moderately convex (Fig. 15a). purple elytra, other than some exceptionally dark-colored Mesepisternum with a few shallow punctures. Abdomen. Sterna V–VII with foveae in basal sulcus coarse. specimens of E. perote. It is readily separable from E. per- Male. Middle tibia with inner face distinctly expanded near apex. ote by the more gentle contours of the submentum and Hind tibia with basal portion of medial ridge represented by a few slight- adjoining part of the transgular groove, and the quite con- ly developed subangulate tubercles. Aedeagus with intermediate asym- vex elytral intervals. Labial palpomere 3 of the male is metric type 2 median lobe, of usual proportions (Figs. 25d–e); preapex securiform, a feature that further distinguishes males of long, broadly tapered to apex (Fig. 25d), area to left of midline somewhat thinner and somewhat less darkly sclerotized than area to right, left mar- E. sallei from those of most other species in the nitidipen- gin near apex even thinner and colorless (Fig. 25f), two primary nis subgroup except for E. nitidipennis, and possibly haemolymph channels present, longest with origin at extreme right side of E. huautla (male unknown). The median lobe of the ostium, other primary channel barely evident but with two or three promi- aedeagus most resembles that of E. cuiyachapa, but the nent secondary channels branched from it (Fig. 25f); left wall of periostial part of median lobe more extensive than and entirely concealing right wall preapex is broader. in lateral view, periostial ventrolateral bulge moderately prominent in profile (Fig. 25e); dorsomedial groove extended entire length of middle part Description.—Body Size. Most individuals relatively large. Apparent M F M F M of median lobe (Fig. 25d). Internal sac with apical band of large body length 11.0 –13.0 mm. Length (n=4 , 6 ): head 0.91–1.15 , microtrichia interrupted ventrally, basal band only evident laterally. pronotum 2.42–3.10F, elytra 5.50M–7.20F mm; width: head 2.02F–2.60F, M F M F pronotum 3.26 –4.25 , elytra 4.00 –5.20 mm. Material Examined.—13 specimens. MEXICO. VERACRUZ: Fortin Body Proportions. Head 2.21M–2.35F× wider than long. Pronotum M F F F M de las Flores, 2900', IX.28.[19]65., George E. Ball, D.R. Whitehead, 2 , 1.31 –1.48 × wider than long. Elytra 1.36 –1.44 × longer than wide. 1F (UASM); the same except, V.27–28.1966, 1F (UASM); the same Color. Dorsum of head and pronotum very dark metallic bluish-pur- except, 3,100', IX.29.65, lot B, 1F (UASM); the same except,

Fig. 24.—Median lobe of aedeagus of E. (Dyschromus) pedernales n. sp. A and C–E, dorsal aspect; B, left lateral aspect; F, right lateral aspect, internal sac everted; A–C and F, specimen from Pedernales; D–E, specimen from Barahona. mt, large microtrichia; pdr, preapical dorsolateral ridge; pvb, periostial ventrolateral bulge. Scale bar = 0.8 mm.

Fig. 25.—Median lobe of aedeagus of subgenus Dyschromus: B and E, left lateral aspect, other figures in dorsal aspect; A–C, E. nitidipennis Putzeys; D–F, E. sallei Chaudoir. dmg, dorsomedial groove; pvb, periostial ventrolateral bulge. Scale bar = 0.8 mm.

26–30.VI.63, leaf litter, D.R. Whitehead, 2M, 1F (UASM); Rio Metlac, 3,200', VIII.I.1973, 512DH, A. Newton, 1F (UASM); Cordoba, Salle Coll., B.C.A. I. 1., Euchroa sallei Chaud., 2F (BMNH); Cordoba, Sa Zongola [Sierra Zongolica], 1F (MNHB). LOCALITY UNKNOWN: Mexico, 1F (SLSC); 42014, 1M (MNHB).

Collecting Notes.—Euchroa sallei has been collected in tropical evergreen forest at about 900 to 1000 m elevation. Geographical Distribution.—This species is known from the eastern part of the Trans-Volcanic Sierra southeast of Volcan Citlaltepetl in Veracruz, and also from the Sierra Zongolica, a mountain range in the extreme northern part of the Sierra Madre de Oaxaca (Fig. 27a). Phylogenetic Relationships.—We consider E. sallei and E. huautla to be sister species, based on both structural and chorological features, and that along with E. nitidipennis, they occupy the most basal positions in the nitidipennis subgroup (see section on phylogeny below for details). Chorological Affinities.—Euchroa sallei occurs at lower elevations and in forest of more tropical aspect than any other species of the subgenus Dyschromus in eastern Mexico, except possibly for E. huautla. As presently known, the ranges of these two species do not overlap, although both live in the Sierra Madre de Oaxaca north of the Rio Santo Domingo.

10. Euchroa (Dyschromus) huautla, new species Figs. 2f, 27a

Type Material.—Two specimens, labeled as follows. HOLOTYPE female, “MEX. Oaxaca/ Rio Santiago, 11.7/ km. e. Huautla de/ Jimenez[,] 1150m,/ cloud forest/ July 13, 1975”; “MEX. EXP. 1975/ George E. Ball &/ H.E. Frania/ collectors,” (USNM). PARATYPE, same labels as holotype, 1F (UASM).

Type Locality.—Rio Santiago, 11.7 km east of Huautla de Jiminez, State of Oaxaca, Mexico.

Diagnosis.—Of the species of the subgenus Dyschromus with reddish-purple elytra, E. huautla is the only one with a coppery-colored head and pronotum. It is further distinguished from the others, except for E. sallei, by the markedly convex elytral intervals.

Description of Adult Female.—Body Size. Individuals relatively large. Apparent body length 13.0 mm. Length (n=2): head 1.07–1.13, pronotum 2.86–3.08, elytra 7.00–7.30 mm; width: head 2.44–2.56, pronotum 3.85–3.97, elytra 4.55–4.75 mm. Body Proportions. Head 2.26–2.28× wider than long. Pronotum 1.29–1.35× wider than long. Elytra 1.54× longer than wide. Color. Dorsum of head and pronotum coppery with a slight greenish reflection, elytra bright reddish-purple (Fig. 2f). Microsculpture. Elytral sculpticells on disc finely impressed, flat. Head. Submentum slightly prominent, posterior face sloped evenly to transgular groove; latter moderately deep posterad submentum. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 Fig. 26.—Scanning electron photomicrograph of median lobe of aedea- subtriangular, 2.14–2.19× longer than wide. gus of subgenus Dyschromus, frontal view. A, E. nitidipennis Putzeys. B, Prothorax. Pronotum with sides moderately sinuate posteriorly; E. perote n. sp. C, E. dimidiata Chaudoir. pdr, preapical dorsolateral hind angles slightly prominent; basolateral impressions moderately ridge; pra, preapex; pvb, periostial ventrolateral bulge. Scale bar = 0.30 mm.

Fig. 27.— Maps of central Mexico, showing geographical distributions of some species of the subgenus Dyschromus belonging to the nitidipennis subgroup.

long. Prosternal intercoxal process not margined. “MEX. Veracruz/ 15.3 km. n. Coscoma/ tepec, betw[een]. Ixtapa/ & Pterothorax. Elytral intervals moderately convex. Mesepisternum Cuiyachapa/ 2300–2400m, cloud/ for[est].[,] VII.6–7.1975”; “MEX. EXP. with numerous, rather coarse punctures. 1975/ George E. Ball &/ H.E. Frania/ collectors,” (USNM). PARATYPES, Abdomen. Sterna V–VII with foveae in basal sulcus coarse. same labels as holotype, 4M, 2F (UASM). “Maltrata/ Mexico/ Salle Coll.”; “471” [blue label]; “Euchroa nitidipennis/ Putz. [illegible] Sallé [hand Material Examined.—The type series only. written]/ B.C.A. I. 1. Euchroa nitidipennis Putz.”, 1M (BMNH). “Haiti”/ Soc. Ent. Belg. Coll.[,] Putzeys”, 1M (IRSNB). Collecting Notes.—The type series of E. huautla was collected in mesophytic forest of tropical aspect, at an Type Locality.—East slope of Volcan Citlaltepetl, between the villages of Ixtapa and Cuiyachapa, state of Veracruz, Mexico. elevation of about 1200 m. Diagnosis.—The very bright luster of the body distinguish- Geographical Distribution.—This species is known es E. cuiyachapa from other species of the subgenus only from the Sierra de Mazateca, Oaxaca, the south- Dyschromus that have a metallic bluish-purple head and ernmost mountain range in the northern part of the pronotum and reddish-purple elytra; so does the condition Sierra Madre de Oaxaca (Fig. 27a). of the microsculpture on the elytra, the sculpticells being Phylogenetic Relationships.—We consider the sister more finely impressed than on the pronotum and effaced in species of E. huautla to be E. sallei (see description of spots. Otherwise, this species is similar in appearance to E. that species). If so, then labial palpomere 3 of the male teotitlan, but the apex of the prosternal intercoxal process of E. huautla should be as broad as that of the male of is not or just indistinctly margined. The aedeagus resembles E. sallei, but we note that it is narrower in the two that of E. sallei, except that the shape of the preapical part females of E. huautla than in females of E. sallei. of the median lobe is reminiscent of E. zongolica. Description.—Body Size. Individuals moderate in size. Apparent body Chorological Affinities.—Only E. sallei occurs at as length 10.5M–11.5F mm. Length (n=5M,2F): head 0.93M–0.99, pronotum low elevations as E. huautla, and it also lives in the 2.42M–2.74F, elytra 5.75M–6.50F mm; width: head 2.06F–2.34F, prono- northern part of the Sierra Madre de Oaxaca, but at the tum 3.10M–3.57F, elytra 3.65M–4.30F mm. opposite end of that system of mountains. Euchroa Body Proportions. Head 2.18M–2.45M× wider than long. Pronotum 1.28M–1.36F× wider than long. Elytra 1.51F–1.59M× longer than wide. teotitlan also occurs in the Sierra de Mazateca, but at Color. Dorsum of head and pronotum brilliant metallic bluish- higher elevations than E. huautla. purple, elytra brilliant reddish-purple and with pronounced greenish reflection in teneral specimens. Microsculpture. Elytra with sculpticells on disc very finely impressed, 11. Euchroa (Dyschromus) cuiyachapa, new species flat, effaced in spots. Figs. 27b, 28a–e Head. Submentum moderately prominent, posterior face sloped evenly to transgular groove; latter deep posterad submentum. Mouthparts. Dorsal mandibular grooves short to long. Labial Type Material.—Nine specimens, labeled as follows. HOLOTYPE male,

Fig. 28.— Median lobe of aedeagus of E. (Dyschromus) cuiyachapa n. sp.: B, left lateral aspect, other figures in dorsal aspect. A–C, specimen from type-locality; D–E, another specimen from type-locality. hs, haemolymph sinus. Scale bar = 0.8 mm.

palpomere 3 narrowly triangular to triangular in males and subtriangular E. zongolica, in combination with the relatively narrow in females, in males 1.24–1.82 and females 2.19–2.20× longer than wide. pronotum and elytra (see key), the microsculpture on the Prothorax. Pronotum with sides moderately to markedly sinuate posteriorly; hind angles slightly to moderately prominent; basolateral impres- disc of the elytra, which consists of sculpticells that are uni- sions moderately long to long. Prosternal intercoxal process not margined formly finely impressed, and the margined prosternal inter- or indistinctly margined laterally. coxal process, distinguish it from the others. In regards to Pterothorax. Elytral intervals nearly flat. Mesepisternum with few to external appearance, D. zongolica is most likely to be mis- many shallow punctures. taken for E. cuiyachapa. The median lobe of the aedeagus Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous. Male. Middle tibia with inner face distinctly expanded near apex. most resembles that of E. lasvigas, but the preapex is more Hind tibia with medial ridge represented nearly to apex by a row of promi- symmetrical in outline and is thinner along the left margin. nent subangulate tubercles. Aedeagus with an intermediate asymmetric type 2 median lobe of usual proportions (Figs. 28a–b, 28d); preapex long, Description of Adult Male.—Body Size. Specimen relatively small. broadly tapered to apex (Figs. 28a, 28d), area to left of midline somewhat Apparent body length 9.5 mm. Length: head 0.87, pronotum 2.26, elytra thinner and somewhat less darkly sclerotized than area to right, left mar- 5.56 mm; width: head 1.95, pronotum 2.96, elytra 3.43 mm. gin near apex even thinner and colorless (Figs. 28c, 28e), one primary Body Proportions. Head 2.22× wider than long. Pronotum 1.31× haemolymph channel present and in most specimens arising at extreme wider than long. Elytra 1.62× longer than wide. right side of ostium (Fig. 28c), one specimen with nearly all of preapex Color. Dorsum of head and pronotum bright metallic bluish-purple, occupied by large haemolymph sinus (Fig. 28e); left wall of periostial part elytra bright reddish-purple. of median lobe more extensive than and entirely concealing right wall in Microsculpture. Elytra with sculpticells on disc finely impressed, flat. lateral view, periostial ventrolateral bulge moderately prominent in profile Head. Submentum prominent, posterior face sloped evenly to transgu- (Fig. 28b); middle part of median lobe without dorsomedial groove (Figs. lar groove; latter moderately deep posterad of submentum. 28a, 28d). Internal sac with apical and basal bands of large microtrichia Mouthparts. Dorsal mandibular grooves short. Labial palpomere 3 both extended entirely around sac. broadly triangular, 1.27× longer than wide. Prothorax. Pronotum with sides moderately sinuate posteriorly; hind Material Examined.—The type series. angles square; basolateral impressions long. Prosternal intercoxal process indistinctly margined at sides near apex. Collecting Notes.—We collected E. cuiyachapa on the Pterothorax. Elytra with basal ridge well developed to level of stria 1; intervals somewhat convex. Mesepisternum with a few faint punctures. east slope of Volcan Citlaltepetl at about 2300 m elevation Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous. in a remnant patch of cloud forest. Male. Middle tibia with inner face distinctly expanded near apex. Hind tibia with basal portion of medial ridge represented by a few moder- Bionomics.—All of the specimens found on 6–7 July, ately developed subangulate tubercles. Aedeagus with intermediate asym- 1975, were teneral. metric type 2 median lobe of usual proportions (Figs. 29a–b); preapex long and broadly tapered toward apex (Fig. 29a), most of preapex stout Geographical Distribution.—This species is known only and darkly sclerotized, extreme left margin very thin and colorless; nearly all of preapex occupied internally by large haemolymph sinus (Fig. from the vicinity of Volcan Citlaltepetl in Veracruz (Fig. 29c); left wall of periostial part of median lobe more extensive than, and 27b). entirely concealing, right wall in lateral view, periostial ventrolateral bulge prominent in profile (Fig. 29b); dorsomedial groove extending entire Phylogenetic Relationships.—We consider that Euchroa length of middle part of median lobe (Fig. 29a). Internal sac not studied. cuiyachapa is most closely related to E. zongolica and E. lasvigas, and is the least derived of the three species. For Material Examined.—Only the holotype. details, see phylogenetic analyses, below. Remarks.—Although nearly all of the preapex of the Chorological Affinities.—The available distributional median lobe of the aedeagus of the holotype of E. zongoli- data are scant, but suggest to us that on the east slope of ca is occupied by a large haemolymph sinus (Fig. 29c), Volcan Citlaltepetl, E. cuiyachapa occupies middle eleva- based upon the situation in E. cuiyachapa (see description tions, with E. sallei and E. nitidipennis occurring at lower, of that species), the usual condition probably is that of one and E. citlaltepetl at higher, elevations. narrow primary haemolymph channel with origin at extreme right side of ostium (as in Fig. 28c).

12. Euchroa (Dyschromus) zongolica, new species Collecting Notes.—While the label indicates that the holo- Figs. 27b, 29a–c type was found in a cave, nothing about the specimen sug- gests that this species lives in a manner different from other Type Material.—Holotype male, labeled “Sótano de Milpa,/ Ver., Mex.[,] species of the subgenus Dyschromus. 4 June 1964,/ T. Raines, W. Bell,” (CMNH). Geographical Distribution.—We know of this species Type Locality.—Five km. s.w. of San Andres de Tenejapa, State of Veracruz, Mexico. Designation of the type locality is based upon informa- only from one specimen from the Sierra Zongolica, a tion that Sótano de Milpa is located 5 km southwest of San Andres de mountain range in the very northernmost part of the Sierra Tenejapa in the Municipo de Tenejapa (Reddell 1981). Madre de Oaxaca in Veracruz (Fig. 27b). Diagnosis.—Of the species of the subgenus Dyschromus Phylogenetic Relationships.—Euchroa zongolica is prob- with a comparatively bright metallic bluish-purple head ably most closely related to E. lasvigas, which is known and pronotum, and reddish-purple elytra, the small size of only from the Cofre de Perote. If so (see section below on

Fig. 29.—Median lobe of aedeagus of subgenus Dyschromus: B and E, lateral aspect; other figures in dorsal aspect. A–C, E. zongolica n. sp.; D–F, E. lasvigas n. sp. Scale bar = 0.8 mm.

phylogenetic relationships), then one or the other of these angles slightly prominent; basolateral impressions long (Fig. 12c). two species, or yet another undescribed one, should occur Prosternal intercoxal process with apex indistinctly margined all around. Pterothorax. Elytra with shoulders more or less prominent and on Volcan Citlaltepetl. humeral angle more or less dentiform (n=2), shoulders not prominent and humeral angle more or less rounded (n=3); basal ridge attaining scutel- Chorological Affinities.—Four species of the subgenus lum or stria 1 (n=3), or only well developed to level of stria 3 and attain- Dyschromus, besides E. zongolica, are known from the ing stria 2 or not (n=2); intervals somewhat convex. Mesepisternum northern part of the Sierra Madre de Oaxaca. Reference to impunctate. World Aeronautical Chart CJ-24 indicates that at the site Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous. Male. Middle tibia with inner face distinctly expanded near apex. where the type specimen was collected, the elevation is Hind tibia with basal portion of medial ridge represented by a few mod- about 1300 m. If correct, then this species occurs at higher erately developed subangulate tubercles. Aedeagus with a variant of an elevations than E. sallei and E. huautla, and lower eleva- intermediate asymmetric type 2 median lobe of usual proportions (Figs. tions than E. teotitlan. 29d–e); preapex long and narrowly tapered near apex, but quite broad towards base (Fig. 29d), stout and darkly sclerotized throughout, nearly all of preapex occupied internally by a large haemolymph sinus (Fig. 29f) or one narrow primary haemolymph channel present with origin at 13. Euchroa (Dyschromus) lasvigas, new species extreme right side of ostium (as in Fig. 28c); left wall of periostial part of Figs. 12c, 27b, 29d–f median lobe more extensive than and entirely concealing right wall in lateral view, periostial ventrolateral bulge prominent in profile (Fig. 29e); Type Material.—Eight specimens, labeled as follows. HOLOTYPE dorsomedial groove extended entire length of middle part of median lobe male, “MEXICO Veracruz/ 15.7 km e[.] Las Vigas/ Rte. 140, cloud/ for- (Fig. 29d). Internal sac of median lobe with apical and basal bands of est, ca. 1930 m./ May 8, 1977”; “MEXICAN EXP. 1977/ J.S. Ashe,/ H.E. large microtrichia both extended around sac. Frania,/ D. Shpeley coll.”, (USNM). PARATYPES, “MEX. Veracruz/ Rancho Viejo/ 6 km. SSW Xalapa/ 1420 m. Riparian for[est]./ w[ith] Material Examined.—The type series. numerous vines/ deep leaf litter/ 21.VII.1998”; “J.R. Spence collector,” 1F (UASM). “MEX. Veracruz, Rancho/ Viejo, 6 km ssw Xalapa/ Collecting Notes.—Euchroa lasvigas has been collected in 19°51.38N 96°58.5W/ riparian mesophytic/ forest; 1420 m/ riparian mesophytic forest, at elevations of about 1400 to 09.VII.1999[,] 99–11”; “COFRE DE PEROTE/ EXPEDITION—1999/ E. Montes de Oca,/ J.R. Spence &/ G.E. Ball collectors,” 1F (UASM); the 1900 m. same except”...5.7 km wnw/ Xico on Tonalaco Rd./ liquidambar-pasture- / bromeliads; 1640 m./ 13.VII.1999[,] 99-14,” 2M (UASM). “MEX. Bionomics.—A teneral female was collected at Rancho Veracruz/ Cuatepec, Rancho/ Viejo, Rancho Aguita/ Fria; 1450 m./ Viejo on July 9, 1999. 07.IX.2000[,] 00-20”; “COFRE DE PEROTE/ EXPEDITION-2000/ E. Montes de Oca,/ J.R. Spence, & G.E. Ball/ collectors,” 2M, 1F (UASM). Geographical Distribution.—This species is known only Type Locality.—Fifteen km. east of Las Vigas, state of Veracruz, from the east side of the Cofre de Perote in northern Mexico. Veracruz (Fig. 27b).

Diagnosis.—Euchroa lasvigas is separable from all other Phylogenetic Relationships.—We consider E. lasvigas to species of the subgenus Dyschromus with a metallic be most closely related to E. zongolica (see description of bluish-purple head and pronotum and reddish-purple ely- that species, as well as section below on phylogenetic rela- tra, including those having the colors as clear and bright tionships). as E. lasvigas, by the combination of proportionately narrow pronotum and elytra (see key), and long pronotal Chorological Affinities.—Euchroa nitidipennis and basolateral impressions. Also, in some specimens the E. flohri also occur on the east side of the Cofre de Perote, basal ridge of the elytra is only slightly developed. The but the latter has been found only at higher elevations than median lobe of the aedeagus most resembles that of E. E. lasvigas. Both E. nitidipennis and E. lasvigas occur at zongolica, but the entire preapex is stout and darkly scle- elevations as low as 1450 m in the vicinity of the Cofre de rotized. Perote, and specimens of both species have been found 5.7 km westnorthwest of Xico at 1640 m, but only E. lasvigas Description.—Body Size. Individuals moderate in size. Apparent body has been collected at elevations higher than this. A consid- length 11.0–12.0 mm. Length (n=3M,2F): head 0.91M–1.05M, pronotum erable distance separates the geographical range of E. lasvi- 2.48M–2.84M, elytra 6.05M–6.65F mm; width: head 2.16M–2.40M, prono- gas from that of its sister species, E. zongolica (see descrip- tum 3.14M–3.45M, elytra 3.75M–4.20M mm. Body Proportions. Head 2.22M–2.37M× wider than long. Pronotum tion of that species). 1.22M–1.33F× wider than long. Elytra 1.53F–1.66M× longer than wide. Color. Dorsum of head and pronotum bright metallic bluish-purple, elytra bright reddish-purple. 14. Euchroa (Dyschromus) flohri, Bates Microsculpture. Elytra with sculpticells on disc moderately finely to Frontispiece, Figs. 30a–c, 31a somewhat deeply impressed, flat. Head. Submentum prominent, posterior face sloped evenly to trans- Euchroa flohri Bates 1882:85. Type material. Two specimens, a male gular groove; latter deep posterad of submentum. and female, in the Oberthür-Chaudoir collection (MNHP). LEC- Mouthparts. Dorsal mandibular grooves short. Labial palpomere 3 in TOTYPE male, labeled: “Quecholac/ Mexico”: “Euchroa/ flohri/ males broadly and females narrowly triangular, in males 1.26–1.29 and Bates” [handwritten]; “H.W. Bates/ Biol. Cent. Amer.”. PARA- females 1.96× longer than wide. LECTOTYPE female, labeled similarly to male, but without the Prothorax. Pronotum with sides moderately sinuate posteriorly; hind determination label. Tschitschérine, 1898:60.

Fig. 30.—Median lobe of aedeagus of E. (Dyschromus) flohri Bates: A and C, dorsal aspect; B, left lateral aspect. Scale bar = 0.8 mm.

Notes about Type Material.—Bates (1882:85) recorded in association Body Proportions. Head 2.30 (2.21M–2.36F)× wider than long. with the original description of this species the locality of “Quecholac.” Pronotum in males 1.31 (1.26–1.38) and females 1.34 (1.31–1.41)× Without reason to doubt that the specimens noted above were those seen wider than long. Elytra in males 1.63 (1.56–1.65) and females 1.60 by Bates, we have selected them here as lectotype and paralectotype. (1.47–1.63)× longer than wide. Color. Dorsum of head and pronotum bright metallic green, elytra Type Locality.—The type locality, as determined from the brassy (Frontispiece). Microsculpture. Elytra with sculpticells on disc moderately finely original description (Bates, 1882:85) and specimen labels impressed, flat. is “Quecholac,” State of Puebla, Mexico, 18°57'N, Head. Submentum prominent, posterior face sloped evenly to trans- 97°40'W. Whether the type series is really from the vicini- gular groove; latter deep posterad submentum. ty of Quecholac is doubtful (see notes concerning geo- Mouthparts. Dorsal mandibular grooves long or short. Labial graphical distribution, below). palpomere 3 in most males broadly and in females narrowly triangular, in males 1.62 (1.28–1.81) and females 2.16 (1.76–2.47)× longer than wide. Diagnosis.—Euchroa flohri is unique among members Prothorax. Pronotum with sides slightly to moderately sinuate pos- of the nitidipennis subgroup in having a metallic green teriorly; hind angles slightly prominent to obtuse; basolateral impres- head and pronotum, and brassy-colored elytra. It is most sions long or moderately so. Prosternal intercoxal process not margined likely to be confused with E. zempoaltepetl (dimidiata or indistinctly margined laterally. Pterothorax. Elytral intervals somewhat convex. Mesepisternum subgroup), but the elytral striae are uninterrupted for with many small punctures. their entire length, and the prosternal intercoxal process Abdomen. Sterna V–VII with foveae in basal sulcus coarse. is not or just very shallowly margined. The median lobe Male. Middle tibia with inner face distinctly expanded near apex. of the aedeagus most resembles that of E. teotitlan, but Hind tibia with medial ridge represented nearly to apex by a row of the preapical dorsolateral ridge is more convex, the left prominent subangulate tubercles. Aedeagus with an intermediate asymmetric type 2 median lobe of usual proportions (Figs. 30a–b); preapex periostial wall is not as high, and the periostial ventro- long, narrowly tapered towards apex (Fig. 30a), most of preapex to left lateral bulge is more prominent. of midline much thinner than area to right and colorless (Fig. 30c), one primary haemolymph channel present, with origin at extreme right side Description.—Body Size. Individuals moderate in size. Apparent body of ostium (Fig. 30c); left wall of periostial part of median lobe more length 11.5M–12.5F mm. Length in males (n=9) and females (n=10): head extensive than, and entirely concealing, right wall in lateral view; 1.00 (0.97–1.03) and 1.02 (0.99–1.07), pronotum 2.61(2.48–2.76) and periostial ventrolateral bulge prominent in profile (Fig. 30b); dorsome- 2.68(2.50–2.82), elytra 6.54 (6.2–7.0) and 6.83 (6.5–7.15) mm; width: dial groove extended entire length of middle part of median lobe (Fig. head 2.26 (2.20–2.28) and 2.35 (2.20–2.46), pronotum 3.42 (3.33–3.75) 30a). Internal sac with basal band of large microtrichia only evident lat- and 3.60 (3.30–3.81), elytra 4.04 (3.9–4.5) and 4.33 (4.05–4.75) mm. erally, no discrete apical band.

Fig. 31.—Maps of central Mexico, showing geographical distributions of some species of the subgenus Dyschromus belonging to the nitidipennis subgroup. RB, Rio Blanco; RSD, Rio Santo Domingo; SZ, Sierra Zacualtipan.

Material Examined.—Thirty-four specimens. VERACRUZ: Cofre de the Cofre de Perote at as high elevations as E. flohri, but on Perote, 27 km. WNW Xico, 2,690 m., pine for[est] on ground, the northwest slope. 15.VII.1998, J.R. Spence, E. Montes de Oca, G.E. Ball, 2M, 3F (UASM); the same except, 05.VII.1999, 99-04A, 3M, 9F (UASM); the same except, ssw slope, under stones and logs, 04.VII.1999, 4M, 2F (UASM); the same except, Tonalaco-Xico road, 2,560 m., pine forest, disturbed area, 15. Euchroa (Dyschromus) teotitlan, new species 11.VIII.2000, 00-04, 1M (IEMM); Ixhuacán (towards El Cofre) Figs. 2e, 10a, 31a, 32a–f 19°25´40''N/ 97°08´30''W, 2,560 m, variously labeled “forest patch, in cow dung bait trap, in pitfall trap, April, June, July,” 2000, E. Montes de Type Material.—One hundred specimens, labeled as follows. HOLO- Oca & Q. Santiago 2M, 6F (IEMM); Las Vigas, Hoge, Euchroa flohri TYPE male, “MEX. OAXACA/ 24.5 km. e. Teotitlan/ Puerto de Soledad/ Bates, 1891-64, 1F (BMNH). LOCALITY UNCERTAIN: PUEBLA: 2240 m. cloud forest/ July 15, 1975”; “MEX. EXP. 1975/ George E. Ball Quecholac, Flohr., 859, 1M (MNHB). &/ H.E. Frania/ collectors,” (USNM). PARATYPES, same labels as holotype, 15M, 15F (UASM); the same, except “22.4 km. e. Teotitlan/ Puerto Collecting Notes.—Specimens of E. flohri have been de Soledad/ 2100 m[,] July 16, 1975,” 1M (UASM); the same, except “MEX. Puebla/ 7.6 km. e. Santa Maria/ del Monte V[icente]. G[uerrero]., found in pine forest at elevations ranging from 2550 to wet/ pine-oak forest 2480 m./ VII.9.1975,” 29M, 12F (UASM); the same, 2700 m. except “7.2 km. w. Santa Maria/ del Monte V.G., oak/ -pine forest 2640m./ July 9, 1975,” 2M, 1F (UASM); the same, except “...7.6 km/ e[.] Santa Geographical Distribution.—This species has been col- Maria del/ Monte V.G.; oak-pine/ leaf litter & under/ logs; 2420 m/ July lected in recent times only on the east side of the Cofre de 16, 1992 16-92/ MEXICAN EXP. 1992/ J.S. Ashe,/ HE. Frania,/ D. Shpeley colls” [some of these specimens also with rearing data], 14M, Perote in northern Veracruz (Fig. 31a). The type locality, 6F (UASM); the same, except “...23.1/ km. e[.] Santa Maria del/ Monte Quecholac, Puebla, is some distance away, being consider- V.G.;...”; “...017-92,” 1M, 2F (UASM); the same, except “...11.1/ km e[.] ably inland of, and at about the same latitude as, Volcan Nicolas Bravo/ dry oak-pine; 2580 m/ under stones & logs/ July 17, Citlaltepetl. As no other species of the subgenus 1992[,] 18-92” 1M (UASM). Dyschromus in eastern Mexico has such a wide range, and Type Locality.—24.5 km. east of Teotitlan Puerto de Soledad, state of the area around Quecholac is much drier than on the east Oaxaca, Mexico. side of the Cofre de Perote, it is likely that the type series was collected elsewhere. Diagnosis.—Among species of the subgenus Dyschromus with a comparatively bright metallic bluish-purple head Phylogenetic Relationships.—We consider E. flohri, and pronotum, and reddish-purple elytra, E. teotitlan is dis- along with E. teotitlan, to be most closely related to a group tinguishable by its moderate size, in combination with its comprised of E. citlaltepetl, E. perote, and E. harrisoni, relatively broad pronotum and elytra, the microsculpture on although the exact relationships of E. flohri and E. teotitlan the disc of the elytra which consists of sculpticells that are are unclear (see section below on phylogeny). moderately finely impressed, and the distinctly margined prosternal intercoxal process. The aedeagus most resem- Chorological Affinities.—Euchroa perote also lives on bles that of E. flohri, but among species with an intermedi-

Fig. 32.—Median lobe of aedeagus of E. (Dyschromus) teotitlan n. sp.: B and E, left lateral aspect; other figures in dorsal aspect. A–C, specimen from type-locality; D–F, specimen from Santa Maria del Monte. Scale bar = 0.8 mm.

Fig. 33.—Median lobe of aedeagus of E. (Dyschromus) citlaltepetl n. sp.: A and C, dorsal aspect; B, left lateral aspect. Scale bar = 0.8 mm.

ate asymmetric type 2 median lobe, the preapical dorsolat- tapered toward apex (Figs. 32a, 32d), area to left of midline somewhat eral ridge is unusually flat in profile, and the left wall of the thinner and somewhat less darkly sclerotized than area to right, nearly all of left margin even thinner and colorless (Figs. 32c, 32f), one primary periostial part of the median lobe is unusually high. haemolymph channel present, with origin at extreme right side of ostium (Figs. 32c, 32f), preapical dorsolateral ridge rather flat in profile (Figs. Description.—Body Size. Individuals moderate in size. Apparent body M F 32b, 32e); left wall of periostial part of median lobe more extensive than, length 10.5 –12.5 mm. Length in males (n=10) and females (n=10): and entirely concealing, right wall in lateral view, periostial ventrolateral head 0.94 (0.89–0.99) and 0.99 (0.95–1.05), pronotum 2.63 (2.44–2.74) bulge moderately prominent in profile (Figs. 32b, 32e); dorsomedial and 2.80 (2.62–2.96), elytra 6.38 (6.05–6.70) and 6.78 (6.5–7.1) mm; groove extended entire length of middle part of median lobe (Figs. 32a, width: head 2.22 (2.06–2.34) and 2.38 (2.20–2.56), pronotum 3.48 32d). Internal sac with apical and basal bands of large microtrichia both (3.24–3.67) and 3.71 (3.39–4.03), elytra 4.07 (3.8–4.25) and 4.49 extended entirely around sac. (4.3–4.7) mm. Body Proportions. Head 2.38 (2.26–2.50)× wider than long. Pronotum Material Examined.—The type series, and one additional female speci- 1.32 (1.28–1.38)× wider than long. Elytra in males 1.57 (1.52–1.61) and men not included in the type series, labeled “Orizaba”; “Mexico./ Salle females 1.51 (1.46–1.55)× longer than wide. Coll.”; “471”; “Euchroa/ nitidipennis/ Putz ?[illegible] Sallé”; “B.C.A. I. Color. Dorsum of head and pronotum bright metallic bluish-purple, 1. Euchroa nitidipennis Putz.”; “E. (Dyschromus)/ teotitlan Frania and elytra bright reddish-purple and usually with green reflection (Fig. 2e). Ball?/ det. Frania & Ball” (BMNH). Microsculpture. Elytra with sculpticells on disc moderately finely impressed, flat. Head. Submentum prominent, posterior face sloped evenly to transgu- Remarks.—The sculpticells on the disc of the elytra of the lar groove; latter deep posterad submentum. female labeled “Orizaba...” are more finely impressed and Mouthparts. Dorsal mandibular grooves short. Labial palpomere 3 in the surface of the elytra has a brighter luster than in any males broadly and in females narrowly triangular, in males 1.33 specimen of E. teotitlan that we have seen from the north- (1.28–1.39) and females 1.92 (1.59–2.21)× longer than wide. Prothorax. Pronotum with sides slightly to moderately sinuate poste- ern part of the Sierra Madre de Oaxaca in the states of riorly; hind angles obtuse to slightly prominent; basolateral impressions Puebla and Oaxaca. The colors of this individual are in fact long or moderately so. Prosternal intercoxal process with entire apical about as brilliant as in E. cuiyachapa, which is known only third distinctly margined. from the vicinity of Mt. Orizaba (Volcan Citlaltepetl) in the Pterothorax. Elytral intervals somewhat convex. Mesepisternum with state of Veracruz. Otherwise, the specimen in question does many small punctures or impunctate. Abdomen. Sterna V–VII with foveae in basal sulcus coarse. not differ in any obvious feature from females of E. teotit- Male. Middle tibia with inner face distinctly expanded near apex. lan found in Puebla and Oaxaca, and so we consider it to be Hind tibia with medial ridge represented nearly to apex by a row of promi- conspecific with them. However, in the absence of a male nent subangulate tubercles. Aedeagus with an intermediate asymmetric from Volcan Citlaltepetl, we do not include it in the type type 2 median lobe of usual proportions (Figs. 32a, 32d), except left wall of periostial part unusually high (Figs. 32b, 32e); preapex long, narrowly series.

Collecting Notes.—We have collected E. teotitlan in oak- Pterothorax. Elytral intervals somewhat convex. Mesepisternum with pine forest and cloud forest at elevations of about 2100 to some small to somewhat coarse punctures. Abdomen. Sterna V–VII with foveae in basal sulcus coarse. 2600 m. Male. Middle tibia with inner face distinctly expanded near apex. Hind tibia with medial ridge represented nearly to apex by a row of Geographical Distribution.—This species occurs in the prominent subangulate tubercles. Aedeagus with intermediate asym- Sierra Madre de Oaxaca, north of the Rio de Santo metric type 2 median lobe of usual proportions (Figs. 33a–b); preapex Domingo, in the states of Puebla and Oaxaca (Fig. 31a), long, narrowly tapered towards apex (Fig. 33a), area to left of midline and it may also occur on Volcan Citlaltepetl in Veracruz. of median lobe reduced in extent compared to rest of preapex and much thinner and colorless (Fig. 33c), one primary haemolymph channel present with origin at extreme right side of ostium (Fig. 33c); right Phylogenetic Relationships.—The closest relative of E. wall of periostial part of median lobe more extensive than left wall, teotitlan could be E. flohri, but whether they are sister ostium visible in left lateral view (Fig. 33b), periostial ventrolateral species is unclear. For details, see description of E. flohri, bulge very prominent in profile (Fig. 33b); dorsomedial groove and also phylogenetic analyses, below. extended entire length of middle part of median lobe (Fig. 33a). Internal sac not studied.

Chorological Affinities.—The data are scanty, but suggest Material Examined.—The type series only. to us that E. teotitlan occurs further inland and in forest of more temperate aspect than any other species of the sub- Collecting Notes.—Euchroa citlaltepetl has been col- genus Dyschromus that lives in the northern part of the lected at an elevation of about 2900 m, probably in pine Sierra Madre de Oaxaca. forest. Geographical Distribution.—This species is known only 16. Euchroa (Dyschromus) citlaltepetl, new species from Volcan Citlatepetl in Veracruz (Fig. 31b). Figs. 10b, 31b, 33a–c Phylogenetic Relationships.—Evidence is reasonably Type Material.—Two specimens, labeled as follows. HOLOTYPE male, “E. Citlaltepetl/ Ver. Mexico/ VII-10-64[,] el. 9500 [ft.]/ L. W. Swan,” convincing that E. perote and E. harrisoni, together com- (CASC). PARATYPE, “Orizaba”; “Mexico/ Salle Coll./ “471”/ B.C.A. I. prise the sister group of E. citlaltepetl (see section below on 1./ Euchroa/ nitidipennis Putz.”; “Euchroa/ nitidipennis/ Putz. ?[illegible] phylogeny for details). Sallé [hand written]”, 1 M (BMNH). Chorological Affinities.—No other species of the sub- Type Locality.—Volcan Citlaltepetl, state of Veracruz, Mexico. The holotype was presumably collected on the east slope of the mountain. genus Dyschromus on Volcan Citlalteptel has been found at as high an elevation as E. citlaltepetl. Its closest relatives Diagnosis.—This is one of the species of the subgenus occur at comparable elevations further north in the moun- Dyschromus with a metallic bluish-purple head and prono- tains of eastern Mexico (Fig. 31b). tum and reddish-purple elytra in which the colors are rather dark in hue. As such, E. citlaltepetl is likely to be mistaken 17. Euchroa (Dyschromus) perote, new species for a small-sized E. perote, but the pronotum is narrower Figs. 2d, 26b, 31b, 34a–c relative to the width of the head (see key). It might also be mistaken for an unusually dark-colored specimen of E. Type Material.—One hundred forty-five specimens, labeled as follows. teotitlan, but the submentum is more prominent. The Holotype male, “MEX. Veracruz/ Cofre de Perote/ n[orth]. slope, 10 mi. aedeagus most resembles that of E. perote, but differs in the s./ Las Vigas, 9600'/ VIII.24.1967”; “Ball, T.L. Erwin/ R.E. Leech/ collectors” (USNM). PARATYPES, same labels as holotype, 14M, 12F greater development of the left side of the preapical part of (UASM). “PEROTE/ Veracruz (México)/ 29[.]VIII[.]1973/ J. Mateu the median lobe. coll.,” 1M (SLSC). “JALAPA/ MEXICO”; “Van Dyke/ Collection,” 1M (CASC). “Jalapa,/ Mexico./ Hoege,” 1M (BMNH), 1M (MNHB), 1M, Description of Adult Male.—Body Size. Individuals of moderate size. (IRSNB). “Las Vigas/ Vera Cruz./ Hoege,” 3F (BMNH), 1F (MCZC), 1M, Apparent body length 11.5–12.0 mm. Length (n=2M): head 0.95–0.99, 3F (MNHB). “MEXICO Veracruz 3.2/ km sw Las Vigas; Rte/ 140; dry pronotum 2.66–2.70, elytra 6.40–6.80 mm; width: head 2.26–2.28, prono- pine-oak/ 2380 m; under/ stone/ July 11, 1992[,] 08-92; “MEXICAN EXP. tum 3.49–3.55, elytra 4.25–4.35 mm. 1992/ J.S. Ashe/ H.E. Frania/ D. Shpeley colls.,” 1M (UASM). “Las Minas Body Proportions. Head 2.28–2.40× wider than long. Pronotum a Las/ Vigas–Veracruz/ (México) J. Mateu,” 1F (SLSC). “MEX Puebla 1.31–1.32× wider than long. Elytra 1.51–1.56× longer than wide. 50.8/ km. s.e. Azumbilla/ 2480 m. oak-pine/ forest, logs/ ground 78B-36a/ Color. Dorsum of head and pronotum metallic bluish-purple, elytra Dec. 24–25, 1978”; “MEXICAN EXP. 1978–1979/ G.E. & K.E. Ball/ col- reddish-purple with slight green reflection, colors rather dark in hue (as in lectors,” 1M, 1F (UASM). “MEX. Puebla/ Honey 2160 m/ oak-pine for- Fig. 2d). est/ June 27, 1975”; “MEX. EXP. 1975/ G.E. Ball &/ H.E. Frania/ collec- Microsculpture. Elytra with sculpticells on disc moderately finely tors,” 12M, 15F (UASM); the same, except “Microondas, 32.7/ km. n. impressed, flat. Zacatlan/ Rte. 119, 2460m/ oak-pine forest/ June 28, 1975,” 2M, 1F Head. Submentum very prominent, posterior face somewhat concave, (UASM); the same, except “km. 11, Rte. 119 to/ Tetela de Ocampo 2740 transgular groove very deep posterad submentum (Fig. 10b). m June 29, 1975,” 5M, 8F (UASM). “6 mi. w. Tezuitlan/ Pueblá, Mex./ Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 Aug. 16 1958/ H.F. Howden,” 1M (UASM), 1M, 2F (CNCI). “MEXICO broadly triangular, 1.33–1.45× longer than wide. Puebla 9mi./ SW of Coacoyunga/ 14 Aug. 1969 9700'/ U.Kansas Mex. Prothorax. Pronotum with sides slightly sinuate posteriorly; hind Exped.,” 2F (SEMC). “MEXICO. HIDALGO/ 10 mi. N. Agua/ Blanca angles slightly prominent; basolateral impressions long. Prosternal inter- [northwest of Honey], 2600 m. 18/ Aug '52. G. Rabb[,] 3”, 3M, 2F coxal process with entire apical third distinctly margined.

Fig. 34.—Median lobe of aedeagus of E. (Dyschromus) perote n. sp.: A and C, dorsal aspect; B, left lateral aspect. pdr, preapical dorsolateral ridge; phc, primary haemolymph channel; pvb, periostial ventrolateral bulge; rwp, right wall of periostial part of median lobe. Scale bar = 0.8 mm.

(MSUC). “Guerrero Mills/ Hidalgo, Mex./ W.M. Mann,” 2M, 2F (MCZC). pronotum 3.93 (3.53–4.13) and 4.23 (3.65–4.64), elytra 4.71 (4.25–5.00) “MEX. Tlaxcala/ 6.8 km. n. Tlaxco [Rte. 119]/ 2820 m[,] pine-fir/ forest/ and 5.14 (4.55–5.65) mm. June 30, 1975”; “MEX. EXP. 1975/ G.E. Ball &/ H.E. Frania/ collectors,” Body Proportions. Head 2.35(2.26–2.50)× wider than long. Pronotum 7M, 6F (UASM); the same, except “...6.8/ km n Tlaxco; Rte 119/ pine- in males 1.33(1.28–1.37) and females 1.35(1.31–1.41)× wider than long. alder-manzanita/ for[est]. edge; 2820 m./ Aug. 22, 1992[,] 72-92”; “MEX- Elytra in males 1.52(1.49–1.55) and females 1.50(1.45–1.53)× longer than ICAN EXP. 1992/ J.S. Ashe,/ H.E. Frania,/ D. Shpeley colls.” [some spec- wide. imens also with rearing data], 2M, 3F (UASM); the same, except “pine-fir; Color. Dorsum of head and pronotum dark metallic bluish-purple, ely- 2775 m/ under stones/ July 9, 1992, 06-92,” 17M, 7F (UASM). “Durango/ tra of most specimens dark reddish-purple (Fig. 2d), few specimens with Mexico,” 1M (SLSC). “LA PLATA,” 1F (SLSC). a greenish reflection, color somewhat brighter in specimens with relatively finely impressed microsculpture on disc of elytra. Type Locality.—About 16 km south of Las Vigas, State of Veracruz, Microsculpture. Elytra of most individuals with surface of disc slight- Mexico. ly beaded, sculpticells slightly convex and moderately deeply impressed, some specimens with sculpticells on disc moderately finely impressed and Diagnosis.—Euchroa perote is not as brightly colored as flat; most individuals with sculpticells on intervals 8 and 9 flat, surface most other species of the subgenus Dyschromus that have beaded and sculpticells convex in individuals from Tlaxcala. Head. Submentum very prominent, posterior face concave; transgular a metallic bluish-purple head and pronotum, and reddish- groove very deep posterad submentum (as in Fig. 9d). purple elytra, except for E. citlaltepetl, and also E. sallei Mouthparts. Mandibular grooves short in most specimens, long in which has an even darker appearance. The very promi- some specimens. Labial palpomere 3 in males broadly and females nar- nent submentum, which is concave posteriorly, further rowly triangular, in males 1.28 (1.18–1.49) and females 1.72 (1.55–1.79)× longer than wide. separates E. perote from the others, again except for Prothorax. Pronotum with sides slightly to moderately sinuate poste- E. citlaltepetl. Nearly all individuals of E. perote are riorly; hind angles in most specimens acute and slightly prominent, obtuse larger in size than E. citlaltepetl, and the pronotum is in a few specimens; basolateral impressions long or moderately so. slightly broader. The median lobe of the aedeagus close- Prosternal intercoxal process with entire apical third deeply margined. ly resembles that of E. harrisoni. Pterothorax. Elytral intervals somewhat convex. Mesepisternum with few to many small punctures or impunctate (some specimens from Description.—Body Size. Most individuals relatively large. Apparent Honey). body length 11.5M–14.0F mm. Length in males (n=10) and females Abdomen. Sterna V–VII with foveae in basal sulcus coarse. (n=10): head 1.06 (0.95–1.11) and 1.09 (0.91–1.19), pronotum 2.97 Male. Middle tibia with inner face distinctly expanded near apex. (2.70–3.10) and 3.11 (2.78–3.35), elytra 7.17 (6.50–7.50) and 7.66 Hind tibia with basal portion of medial ridge represented by few moder- (6.80–8.30) mm; width: head 2.47 (2.24–2.60) and 2.59 (2.20–2.76), ately developed subangulate tubercles. Aedeagus with extreme asymmetric type 2 median lobe of usual proportions (Figs. 34a–b); preapex long,

narrowly tapered to apex, and developed only to right of midline of median lobe (Fig. 34a), stout and darkly sclerotized throughout (Fig. 34c), one primary haemolymph channel present, with origin at extreme left side of ostium (Fig. 34c); right wall of periostial part of median lobe more extensive than left wall, ostium visible in left lateral view (Fig. 34b); periostial ventrolateral bulge very prominent in profile (Fig. 34b); dorsomedial groove extended entire length of middle part of median lobe (Fig. 34a). Internal sac with apical and basal bands of large microtrichia both extended entirely around sac.

Material Collected.—The type series.

Collecting Notes.—Euchroa perote has been collected in comparatively dry oak-pine, pine, and pine-fir forest at elevations of about 2100 to 3000 m. Geographical Distribution.—This species is known from the inland side of the Cofre de Perote and the mountains to the west and northwest, in Veracruz, Puebla, Hidalgo, and Tlaxcala (Fig. 31b). Phylogenetic Relationships.—Euchroa harrisoni is the sister species of E. perote (see phylogenetic analyses below for details). Chorological Affinities.—Euchroa perote occurs at higher elevations and in forest of more temperate aspect than any other species of Dyschromus in its range, except for Fig. 35.— Median lobe of aedeagus of E. (Dyschromus) harrisoni n. sp.: E. flohri. Both species occur on the Cofre de Perote, but dorsal aspect. Scale bar = 0.8 mm. E. perote has only been found on the north and west slopes, E. flohri on the east slope. also with rearing data], 2M, 8F (UASM). “MEX. Hidalgo/ Rte. 105[,] The most northerly locations for E. perote (Guerrero 3.2 km. n./ Tlanchinol[,] 1540 m/ ground cloud forest/ June Mills in Hidalgo, and Honey in Puebla) are between 50 21–23.1975”/ “MEX. EXP. 1975/ G.E. Ball/ H.E. Frania/ collectors,” and 75 km from the most southerly locality for its sister 2M, 3F (UASM). “MEX. Hidalgo/ Rte. 85 23.8 mi./ n.e. Jacala/ 5100' species, E. harrisoni (Tlanchinol in Hidalgo). No topo- XI.13.65”; “George E. Ball/ D.R. Whitehead/ collectors,” 1 F (UASM). “MEX. HGO[.] 5200'/ 10 mi. N.E. Rancho/ Viejo graphic barriers to dispersal are evident in the intervening VI.29.[19]71/ A. Newton[,] 257,” 1M (UASM). “Cueva de Puerto de/ area (World Aeronautical Chart CJ-24), but we do not la Zorra [10 km. n.e. of Jacala (Reddell 1981)], Hgo., Mex./ know whether suitable forest habitat is present throughout 11.VIII.1966[,] J. Fish,/ J. Reddell, leg.,” 1F (CMNH). the area. Type Locality.—About 12.8 km northwest of Gomez Farias, state of Tamaulipas, Mexico. 18. Euchroa (Dyschromus) harrisoni, new species Figs. 11c–d, 31b, 35a–b Specific Epithet.—The specific epithet is an eponym, Latinized, genitive case, based on the surname of William Type Material.—Sixty-nine specimens, labeled as follows. HOLOTYPE Francis (Frank) Harrison, a Canadian teacher, naturalist male, “MEX. Tamaulipas. Sierra de Guat-/ emala, Rancho del/ Cielo, 8 and farmer, who made his home for some two decades on mi. n. w./ Gomez Farias/ 3800' X.6-10.65”; “George E. Ball/ D.R. the slopes of the Sierra de Guatemala, in eastern Whitehead/ collectors,” (USNM). PARATYPES, same labels as holotype, 8M, 10F (UASM); the same, except “July 24–29, 1971”; “George E. and/ Tamaulipas. His small mountain ranch (Rancho del Cielo), Kathleen E. Ball collectors,” 2F (UASM); the same, except “Agua located in the northernmost cloud forest in the Neotropical Linda”/ ca. 12 mi. n.w./ Gomez Farias/ 5800'/ X.9.[19]65”; “George E. Region, served as a base for numerous field biologists Ball and D.R. Whitehead,” 3M, 3F (UASM); the same, except “8.1 mi. w. (see, for example, Sutton 1972: 138–142, and Martin Encino, 3100'/ X.II.65" (elytron, UASM). “5 mi. n. Gomez Farias[,] 3200'[,] Rancho Cielo/ cloud forest Martin.Robbins,” 1M (UASM). 1958) who came to study the rich biota of the Sierra de “MEX. S.L.P./ 24.7 mi. e. Landa/ de Matamoros/ Qro.[,] 5000'/ Guatemala from 1948 to 1965. His life was ended late in XI.18–19.65”; “George E. Ball/ D.R. Whitehead/ collectors,” 8M, 6F 1965 by an unexpected, unfortunate encounter with an (UASM). “MEX. Queretaro/ 17.8 mi. e. Landa/ de Matamoros/ 5300' uneducated, determined campesino (peasant farmer) who, M F XI.17-18./ 65”; “George E. Ball/ D. R. Whitehead/ collectors,” 3 , 2 for his ejido (rural collective), wanted Frank’s land. We (UASM). “MEX. Qro., 29 km e/ Landa de Matamoros/ oak-pine litter/ under logs; 1520 m/ 20.VIII.1985 64–85”; “MEXICAN EXP. 1985/ dedicate this species to the memory of Frank Harrison, a H.E. Frania &/ D. Shpeley/ collectors,” 2M, 1F (UASM); “the same, gentle, friendly man, who was an environmentalist before except “...litter;/ “oak-pine; 1520 m/ 24–25.VII.92[,] 76–92/ MEXI- that term was invented. CAN EXP. 1992/ G.E. Ball and/ H.E. Frania colls.” [some specimens

Fig. 36.—Median lobe of aedeagus of subgenus Dyschromus: B, left lateral aspect, other figures in dorsal aspect. A–C, E. soladevega n. sp.; D–E, E. juchatengo n. sp.; F–G, E. nizavaguiti n. sp. dmg, dorsomedial groove. Scale bar = 0.8 mm.

Diagnosis.—Euchroa harrisoni is the only member of the south in the eastern part of the Trans-Volcanic Sierra (see subgenus Dyschromus in eastern Mexico that is uniformly discussion of that species). metallic bluish-purple in color dorsally. The aedeagus is nearly identical in appearance to that of E. perote. Soladevega Subgroup Description.—Body Size. Most individuals relatively large. Apparent body length 11.0M–14.0F mm. Length in males (n=10) and females Diagnosis.—Species belonging to soladevega subgroup (n=10): head 1.03 (0.91–1.11) and 1.09 (0.99–1.19), pronotum 2.93 are uniformly metallic bluish-purple in color dorsally, and (2.58–3.13) and 3.07 (2.86–3.28), elytra 7.06 (6.30–7.40) and 7.43 are characterized further by the elytral striae, which are (6.80–7.80) mm; width: head 2.40 (2.08–2.56) and 2.56 (2.36–2.74), pronotum 3.80 (3.33–3.97) and 4.09 (3.73–4.53), elytra 4.57 (4.00–4.80) entire and moderately impressed on the disc of the elytra, and 4.99 (4.65–5.45) mm. and by the microsculpture on elytral interval 9 and the outer Body Proportions. Head 2.36 (2.22–2.48)× wider than long. Pronotum half of interval 8, which consists of sculpticells that are in males 1.30 (1.27–1.33) and females 1.33 (1.31–1.38)× wider than long. markedly convex, so that the surface of both intervals is Elytra in males 1.54 (1.50–1.58) and females 1.50 (1.43–1.54)× longer than wide. distinctly beaded. Color. Dorsum clear metallic bluish-purple. Microsculpture. Elytra with sculpticells on disc and at sides moderate- Description.—Color. Dorsum metallic bluish-purple. ly finely impressed and flat, or elytral surface slightly beaded, with sculp- Microsculpture. Elytra with sculpticells on disc isodiametric or elon- ticells somewhat deeply impressed and slightly convex. gate, some species with portions or entire surface of disc beaded and Head. Submentum very prominent, posterior face concave; transgular sculpticells convex; outer half of interval 8, all of 9, and apex of elytra groove very deep posterad submentum. with surface distinctly beaded (Fig. 15b), sculpticells isodiametric and Mouthparts. Dorsal mandibular grooves short or long. Labial markedly convex (Fig. 6c). palpomere 3 in males broadly (Fig. 11d) and females narrowly triangular Head. Postocular transverse groove shallow but distinct dorsomedi- (Fig. 11c), in males 1.12 (1.07–1.33) and females 1.60 (1.49–1.81)× ally. Submentum with posterior face sloped evenly to transgular groove. longer than wide. Mouthparts. Dorsal mandibular grooves long. Mentum with parame- Prothorax. Pronotum with sides slightly to moderately sinuate poste- dian pits small. riorly; hind angles obtuse to slightly prominent; basolateral impressions Prothorax. Pronotal basolateral impressions short. Prosternal inter- moderately long in most individuals, long in a few individuals. Prosternal coxal process with entire apical third deeply margined. intercoxal process with entire apical third deeply margined. Pterothorax. Elytra with basal punctures; striae moderately deeply Pterothorax. Elytral intervals somewhat convex. Mesepisternum impressed, entire on disc. Mesepisternum coarsely punctate. impunctate. Abdomen. Sterna V–VII with basal sulcus only foveate laterally, Abdomen. Sterna V–VII with foveae in basal sulcus coarse. foveae inconspicuous to coarse. Male. Middle tibia slightly expanded medially near apex. Hind Male. Middle tibia with inner face distinctly expanded near apex. tibia with basal portion of medial ridge represented by few slightly Median lobe of aedeagus with preapex thicker to right of midline, or developed subangulate tubercles. Median lobe of aedeagus as in E. uniformly thick throughout; periostial ventrolateral bulge rounded or perote, except somewhat more slender, and dorsomedial groove of subangulate in cross-section. Internal sac without constrictions. middle part of median lobe comparatively shallow (Figs. 35a–b). Internal sac with apical and basal bands of large microtrichia both Geographical Distribution.—The soladevega sub- extended entirely around sac. group is known only from the Sierra de Quatro Venados and Sierra de Miahuatlan in southwestern Oaxaca (Fig. Material Examined.—The type series. 37a). Collecting Notes.—Euchroa harrisoni has been found in Phylogenetic Relationships.—We consider the dimidiata oak-pine forest and cloud forest at elevations of about 900 subgroup to be the sister group of the soladevega subgroup, to 1800 m. but evidence for monophyly of the soladevega subgroup is Bionomics.—A teneral female was collected at Rancho del weak. For details, see phylogenetic analyses, below. Cielo on July 24–29, 1971. Chorological Affinities.—The geographical range of the Geographical Distribution.—This species is found in the soladevega subgroup is encompassed within that of the southern portion of the Sierra Madre Oriental, in the states dimidiata subgroup. of Tamaulipas, San Luis Potosi, Hidalgo, and Queretaro Taxonomic Composition.—Three species are included in (Fig. 31b). The northern limit of the distributional range of this subgroup: E. soladevega n. sp., E. juchatengo n. sp., E. harrisoni may correspond to that of cloud forest in and E. nizavaguiti n. sp. Mexico (Martin 1958).

Phylogenetic Relationships.—The sister species of 19. Euchroa (Dyschromus) soladevega, new species E. harrisoni is E. perote (see section on phylogeny below Figs. 6a–c, 15b, 36a–c, 37a for details). Type Material.—Twenty-two specimens, labeled as follows. HOLO- Chorological Affinities.—No other species of TYPE male, “MEXICO. Oaxaca./ 9.3 mi. n. Sola de Vega 6000'/ Dyschromus is known to live in the Sierra Madre Oriental. VII.18.1966; “George E. Ball/ D.R. Whitehead/ collectors,” (USNM). PARATYPES, same labels as holotype, 3M, 1F (UASM); the same, except Its closest relative, E. perote, lives a short distance to the “7100' 21.8 mi./ n. Juchatengo./ VII.18–19.1966.,” 3M, 1F (UASM); the

Fig. 37.—Maps of central Mexico, showing geographical distributions of some species belonging to the subgenus Dyschromus. A, soladevega subgroup; B, dimidiata subgroup, in part.

same, except “Rte. 131, 13.1/ mi. n. Jucha-/ tengo 4400'/ cypress creek/ females 1.81 (1.59–1.97)× longer than wide. August 7, 1972”; “B.S. Heming/ G.E. Ball COLLECTORS,” 1M (UASM). Prothorax. Pronotum with sides slightly sinuate very near base; hind “MEXICO Oaxaca 14.9/ km[.] n[.] Sola de Vega/ dry oak-pine forest/ leaf angles obtuse. litter; 1820 m/ July 20, 1992[,] 24-92,” “MEXICAN EXP. 1992/ J.S. Pterothorax. Elytral intervals nearly flat (Fig. 15b). Ashe,/ H.E. Frania,/ D. Shpeley colls.” [some specimens also with rearing Male. Hind tibia with medial ridge represented nearly to apex by row data], 4M, 6F, 2? (UASM). of moderately prominent, subrectangular tubercles. Aedeagus with intermediate asymmetric type 2 median lobe of usual proportions (Figs. Type Locality.—About 14.9 km north of Sola de Vega, state of Oaxaca, 36a–b); preapex long and broadly tapered to apex (Fig. 36a), with narrow Mexico. dorsolateral ridge terminated in small apical lobe (Fig. 36a), convex in profile (Fig. 36b), entire preapex to left of midline somewhat thinner and Diagnosis.—Euchroa soladevega is distinguishable from somewhat less darkly sclerotized than area to left (Fig. 36c), one primary E. juchatengo and E. nizavaguiti, the other two species of haemolymph channel present, narrow, with origin at extreme left side of ostium (Fig. 36c); left wall of periostial part of median lobe more exten- the subgenus Dyschromus in southwestern Mexico that are sive than and entirely concealing right wall in lateral view (Fig. 36b); uniformly metallic bluish-purple in color dorsally, by its periostial ventrolateral bulge slightly prominent in profile (Fig. 36b) and larger size, nearly flat elytral intervals, and the microsculp- rounded in cross-section (as in Fig. 26a); dorsomedial groove extended ture on the disc of the elytra, which consists of sculpticells entire length of middle part of median lobe (Figs. 36a). Internal sac with apical and basal bands of large microtrichia both extended entirely around that are as deeply impressed as those along the sides of the sac. elytra. The median lobe of the aedeagus has a general resemblance to that of species belonging to the other two Material Examined.—The type species. subgroups that also have an intermediate asymmetric type 2 median lobe. Collecting Notes.—We have collected E. soladevega in dry oak forest at elevations of about 1300 to 2200 m. Description.—Body Size. Individuals relatively large. Apparent body length 12.0M–13.5F mm. Length (n=10M,7F): head 1.01 (0.91M–1.07M), Geographical Distribution.—This species is known only F M M F pronotum 3.25 (3.04 –3.47 ), elytra 7.02 (6.55 –7.60 ) mm; width: from the southernmost part of the Sierra de Quatro head 2.44 (2.22M–2.68F), pronotum 4.16 (3.81M–4.51F), elytra 4.82 (4.40M–5.30F) mm. Venados, in southwestern Oaxaca (Fig. 37a). Body Proportions. Head 2.42 (2.31M–2.60F)× wider than long. Pronotum 1.27 (1.23–1.37F)× wider than long. Elytra in males 1.47 Phylogenetic Relationships.—Euchroa soladevega does (1.43–1.54) and in females 1.43 (1.39–1.45)× longer than wide. not appear to be closely related to either of the other two Microsculpture. Elytra with sculpticells on disc slightly elongate and members of the soladevega subgroup, if indeed they belong deeply impressed, entire surface of disc distinctly beaded, sculpticells markedly convex (Figs. 6a–b). to the same subgroup (see phylogenetic analyses below for Head. Submentum prominent. Transgular groove deep posterad sub- details). mentum. Mouthparts. Labial palpomere 3 in males subsecuriform to broadly Chorological Affinities.—Euchroa juchatengo, and also triangular, in females narrowly triangular, in males 1.11 (1.02–1.26) and E. santacatarina (dimidiata subgroup), both occur on the

Pacific side of the western part of the Sierra de Miahuatlan extended entire length of middle part of median lobe, groove shallow (Fig. near the localities where E. soladevega has been found in 36d). Internal sac with apical and basal bands of large microtrichia both the Sierra de Quatro Venados. The two mountain ranges are extended entirely around sac. separated here by only a narrow valley (World Aeronautical Material Examined.—The type series. Chart CJ-24). We (Ball) have collected a specimen of E. soladevega in the bottom of that valley (13.1 miles north Collecting Notes.—We (Ball) collected some individuals of Juchatengo), and so perhaps E. soladevega also occurs in of E. juchatengo in a steep-sided, narrow ravine with forest the western part of the Sierra de Miahuatlan along with of tropical aspect providing dense shade, at an elevation of E. juchatengo and E. santacatarina, but presumably in about 1700 m. Since then, the ravine has been cleared of drier forest, and further inland, than those two species. much of its trees. Bionomics.—A teneral male was collected on July 21–22, 20. Euchroa (Dyschromus) juchatengo, new species 1966. Figs. 36d–e, 37a Geographical Distribution.—This species is known only Type Material.—Six specimens, labeled as follows. HOLOTYPE male, from a small area on the seaward side of the western part of “MEX. OAX.; 20 mi S/ Juchatengo. 6000'/ 28-30.v.[19]71[,] S. Peck,” (USNM). PARATYPES, “MEXICO. Oaxaca/ 5800'. 22.2 mi./ s. the Sierra de Miahuatlan, Oaxaca (Fig. 37a). Juchatengo./ VII.21–22.1966”; “George E. Ball/ D.R. Whitehead/ collectors,” 3M, 2F (UASM). Phylogenetic Relationships.—The nearest relative of E. juchatengo is E. nizavaguiti (see section below on phy- Type Locality.—About 32 km south of Juchatengo, state of Oaxaca, logeny for details). Mexico. Chorological Affinities.—Euchroa santacatarina (dimidi- Diagnosis.—Besides E. juchatengo, two other species of ata subgroup) lives in the same area as E. juchatengo. We the subgenus Dyschromus in western Mexico are uniform- (Ball) have collected individuals of both species in virtually metallic bluish purple in color dorsally. Of these, ly the same handful of leaf litter. Euchroa soladevega E. juchatengo is separable from E. soladevega by its small- might also live in the vicinity, although probably in drier er size, and the microsculpture on the disc of the elytra, forest (see description of that species). A considerable gap which consists of sculpticells that are isodiametric, and separates the geographical ranges of E. juchatengo and its more finely impressed than in E. soladevega. It is distin- presumed sister species, E. nizavaguiti (Fig. 37a). guishable from E. nizavaguiti by the shallower transgular groove, and slightly prominent hind angles of the pronotum. The median lobe of the aedeagus most resembles that 21. Euchroa (Dyschromus) nizavaguiti, new species of E. nizavaguiti, but the preapex is shorter and broadly Figs. 12d, 36f–g, 37a rounded in outline. Type Material.—Nine specimens, labeled as follows. HOLOTYPE male, Description.—Body Size. Individuals relatively small to moderate in size. “MEX. OAXACA[.] n.w. Santa/ Maria Nizavaguiti/ 16º41'N 95º50'W[,] Apparent body length 9.5M–10.5F mm. Length (n=4M,2F): head oak-/ pine for[est]. (dry)/ in litter[,] 1935m./ June 20, 1979[,] 79-43”; 0.83M–0.91F, pronotum 2.28M–2.54, elytra 5.30M–5.95F mm; width: “MEXICAN EXP. 1979/ J.S. Ashe, G.E. Ball,/ & D. Shpeley/ collectors,” head 1.97M–2.20F, pronotum 3.04M–3.37F, elytra 3.40M–4.00F mm. (USNM). PARATYPES, same labels as holotype, 7M, 1F (UASM). Body Proportions. Head 2.3M–2.41F× wider than long. Pronotum 1.29M–1.36F× wider than long. Elytra 1.49F–1.56M× longer than wide. Type Locality.—A few kilometers northwest of Santa Maria Nizavaguiti, Microsculpture. Elytra with sculpticells on disc moderately finely state of Oaxaca, Mexico. impressed, isodiametric, portions of disc with surface slightly beaded and sculpticells slightly convex. Diagnosis.—Euchroa nizavaguiti, one of three species of Head. Submentum slightly prominent. Transgular groove moderately the subgenus Dyschromus in western Mexico that are uni- deep posterad submentum. Mouthparts. Labial palpomere 3 in males narrowly triangular and formly metallic bluish-purple in color dorsally, is only like- females subtriangular, in males 1.67–2.04 and females 2.28–2.33× longer ly to be confused with E. juchatengo, from which it is dis- than wide. tinguishable by the deeper transgular groove and obtuse Prothorax. Pronotum with sides slightly sinuate just at very base of, hind angles of the pronotum. It differs from E. soladevega or along posterior third of, pronotum; hind angles slightly prominent. in the same ways as does E. juchatengo. Pterothorax. Elytral intervals slightly convex. Male. Hind tibia with basal half of medial ridge represented by row of Description.—Body Size. Individuals moderate in size. Apparent body moderately prominent, subangular tubercles. Aedeagus with intermediate length 10.0M–11.0M mm. Length (n=8M,1F): head 0.95 (0.87M–0.97M), symmetric type 4 median lobe of usual proportions (Fig. 36d); preapex pronotum 2.57 (2.38M–2.68M), elytra 6.08 (5.55M–6.30M) mm; width: moderately long and broadly rounded in outline (Fig. 36d), and moderate- head 2.23 (2.0M–2.32M), pronotum 3.53 (3.08M–3.65M), elytra 4.1 ly thick and evenly sclerotized throughout (Fig. 36e), with one short cen- (3.60M–4.25M) mm. trally located primary haemolymph channel and several secondary chan- Body Proportions. Head 2.37 (2.27–2.43)× wider than long. Pronotum nels branched from it (Fig. 36e); left wall of periostial part of median lobe in males 1.37 (1.29–1.40) and female 1.44× wider than long. Elytra in more extensive than and entirely concealing right wall in lateral view, males 1.49 (1.47–1.54) and female 1.43× longer than wide. periostial ventrolateral bulge slightly prominent in profile (as in Fig. 43b) Microsculpture. Elytra with sculpticells on disc moderately finely and subangulate in cross-section (as in Fig. 26c); dorsomedial groove

Phylogenetic Relationships.—Euchroa nizavaguiti is most closely related to E. juchatengo (see description of that species, and section below on phylogeny, for details). Chorological Affinities.—The type-locality for E. nizavaguiti overlooks the Isthmus of Tehuantepec; no other species of the subgenus Dyschromus has been found so far east in the Sierra de Miahuatlan. The other two members of the soladevega subgroup are known only from the westernmost part of the Sierra de Miahuatlan or nearby, so that a considerable gap separates their known geographical ranges from that of E. nizavaguiti (Fig. 37a).

Dimidiata Subgroup Diagnosis.—Species belonging to the dimidiata subgroup have a metallic green head and pronotum and brassy-colored elytra, or the dorsum is entirely metallic green, brassy, or coppery-colored, or the head and pronotum are coppery and the elytra bronze. At least stria 1 is interrupted on the disc of the elytra, except for some specimens of E. zempoalteptl and E. tenancingo, and stria 1 and/ or stria 2 are interrupted several times toward the apex of the elytra.

Description.—Color. Dorsum of head and pronotum metallic green and Fig. 38.—Median lobe of aedeagus of E. (Dyschromus) jalisco n. sp.: elytra brassy, or head and pronotum coppery and elytra bronze, or dorsum dorsal aspect. phc, primary haemolymph channel. Scale bar = 0.8 mm. entirely metallic green, brassy or coppery. Microsculpture. Elytra with most or all sculpticells on disc isodiametric, some slightly transverse in some species, only E. dimidiata with sur- impressed, isodiametric, flat. face of disc beaded and sculpticells convex, and only slightly so (Fig. 5e), Head. Submentum prominent. Transgular groove deep posterad sub- other species with sculpticells on disc flat; extent of beading at sides and mentum. apex of elytra variable, at one extreme some species without beading, all Mouthparts. Labial palpomere 3 in males broadly triangular and in sculpticells flat, at other extreme in a few species outer half of interval 8, female subtriangular, in males 1.32 (1.23–1.49) and female 2.18× longer all of 9, and apex with entire surface beaded, sculpticells markedly con- than wide. vex (as in Fig. 15b). Prothorax. Pronotum with sides entire posteriorly or sinuate very near Head. Frontal furrows with narrow and deeply impressed portion typ- base; hind angles obtuse (Fig. 12d). ically extended posteriorly to level of anterior supraorbital setigerous Pterothorax. Elytral intervals slightly convex. punctures, shorter in E. yucuyacua, longer in some specimens of E. Male. Hind tibia with medial ridge represented nearly to apex by row dimidiata. Postocular transverse groove evident dorsomedially or not. of moderately prominent, subangular tubercles. Aedeagus with extreme Submentum slightly to moderately prominent and sloped evenly to trans- symmetric type 4 median lobe of usual proportions (Fig. 36f); preapex gular groove, except prominent and with posterior face concave in most long and subtruncate apically (Fig. 36f), moderately thick and evenly scle- specimens of E. dimidiata. rotized throughout (Fig. 36g), with one short centrally located primary Mouthparts. Mentum with paramedian pits small to moderate in size. haemolymph channel and several secondary channels branched from it Prothorax. Pronotum with sides entire to moderately sinuate posteri- (Fig. 36g); left wall of periostial part of median lobe more extensive than orly; basolateral impressions short, moderately long, or long. and entirely concealing right wall in lateral view, periostial Pterothorax. Elytra with basal punctures present or not; striae moder- ventrolateral bulge slightly prominent in profile (as in Fig. 43b) and sub- ately deeply or shallowly impressed, at least stria 1 in most specimens angulate in cross-section (as in Fig. 26c); middle part of median lobe with interrupted on disc; intervals flat or slightly convex; apical declivity mod- dorsomedial groove short (Fig. 36f). Internal sac with apical band of large erately steep to nearly perpendicular. microtrichia extended entirely around sac, basal band only evident Abdomen. Sterna V–VIII with basal sulcus only foveate laterally, or laterally. also with median fovea, or entirely foveate; foveae coarse. Male. Middle tibia with inner face expanded near apex or not. Median Material Examined.—The type series. lobe of aedeagus with preapex thicker to right of midline, or uniformly thick throughout, or preapex only developed to right of midline; periostial Collecting Notes.—Euchroa nizavaguiti has been found in ventrolateral bulge slightly to markedly prominent. Internal sac with or dry oak-pine forest at an elevation of about 1900 m. without one or more bulges. Geographical Distribution.—The geographical range of Geographical Distribution.—This species is known only the dimidiata subgroup corresponds to that of the nitidipen- from the very easternmost part of the Sierra de Miahuatlan, nis group as a whole, except that it is not known from east- Oaxaca (Fig. 37a). ern Mexico, north of the Rio Santo Domingo, other than a

Fig. 39.—Median lobe of aedeagus of subgenus Dyschromus, dorsal aspect. A–B, E. tenancingo n. sp.; C–D, E. ixtapa n. sp. Scale bar = 0.8 mm.

specimen of E. dimidiata that is doubtfully from Veracruz Type Locality.—Ca. 19.8 km south of Tecalitlan, state of Jalisco, Mexico. (see description of that species). Diagnosis.—Of the three species of the subgenus Phylogenetic Relationships.—We consider the soladeve- Dyschromus that are uniformly metallic green in color dor- ga subgroup to be the sister group of the dimidiata sub- sally, E. jalisco most resembles E. atoyac, but compared to group. See phylogenetic analyses below for details. both E. atoyac and E. suchixtepec, the green color is clear- er and brighter, the basolateral impressions of the pronotum Chorological Affinities.—The geographical range of the are longer, the mesepisternum has fewer punctures, and dimidiata subgroup encompasses that of the soladevega comparing the same sex, labial palpomere 3 is much broad- subgroup in western Oaxaca, but does not overlap at all er. The median lobe of the aedeagus does not closely with that of the nitidipennis subgroup in eastern Mexico. resemble that of any other species that has an extreme symmetric type 4 median lobe. Taxonomic Composition.—The dimidiata subgroup contains 14 species: E. jalisco n. sp.; E. tenancingo n. sp.; Description.—Body Size. Individuals moderate to relatively large in size. E. ixtapa n. sp.; E. dimidiata Chaudoir; E. atoyac n. sp.; Apparent body length 10.5M–13.0F mm. Length (n=2M,2F): head M F M F M F E. puertogallo n. sp.; E. filodecaballo n. sp.; E. chryso- 0.83 –0.95 , pronotum 2.54 –2.82 , elytra 5.60 –6.50 mm; width: head 1.97M–2.34F, pronotum 3.18M–3.97F, elytra 3.70M–4.65F mm. phana Bates; E. yucuyacua n. sp.; E. santacatarina n. sp.; Body Proportions. Head 2.36M–2.48F× wider than long. Pronotum E. zempoaltepetl n. sp.; E. carbonera n. sp.; E. miahuatlan 1.25M–1.41F× wider than long. Elytra 1.40F–1.51M× longer than wide. n. sp.; and E. suchixtepec n. sp. The last four or five species Color. Dorsum uniformly bright metallic green. may actually belong to the nitidipennis subgroup (for Microsculpture. Dorsum of head smooth in spots, sculpticells effaced. Elytra with most sculpticells on disc isodiametric, some slightly trans- details, see phylogenetic analyses, below). verse, finely impressed, flat; interval 9 and apex with sculpticells moderately deeply impressed, flat. Head. Postocular transverse groove obsolete dorsomedially or barely 22. Euchroa (Dyschromus) jalisco, new species evident. Submentum slightly prominent, posterior face sloped evenly to Figs. 13a, 37b, 38a–b transgular groove; latter moderately deep posterad submentum. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in Type Material.—Four specimens, labeled as follows. HOLOTYPE male, males subsecuriform and female subtriangular, in males 1.02–1.13 and “MEX. Jalisco/ 12.4 mi. s./ Tecalitlan[,] 5300'/ August 3, 1967”; “Ball, female 2.1× longer than wide. Mentum with paramedian pits small. T.L. Erwin/ R.E. Leech/ collectors,” (USNM). PARATYPES, same labels Prothorax. Pronotum with sides moderately sinuate posteriorly; hind as holotype, 1M, 2F (UASM). angles obtuse; basolateral impressions long (Fig. 13a). Prosternal inter-

Fig. 40.—Median lobe of aedeagus of E. (Dyschromus) dimidiata Chaudoir: B, left lateral aspect, other figures in dorsal aspect. A–D, two specimens from Rte. 135, 64.4 km. n.w. Ciudad Oaxaca; E–F, two specimens from microondas station, 0.5 mi. e. jct. Rtes. 190 and 195, Oaxaca; G, specimen from 20 km. n.w. Chilpancingo, Guerrero; H, specimen from Buenavista de Cuéllar, Gro. phc, primary haemolymph channel. Scale bar = 0.8 mm.

coxal process not or indistinctly margined apically. inal sterna V–VII extended to the lateral margins of the Pterothorax. Elytra with basal punctures present; striae 1-2 interrupt- abdomen. It differs further from similarly colored species ed on disc, 1 interrupted 2–3 times (n=4), 2 interrupted 1–2 times (n=2); intervals slightly convex; apical declivity moderately steep. in that the entire basal sulcus of sternum VII is coarsely Mesepisternum with few shallow punctures. foveate. The median lobe of the aedeagus closely resem- Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous; bles that of E. ixtapa. sternum VII without median fovea. Male. Middle tibia with inner face expanded near apex. Hind tibia Description.—Body Size. Individuals moderate in size. Apparent body with most of medial ridge represented by row of prominent subangular length 11.0–12.0 mm. Length (n=5M,6F): head 0.95 (0.87F–1.01M), tubercles. Aedeagus with extreme symmetric type 4 median lobe of usual pronotum 2.80 (2.58F–2.92M), elytra 6.30 (6.00–6.70F) mm; width: head proportions (Fig. 38a); preapex long, broad, and moderately thick and 2.31 (2.18F–2.38F), pronotum 3.67 (3.49F–3.91F), elytra 4.24 evenly sclerotized throughout, apex obliquely subtruncate (Figs. 38a–b), (4.05–4.55F) mm. with one large centrally located primary haemolymph channel (Fig. 38b); Body Proportions. Head 2.44 (2.28M–2.57F)× wider than long. left wall of periostial part of median lobe more extensive than and entire- Pronotum 1.31 (1.27M–1.36F)× wider than long. Elytra 1.48 ly concealing right wall in lateral view, periostial ventrolateral bulge (1.39F–1.57F)× longer than wide. slightly prominent in profile (as in Fig. 43b) and subangulate in cross- Color. Dorsum of head and pronotum metallic green, elytra brassy, or section (as in Fig. 26c); middle part of median lobe without dorsomedial brassy with marked greenish sheen (n=1). groove (Fig. 38a). Internal sac with apical band of large microtrichia Microsculpture. Dorsum without smooth spots, sculpticells evident extended dorsally along midline towards basal band, both bands extended over entire surface. Elytra with sculpticells on disc isodiametric to slight- around sac. ly transverse, finely impressed, flat; portions of interval 9 and apex of elytra beaded, sculpticells convex. Material Examined.—The type series. Head. Postocular transverse groove shallow but distinct dorsomedially. Submentum slightly prominent, posterior face sloped evenly to trans- Collecting Notes.—The type series of E. jalisco was col- gular groove; latter moderately deep posterad submentum. lected in mesophytic forest at about 1600 m elevation. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in males broadly triangular and females subtriangular, in males 1.31 (1.27–1.35) and females 2.25 (2.08–2.33)× longer than wide. Mentum Geographical Distribution.—This species is known from with paramedian pits moderate in size. the western part of the Trans-Volcanic Sierra in Jalisco Prothorax. Pronotum with sides moderately sinuate posteriorly; hind (Fig. 37b). angles slightly prominent; basolateral impressions long, deeply impressed portion nearly extended to hind margin of pronotum. Prosternal intercox- Phylogenetic Relationships.—The closest relatives of al process not margined, or entire apical third shallowly margined (Buenavista Cuellar). E. jalisco could be a group comprising E. tenancingo, Pterothorax. Elytra with basal punctures present; striae entire on disc E. ixtapa, and E. dimidiata, but perhaps E. jalisco occupies (n=3) or stria 1, 1–2, 1–3, 1–4, or 1–5 interrupted on disc, stria 1 interrupt- a more basal position within the dimidiata subgroup. For ed 1–9, 2 and 3 interrupted 0–4, and 4 and 5 interrupted 0–2 times; inter- details, see phylogenetic analyses, below. vals slightly convex; apical declivity moderately steep to nearly perpendicular. Mesepisternum with few or many shallow punctures or nearly Chorological Affinities.—This is the only species of the smooth. Abdomen. Sterna V–VII with basal sulcus extended to lateral margins subgenus Dyschromus known from the western part of the of abdomen; sternum VII with entire basal sulcus coarsely foveate. Trans-Volcanic Sierra. Male. Middle tibia with inner face slightly expanded near apex. Hind tibia with most of medial ridge represented by row of prominent subangular tubercles. Aedeagus with extreme symmetric type 4 median lobe, com- 23. Euchroa (Dyschromus) tenancingo, new species paratively slender in dorsal aspect (Fig. 39a), otherwise of usual propor- Figs. 14d, 37b, 39a–b tions; preapex long, broad, and moderately thick and evenly sclerotized throughout, apex subtruncate (Figs. 39a–b), with one large centrally locat- Type Material.—Twelve specimens, labeled as follows. HOLOTYPE ed primary haemolymph channel (Fig. 39b); left wall of periostial part of male, “MEXICO. Mexico./ Rio de Molino/ nr. Valle de/ Bravo, 6500'/ median lobe more extensive than, and entirely concealing, right wall in August 2–3, 1962”; “H.E. Evans Exp./ George E. Ball Collector,” lateral view; periostial ventrolateral bulge slightly prominent in profile (as (USNM). PARATYPES, same labels as holotype, 1F (UASM). in Fig. 43b) and subangulate in cross-section (as in Fig. 26c); middle part “Tejupilco, MEX./ Temescaltepec”; “H.E. Hinton/ Collector,” 2M of median lobe without dorsomedial groove (Fig. 39a). Internal sac with (MCZC). “H.E. Hinton/ Collector/ Rincon,” 1M (MCZC). “MEX. Mex. apical band of large microtrichia extended dorsally along midline toward 2.7 km ne/ Temazcaltepec, Rte/ 130, stream arroyo/ oak; litter; 1765 m/ basal band; basal band only evident on left side. Aug. 3, 1983[,] 83-61”; “MEXICO EXPED. 1983/ H.E. Frania &/ R.J. Jaagumagi/ collectors,” 1M, 3F (UASM). “MEX. Mexico. 4/ mi. s. Material Examined.—The type series. Tenancingo/ Rte. 55, 7100'/ VIII.7.65”; “George E. Ball/ D.R. Whitehead/ collectors,” 1F (UASM). “MEXICO: GUERRERO/ B[uena]v[is]ta de Remarks.—The two females of E. tenancingo from Cuéllar/ “La Estancia”/ 6.09.1992[,] RN14-1/ 1890 m, R. Nieto”; “157,” Buenavista de Cuellar differ from the others in that the 1F (UAEM); the same, except “13.06.93[,] RN43-1”; “320,” 1F (UAEM). basal sulcus of sternum VII has coarser foveae. Type Locality.—Rio de Molino, near Valle de Bravo, state of Mexico, Mexico. Collecting Notes.—Euchroa tenancingo has been collected in pine-oak forest at elevations of 1800 to 2000 m. Diagnosis.—Euchroa tenancingo, one of seven species of the subgenus Dyschromus in Mexico with a metallic green Geographical Distribution.—This species is known from head and pronotum, and brassy-colored elytra, is the only the central part of the Trans-Volcanic Sierra in the states of species of Dyschromus that has the basal sulcus of abdom- Mexico and Guerrero (Fig. 37b).

Fig. 41.—A, Distal portion of median lobe and everted internal sac of E. (Dyschromus) dimidiata Chaudoir from 64.4 km. n.w. Ciudad Oaxaca, right lateral aspect, except individual microtrichia illustrated from regions indicated on left side of sac. B, individual microtrichia from area of basal band indicated in Fig. 41A, specimen from microondas station, 0.5 mi. e. jct. Rtes. 190 & 195, Oaxaca. Scale bar = 0.8 mm.

Phylogenetic Relationships.—Euchroa tenancingo is Diagnosis.—Euchroa ixtapa has a metallic green head and most closely related to E. ixtapa and E. dimidiata (see phy- pronotum, and brassy-colored elytra. It is distinguishable logenetic analyses below for details). from the six other species of the subgenus Dyschromus from Mexico with this combination of colors by stria 7, as Chorological Affinities.—Euchroa dimidiata also occurs well as striae 1–6 each being interrupted on the disc of the in the central part of the Trans-Volcanic Sierra, but E. ten- elytra, with stria 7 being interrupted at least five times, and ancingo is found at higher elevations and in more mesic in most specimens about 25 times. Otherwise, only some forest. specimens of E. dimidiata have stria 7 interrupted, and then only once. The median lobe of the aedeagus most resem- 24. Euchroa (Dyschromus) ixtapa, new species bles that of E. tenancingo. Figs. 37b, 39c–d Description.—Body Size. Individuals moderate in size. Apparent body length 11.0M–12.0M mm. Length (n=5M,3F): head 0.89F–0.99M, prono- Type Material.—Seven specimens, labeled as follows. HOLOTYPE tum 2.56F–2.92M, elytra 5.80M–6.20M mm; width: head 2.16M–2.32M, male, “MEX. Gro. 78.5 km/ n jct Rte 200 on/ Rte 134[,] ridge top/ oak- pronotum 3.43M–3.81M, elytra 4.00M–4.35F mm. pine; 1770 m./ 11.VIII.1985[,] 46-85”; “MEXICAN EXP. 1985/ H.E. Body Proportions. Head 2.24M–2.53F× wider than long. Pronotum Frania &/ D. Shpeley/ collectors,” (USNM). PARATYPES, same labels as F F M M F 1.28–1.37 × wider than long. Elytra 1.36 –1.50 × longer than wide. holotype, 1 , 1 (UASM). “MEX. Guerrero 78.5/ n jct Rte 200 on Rte/ Color. Dorsum of head and pronotum metallic green with bluish tint, 134 to Ciudad Altami/ rano; oak-pine; 1770 m/ leaf litter; under logs/ July elytra brassy, with (n=3) or without greenish tint (n=2). 30 [or 31], 1992[,] 31-92”; MEXICAN EXP. 1992/ J.S. Ashe/ H.E. Frania/ M F Microsculpture. Dorsum without smooth spots, sculpticells evident D. Shpeley colls.,” 3 , 1 (UASM). over entire surface. Elytra with sculpticells on disc isodiametric to slightly transverse, finely impressed, flat; small portions of interval 9 and apex Type Locality.—Crest of Sierra Madre del Sur at Route 134, northeast of of elytra beaded (sculpticells convex). Ixtapa, state of Guerrero, Mexico. Our choice of the coastal town of Head. Postocular transverse groove shallow but distinct dorsomedial- Ixtapa as a map reference for the type locality is due to the fact that ly. Submentum slightly prominent, posterior face sloped evenly to trans- although we (Frania) travelled the length of Route 134 both in 1985 and gular groove; latter moderately deep posterad submentum. 1992, we did not pass through any towns or villages, nor did we see any Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in signs indicating roads to communities located off the highway. males broadly triangular and in females subtriangular, in males 1.09–1.32 and females 2.25× longer than wide. Mentum with paramedian pits small

Fig. 42.—Maps of central Mexico, showing geographical distributions of some species of the subgenus Dyschromus belonging to the dimidiata subgroup.

to moderate in size. num VII, the entire basal sulcus is foveate; the inner face Prothorax. Pronotum with sides moderately sinuate posteriorly; hind of the subapical part of the hind tibia of this male is not angles slightly prominent; basolateral impressions long. Prosternal intercoxal process not margined. expanded medially. The median lobe of the aedeagus is Pterothorax. Elytra with basal punctures; striae 1–7 all interrupted reminiscent of that of E. puertogallo in that the preapex is about 25 times on disc; intervals nearly flat; apical declivity steep to near- comparatively short. The peculiar nature of the specimen ly perpendicular. Mesepisternum with many shallow to coarse punctures. is evident in the condition of the elytral striae because in Abdomen. Sterna V–VII with foveae in basal sulcus coarse; basal sul- all other individuals of the subgenus Dyschromus that cus of sternum VII without median fovea. Male. Middle tibia with inner face very slightly expanded near apex. have interrupted elytral striae, the innermost striae are Hind tibia with most of medial ridge represented by row of prominent sub- broken at least as many times and usually more times than angular tubercles. Aedeagus with extreme symmetric type 4 median lobe, the outer ones. It remains to be determined what the sta- comparatively slender in dorsal aspect (Fig. 39c), otherwise of usual pro- tus is of this individual. portions; preapex long, broad, and moderately thick and evenly sclerotized throughout, apex subtruncate (Figs. 39c-d), with one large centrally located primary haemolymph channel (Fig. 39d); left wall of periostial Collecting Notes.—All the specimens of E. ixtapa were part of median lobe more extensive than and entirely concealing right wall found in rather open oak-pine forest at about 1800 m ele- in lateral view, periostial ventrolateral bulge slightly prominent in profile vation, either under logs, or in leaf litter near a small (as in Fig. 43b) and subangulate in cross-section (as in Fig. 26c); middle stream. part of median lobe without dorsomedial groove (Fig. 39c). Internal sac with microtrichia on left side of basal band of large microtrichia mostly triangular or transverse in shape, a few drawn out into a very short spine. Geographical Distribution.—This species is known from the westernmost Sierra Madre del Sur in Guerrero Material Examined.—The type series, and one additional male collect- (Fig. 37b). The type locality is on the continental divide. ed at the type locality on July 31, 1992. This specimen differs sufficiently from the others that it is not included in the type series (see below). Phylogenetic Relationships.—Euchroa ixtapa is most closely related to either E. tenancingo or E. dimidiata (see Remarks.—The specimen not included in the type series phylogenetic analyses below for details). of E. ixtapa is smaller than the others, labial palpomere 3 is broader, and so is the pronotum, as follows: apparent Chorological Affinities.—No other member of the sub- body length, 10.0 mm; labial palpomere 3, 0.93× wider genus Dyschromus has been found so far west in the than long; pronotum 1.40× wider than long. The elytral Sierra Madre del Sur, but E. dimidiata probably also striae are not so interrupted on the disc of the elytra, with occurs in the vicinity, considering its geographical distri- stria 1 interrupted 1–4 (right elytron), striae 2–4 interrupt- bution and habitat preferences. ed 0–1, striae 5–6 interrupted 2–4, and stria 7 interrupted 6–13 times. Other differences are that on abdominal ster-

25. Euchroa (Dyschromus) dimidiata Chaudoir palpomere 3 in males broadly and females narrowly triangular, in males Figs. 1b, 3d, 5d–f, 9d, 14b–c, 16a–b, 17b, 26c, 1.32(1.21–1.53) and females 1.82(1.66–1.90)× longer than wide. Mentum with paramedian pits small. 40a–h, 41a–b, 42a Prothorax. Pronotum with sides in most specimens slightly or moderately sinuate posteriorly (Fig. 1b) and hind angles slightly prominent, Euchroa dimidiata Chaudoir 1874:17. Type material. Three speci- sides in a few individuals oblique with hind angles obtuse; basolateral mens, in the Oberthür-Chaudoir collection (MNHP), associated impressions short. Prosternal intercoxal process with apical third deeply with the following handwritten label that was pinned originally in margined all around (n=22) or just laterally (n=1). the bottom of an insect box: “dimidiata/ Chaud/ Mexique/ Oaxaca Pterothorax. Elytra with basal punctures; stria 1-2, 1-3, 1-4, or 1-5 Sallé.” LECTOTYPE male, two PARALECTOTYPES, male and interrupted on disc, 1 interrupted 24–32 times, 2 interrupted 3–28, 3 and 4 female, each labeled “Ex Musaeo/ Chaudoir” [red print]. Bates, each interrupted 0–19, 5 interrupted 0–12 times, 6 nearly interrupted 3 1882:85. times on one elytron (n=1); intervals nearly flat; apical declivity nearly Dyschromus dimidiatus (Chaudoir): Tschitschérine, 1898:60. Liebherr, perpendicular (Fig. 14b). Mesepisternum with a few shallow punctures. 1994:842. Abdomen. Sterna V–VII with foveae in basal sulcus coarse; basal sulcus of sternum VII without median fovea (Fig. 14c). Notes about Type Material.—Chaudoir (1874:17), recorded in associa- Male. Middle tibia with inner face distinctly expanded near apex (Fig. tion with the original description of this species, three specimens, from 16a). Hind tibia with most of medial ridge represented by a row of promi- Oaxaca, received from Augustin Sallé. The lectotype (here selected) is the nent subangular tubercles (Fig. 16b). Aedeagus with extreme symmetric first male in the type series. type 4 median lobe of usual proportions (Figs. 40a, 40d); preapex long, broad, and moderately thick and evenly sclerotized throughout, apex sub- Type Locality.—The type area, as determined from the original descrip- truncate (Figs. 40a, 40c–d), with one large centrally located primary tion (Chaudoir, 1874:18) is “Oaxaca,” the name of one of the larger of the haemolymph channel (Fig. 40c); left wall of periostial part of median lobe Mexican states. The largest city in that state is also named Oaxaca, and the more extensive than, and entirely concealing, right wall in lateral view type specimens probably were collected near that city. Accordingly, we (Fig. 40b); periostial ventrolateral bulge moderately prominent in profile arbitrarily restrict the type locality to the environs of Oaxaca city. (Fig. 40b) and subangulate in cross-section (Fig. 26c); middle part of median lobe with dorsomedial groove not evident in most specimens, Diagnosis.—Specimens of E. dimidiata from most locali- present but indistinct in some individuals. Internal sac with microtrichia ties are distinguishable from the six other species of the on left side of basal band of large microtrichia each extended into a long subgenus Dyschromus in Mexico that have a metallic green spine (Fig. 41a). head and pronotum and brassy-colored elytra by the unusu- Material Examined.—82 specimens. OAXACA. Rte. 131[135], 64.4 ally long frontal furrows on the head, the very prominent km. n.w. Oaxaca, oak forest, 1930 m, July 17, 1975, G.E. Ball & H.E. submentum that is concave posteriorly, the microsculpture Frania, 7M (UASM); 38.4 km. n. Telixlahuaca [same site as above], oak- on the disc of the elytra which consists of sculpticells that palmetto woodland, 1,880 m., under stones, July 19, 1992, 23-92, J.S. are relatively deeply impressed, and the nearly perpendicu- Ashe, H.E. Frania, D. Shpeley, 3M, 7F, 1? (UASM); 24.2 km. s.w. Yolomecatl, Rte. 125, oak-pine madrono, 2,120 m., 20 August 1992, 64- lar apical declivity of the elytra. The large number of times 92, G.E. Ball & H. Frania, 1M, 1F (UASM); microondas, 4.2 km. N. jct that the elytral striae are interrupted on the disc is only Hwys. 125 & 190, 2900 m., Aug. 10, 1988, J.K. Liebherr, D.A. Yager, 3M matched by E. tenancingo, and exceeded by E. ixtapa. The (CUIC); Microondas Sta[tion], 0.5 mi. e. jct. Rtes. 190 & 195, ca. 8,300', median lobe of the aedeagus of E. dimidiata most resem- oak forest, August 23 & 24, 1972, B.S. Heming, G.E. Ball, 6M, 4F bles that of E. ixtapa and E. tenancingo, but in nearly all (UASM); the same except, July 31, 1974, D.R. Whitehead, H. Frania & G.E. Ball, 1M, 3F (UASM); the same except, VI.10.1979, 79-27, J.S. Ashe, specimens is not so slender. G.E. Ball, & D. Shpeley, 3M, 6F (UASM); the same except, 7.4 km. n. Santiago Tejapan, oak forest (dry) on ground, 2,350 m., in litter, 79-29, Description.—Based on specimens from eastern Oaxaca west to June 11, 1979, 3M (UASM); the same except, 7.9 km. n.w. La Carbonera, Nochixtlan. Body Size. Most individuals moderate to relatively large in Rte. 190, oak-pine for[est]. (dry), 2,250 m., in litter, 79-29 June 11, 1979, M M M F size. Apparent body length 9.0 –13.0 mm. Length (n=15 ,8 ): head 2M, 1F (UASM); the same except, 15.7 km. s. Rte. 190, rd. to Ojo de M M M M M 1.00(0.83 –1.11 ), pronotum 2.89(2.50 –3.14 ), elytra 6.58(5.50 – Agua, oak-pine zone, Alnus near stream in litter, 2,320 m., June 12, 1979, M M M F 7.20 ) mm; width: head 2.48(2.12 –2.70), pronotum 4.00(3.45 –4.29 ), 79-31, 1M (UASM); Hwy. 190, 9.4 km. S. Nochixtlan, 2,300 m., dry oak M F elytra 4.57(3.75 –5.00 ) mm. forest, Aug. 11, 1988, J. K. Liebherr & D. A. Yager, 2M (CUIC); the same M F Body proportions. Head 2.47(2.41 –2.62 )× wider than long. except, Hwy. 135, 23.6 km N Jct. 190 at Huitzo, 20 Aug 1988, el. 2,210 Pronotum in males 1.37(1.34–1.44) and females 1.42(1.38–1.44)× wider m., 1M, 1F (CUIC); 10.6 km. n. jct. #190 & 135, on #135, wet oak than long. Elytra in males 1.46(1.40–1.53) and females 1.41(1.40–1.46)× for[est]., 1,920 m., 21.VII.1987, R.S. Anderson, 1F (UASM); 14.3 km. e. longer than wide. Ixtlan de Juarez, 2,030 m., dry pine-oak for[est]., July 21, 1975, G.E. Ball Color. Dorsum of head and pronotum bright metallic green, bluish & H.E. Frania, 4M, 3F (UASM); 7.2 mi. E. Oaxaca on Papaloapan Road reflection evident in some specimens, elytra brassy, often with greenish at km. 224, 6,140 ft., 10 Sept. 1961, Hubbell, Cantrall, Cohn, #90, 2M reflection (Fig. 3d). (MSUC); 2 km. w. Capulalpam, oak-pine for[est]. (dry) in litter, 2,010 m., Microsculpture. Dorsum without smooth spots, sculpticells evident VI.13.1979, 79-32, J.S. Ashe, G.E. Ball & D. Shpeley, 1F (UASM); Hwy. over entire surface. Elytra with sculpticells on disc isodiametric, moder- 175, 7.5 km. N. El Punto, 2,130 m., Aug. 18, 1988, J.K. Liebherr, D.A. ately deeply impressed, surface of disc slightly beaded, sculpticells slight- Yager, 1M (CUIC); the same except, 18.3 km. S. Gueletao, 2,300 m., 1F ly convex (Figs. 5d–e); small portions of posterior third of interval 9 and (CUIC); Sierra Mixteca, O.A. Purpus S.V., 1F (MNHB). GUERRERO. apex of elytra distinctly beaded (sculpticells moderately convex, Fig. 5f). Amula, 6,000 ft., Aug., H.H. Smith, 1891-64, Euchroa dimidiata Chaud., Head. Frontal furrows with narrow and deeply impressed portion 1M (BMNH); Buenavista de Cuéllar, Venta de la Negra, 8-II-1992, RN extended posteriorly two-thirds to almost entire distance from clypeus to 35.3, 1,400 m., R. Nieto, 2M, 1F (UAEM); 20 km. n.w. Chilpancingo, rd. postocular transverse groove (Fig. 1b). Postocular transverse groove shal- to Chichihualco, oak-acacia, arroyo, 1,600 m, Aug. 7, 1983, 83-66, H.E. low but distinct dorsomedially. Submentum very prominent (Fig. 9d), Frania & R.J. Jaagumagi, 1M (UASM); the same except, oak-palmetto- posterior face concave (as in Fig. 10b); transgular groove deep posterad Agave, under stones, 1,540 m, 12-13.VIII.92, 48-92, G.E. Ball, H.E. submentum. Frania, 1M (UASM); Omilteme, 8,000 ft., July, H.H. Smith, Euchroa Mouthparts. Dorsal mandibular grooves long (Fig. 1b). Labial dimidiata Chaud., 1F (BMNH). MORELOS: Tepoztlan, 1,650 m.,

22.VI.1941, C. Bolivar, 1F (SLSC); 3.6 mi. e. of Cuernavaca, 4,600', intercoxal process margined only laterally, indistinctly so pedregal, 24.XI.1965, George E. Ball, D.R. Whitehead, 1F (UASM). in some specimens. LOCALITY UNCERTAIN: VERACRUZ: El Encanto, Tlapacoyan, 8.VIII.1953, A. Bolivar (Alo-luz), 1M (UASM). LOCALITY A difference common to all specimens from outside UNKNOWN: Istepec, Sallé collection, Euchroa dimidiata Chaud., 1F Oaxaca is that the microsculpture on the body and espe- (BMNH). cially the elytra consists of sculpticells that are not as deeply impressed, hence the specimens have a brighter Remarks.—The specimen of E. dimidiata labeled as being luster. Differences from one area to another outside of from “El Encanto, Tlapacoyan, Veracruz,” is distinctive Oaxaca are also apparent. morphologically (see below), but we suspect that it was The two males and one female from the Sierra Madre collected elsewhere because in our experience, E. dimidia- del Sur west of Chilpancingo, Guerrero (i.e., road to ta inhabits much drier forest formations than we have seen Chichihualco; Omilteme) are unique in the extent to which in the mountains around Tlapacoyan. the elytral striae are interrupted, these being the only specimens of E. dimidiata that have elytral stria 6 and 7 inter- Collecting Notes.—All collections of this species that we rupted: 6 interrupted 2–9 and 7 interrupted 0–1 times; other have made were in the dry, scrubby oak-palmetto, oak, and striae interrupted 23 (stria 5) to 34 (stria 1) times. They also oak-pine woodlands that fringe the Rio Balsas Basin, and differ from all but one other specimen (from Amula, Valley of Oaxaca, at elevations of about 1400 to 2500 m. Guerrero) in that the posterior face of the submentum Bionomics.—During 1992, a teneral adult female was col- slopes evenly to the transgular groove. Next to the speci- lected on July 19 (38.4 km north of Telixlahuaca, Oaxaca), men supposedly from Tlapacoyan, Veracruz, they have the M F and a male on November 8 (Buenavista de Cuéllar, longest and narrowest pronotum: length 2.92 –3.22 mm, M Guerrero). 1.32–1.33 × wider than long. Labial palpomere 3 of the two males is broader than in any other males of E. dimidi- Geographical Distribution.—This species is known from ata, being 1.06× longer than wide. As in some specimens the interior of Oaxaca and Guerrero, and also from from eastern Oaxaca, the sides of the pronotum are oblique Morelos, and possibly Veracruz (Fig. 42a). A curious fea- posteriorly; also, the large microtrichia at the base of the ture of its distribution is that it is known from the northern, internal sac are spinose, although the spines are shorter. As but not the southern, fringes of the Valley of Oaxaca. in many specimens from westernmost Oaxaca, the prosternal intercoxal process is just margined laterally and indis- Geographical Variation.—Chaudoir described E. dimidi- tinctly so. Finally, the preapex is wider relative to the rest ata on the basis of specimens that were probably collected of the median lobe (Fig. 40g) compared to males from in the Sierra Mixteca Alta of east-central Oaxaca, and indi- Oaxaca (Figs. 40a–f). viduals from that state appear to be uniform morphological- The male specimen from the Sierra Madre del Sur east ly as far west as Nochixtlan. Specimens from westernmost of Chilpancingo (Amula) differs from most other speci- Oaxaca differ somewhat, and those from other parts of the mens in that the apical declivity of the elytra is only mod- range even more so, as follows. erately steep, while as in those from west of Chilpancingo, Males from the westernmost localities in Oaxaca for the posterior face of the submentum slopes evenly to the this species (i.e., 7.4 km north of. Santiago Tejapan, 0.5 transgular groove, and the prosternal intercoxal process is miles east of jct. Rtes. 190 and 125, and 24.2 km southwest margined only laterally. of Yolomecatl) have a more slender median lobe of the The five specimens from the central part of the Trans- aedeagus than other males from Oaxaca (Figs. 40e–f), and Volcanic Sierra (i.e., state of Morelos, and Buenavista de on the left side of the internal sac the large microtrichia Cuéllar, Guerrero) resemble the male from Amula, that comprise the basal band are mostly triangular to trans- Guerrero in that the apical declivity of the elytra is only verse in shape, only a few being drawn out into a very moderately steep, but as in many specimens from western- short spine (Fig. 41b). On average, individuals of both most Oaxaca, the head and pronotum have a bluish reflec- sexes are smaller, the pronotum is broader, and labial tion, and the posterior face of the submentum is only palpomere 3 is narrower, as follows: Length (n=13M,10F): slightly concave. The median lobe of the aedeagus of one head 0.91(0.77F–0.99M), pronotum 2.57(2.16F–2.78), ely- of the two males from Buenavista de Cuéllar has a unique tra 6.01(4.95F–6.5) mm; width: head 2.27(1.89F–2.46F), shape (Fig. 40h), but the form of the other is typical of pronotum 3.70(3.18F–4.07F), elytra 4.17(3.40F–4.65F) males from westernmost Oaxaca (as in Fig. 40e). mm. Pronotum 1.44(1.34M–1.56M)× wider than long. Otherwise, except that one is much larger in size, the two Labial palpomere 3 in males 1.41(1.28–1.60) and females males cannot be distinguished from one another, and so we 1.93(1.64–2.11)× longer than wide. More than half of the consider them to be conspecific. specimens, but not necessarily the same ones, exhibit sev- The specimen supposedly from near Tlapacoyan, eral other differences: head and pronotum with bluish Veracruz, a male, resembles individuals from eastern reflection; posterior face of submentum not so concave; Oaxaca in most respects, but is intermediate between them sides of pronotum more sinuate posteriorly; prosternal and those from westernmost Oaxaca for narrowness of the

Fig. 43.—Median lobe of aedeagus of E. (Dyschromus) atoyac n. sp.: B, left lateral aspect, other figures in dorsal aspect; A–C, specimen from type- locality; D–E, another specimen from type-locality. Scale bar = 0.8 mm.

aedeagus and shape of the microtrichia of the internal sac. often evident, the pronotal basolateral impressions are It has the largest and proportionately narrowest pronotum shorter, the mesepisternum has more punctures, and com- of any specimen of this species, and the longest elytra: paring the same sex, labial palpomere 3 is narrower. The pronotum 3.35 mm long, 4.37 mm wide, 1.30× wider than median lobe of the aedeagus most resembles that of long; elytra 7.4 mm long; and only one of two specimens E. puertogallo, but is somewhat stouter. from eastern Oaxaca have the pronotum as broad at the base. Description.—Body Size. Individuals moderate in size. Apparent body M F M F M F Despite the morphological differences between speci- length 10.5 –12.5 mm. Length (n=10 ,10 ): head 0.93(0.81 –1.01 ), pronotum 2.73(2.42M–2.94F), elytra in males 6.01(5.40–6.40) and in mens from westernmost Oaxaca and those from further females 6.60(6.30–6.80) mm; width: head 2.26(1.95M–2.42F), pronotum east in that state, mDNA sequence data indicates that they 3.68(3.24M–3.97F), elytra in males 4.05(3.65–4.35) and females are conspecific (Sperling, Frania, and Ball, in prep.). 4.56(4.40–4.75) mm. Specimens from around Chilpancingo and from the Trans- Body Proportions. Head 2.38(2.26M–2.55M)× wider than long. Pronotum 1.34(1.29M–1.40F)× wider than long. Elytra 1.47(1.42F– Volcanic Sierra should also be examined in this way to 1.54M)× longer than wide. verify that they too are conspecific with the others. Color. Dorsum uniformly dark metallic green, with faint reddish purple reflection (n=1), head with obvious reddish purple reflection (n=1), Phylogenetic Relationships.—Euchroa ixtapa and E. ten- lateral and basal portions of pronotum and lateral portions of elytra also ancingo are the closest relatives of E. dimidiata (see phylo- with obvious reddish purple reflection (n=1); dorsum except for disc of genetic analysis below for details). pronotum with reddish purple reflection (n=1); dorsum with pronounced reddish purple reflection, especially disc of pronotum (Fig. 3a, n=5). Microsculpture. Dorsum of head of some specimens smooth in spots, Chorological Affinities.—Euchroa dimidiata occurs for sculpticells effaced. Elytra with sculpticells on disc isodiametric to slight- the most part, in drier, more scrubby woodlands, either fur- ly transverse, finely impressed, flat; portions of apex of elytra beaded, ther inland, or at lower elevations, than other species of sculpticells convex. Dyschromus in its range, but it has been collected on sever- Head. Postocular transverse groove shallow but distinct dorsomedially. Submentum slightly prominent, posterior face sloped evenly to trans- al occasions together with E. carbonera in dry oak-pine gular groove; latter moderately deep posterad submentum. forest in western Oaxaca. Mouthparts. Dorsal mandibular grooves long. Mentum with paramedian pits small. Labial palpomere 3 in males narrowly triangular and females subtriangular, in males 1.67(1.50–1.81) and females 2.41 26. Euchroa (Dyschromus) atoyac, new species (2.23–2.58)× longer than wide. Figs. 3a, 42a, 43a–e Prothorax. Pronotum with sides moderately sinuate posteriorly; hind angles obtuse to slightly prominent; basolateral impressions usually mod- Type Material.—Twenty-three specimens, labeled as follows. HOLO- erately long, rarely long. Prosternal intercoxal process with apical third TYPE male, “MEX. Guer. 66.4 km/ ne Atoyac de Alvarez/ cl[ou]d distinctly margined all around, or indistinctly or not margined. for[est].-mont[ane]. rain/ for[est]. [transition]; litter; 1925 m/ Aug. 13, Pterothorax. Elytra with basal punctures present; stria 1 not interrupt- 1983[,] 83-77”; “MEXICO EXPED. 1983 H.E. Frania &/ R.J. Jaagumagi/ ed on disc (n=3), or interrupted once (n=2), or 2–3 times and with numer- collectors,” (USNM). PARATYPES, same labels as holotype, 1M, 2F ous near breaks (n=4), other striae not interrupted on disc; intervals slight- (UASM); the same except, “58.4 km/ ne Atoyac de Alvarez/ mont[ane]. ly convex or nearly flat on disc, slightly convex posterolaterally; apical tropical forest/ arroyo; litter; 1400 m/ ... 83-78,” 1M (UASM); the same declivity moderately steep. Mesepisternum coarsely punctate. except, “75.1 km ne Atoyac de/ Alvarez, cloud for[est]./ for. litter, 1890 Abdomen. Sterna V–VII with foveae in basal sulcus coarse; basal sul- m./ 15.VIII.1986[,] 86-54”; “MEXICO EXPED./ G.E. Ball,/ H.E. Frania cus of sternum VII rarely with small median fovea (n=1F). &/ D.S. Mulyk colls,” 1M, 1F (UASM); the same except, “64.9 km. ne. Male. Middle tibia with inner face slightly to markedly expanded near Atoyac/ de Alvarez, trop[ical]. ever-/ green [forest], ravine, lit-/ ter, 1402 apex. Hind tibia with most of medial ridge represented by row of promi- m. 86-44/ August 14, 1986,” 1M, 1F (UASM). “MEX. Guerrero 63.2 km/ nent subangular tubercles. Aedeagus with extreme symmetric type 4 ne Atoyac de Alvarez/ trop[ical] evergreen for[est]./ leaf litter; 1300 m/ median lobe of usual proportions (Figs. 43a–b, 43d); preapex moderately July 28, 1992[,] 27-92”; “MEXICAN EXP. 1992/ J.S. Ashe,/ H.E. Frania,/ long, broad, and moderately thick and evenly sclerotized throughout (Figs. D. Shpeley colls.,” 1M (UASM); the same except, “...71 km/ ne Atoyac de 43a, 43c–d), apex subtruncate (Figs. 43a, 43d), with one large centrally Alvarez/ cloud forest; 1700 m/ leaf litter; under logs/ July 26, 1992[,] 28- located primary haemolymph channel (Figs. 43c, 43e); left wall of perios- 92” [and with rearing data label], 2M, 5F (UASM). “MEXICO GUER- tial part of median lobe more extensive than and entirely concealing right RERO/ 24 km NW El Paraiso/ 1680 m. 7 Aug 1986/ R. Davidson, J. wall in lateral view, periostial ventrolateral bulge slightly prominent in Rawlins, 3M, 3F (CMNH). profile (Fig. 43b) and subangulate in cross-section (as in Fig. 26c); middle part of median lobe without dorsomedial groove (Figs. 43a, 43d). Type Locality.—66.4 km northeast of Atoyac de Alvarez, state of Internal sac with apical band of large microtrichia connected dorsally Guerrero, Mexico. along midline with basal band, both extended around sac. Material Examined.—The type series. Diagnosis.—Euchroa atoyac is one of three species of the subgenus Dyschromus in Mexico that are entirely metallic Collecting Notes.—Euchroa atoyac has been collected in green in color dorsally. Of these, E. atoyac is distinguished tropical evergreen and mesophytic forest at elevations of from E. suchixtepec by the elytral striae, which are uninter- about 1300 to 1900 m. rupted or nearly so on the disc, and are all moderately deeply impressed, and by the presence on the disc of mesh Bionomics.—Two teneral specimens, a male and a female, microsculpture. Compared to E. jalisco, E. atoyac is col- were collected on August 14–15, 1986. ored a darker shade of green, and a purple reflection is

Fig. 44.—Median lobe of aedeagus of E. (Dyschromus) puertogallo n. sp.: B and E, left lateral aspect, other figures in dorsal aspect; A–C, specimen from type-locality; D–F, specimen from 70.8 km n.e. of Atoyac de Alvarez that is not included in type-series. Scale bar = 0.8 mm.

Geographical Distribution.—This species is known from not at all on one elytron, other striae usually entire on disc, stria 2 some- the western versant of the Sierra Madre del Sur in Guerrero, times interrupted once on one elytron; intervals flat on disc, slightly convex posterolaterally; apical declivity with gentle slope. Mesepisternum and more precisely, the lower slopes of the Sierra de Atoyac coarsely punctate. (Fig. 42a). Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous to moderately coarse; basal sulcus of sternum VII without median fovea. Phylogenetic Relationships.—We propose that Euchroa Male. Middle tibia with inner face not expanded near apex. Hind tibia atoyac occupies the basal position in a lineage that also with most of medial ridge represented by row of prominent subangular includes E. puertogallo, E. filodecaballo, E. chrysophana, tubercles. Aedeagus with intermediate symmetric type 4 median lobe of usual proportions (Figs. 44a–b); preapex moderately long, broad, moder- E. yucuyacua, and E. santacatarina (see phylogenetic ately thick and evenly sclerotized throughout, apex subtruncate (Figs. 44a, analyses below for details). 44c), with one large, centrally located primary haemolymph channel (Fig. 44c); left wall of periostial part of median lobe more extensive than, and Chorological Affinities.—Euchroa atoyac occurs at lower entirely concealing, right wall in lateral view, periostial ventrolateral bulge elevations, and in forest of more tropical aspect than any slightly prominent in profile (Fig. 44b) and subangulate in cross-section (as in Fig. 26c); dorsomedial groove absent (Fig. 44a). Internal sac with other species of the subgenus Dyschromus in the Sierra de large bulge on left side at middle; apical and basal bands of large Atoyac. microtrichia connected dorsally and ventrally, both extended around sac.

Material Examined.—The type series, and one other male specimen 27. Euchroa (Dyschromus) puertogallo, new species labeled “MEX. Guer. 70.8 km/ ne Atoyac de Alvarez/ cloud for[est].; Figs. 42b, 44a–f slope;/ leaf litter[,] 2150 m./ Aug. 12, 1983[,] 83-76”; “MEXICO EXPED. 1983/ H.E. Frania &/ R.J. Jaagumagi/ collectors,” (UASM). Type Material.—Seven specimens, labeled as follows. HOLOTYPE male, “MEXICO[,] Guerrero[,] 32 km/ nw Filo de Caballo/ oak-pine Remarks.—The male not included in the type series of E. for[est]., litter/ & roadside; 1950 m,/ Aug. 5, 1992[,] 34-92”; “MEXICAN puertogallo differs considerably from the other specimens. EXP. 1992/ G.E. Ball and/ H.E. Frania colls.,” (USNM). PARATYPES, It is smaller in size (apparent body length 9.5 mm), and GUERRERO: same labels as holotype, and also longevity and egg-laying data for four of the specimens, 4M, 2F (UASM). compared to the other males, the pronotum is quite broad (PL/ PWm: 1.41). Also, the pronotal medial impression Type Locality.—32 km northwest of Filo de Caballo, state of Guerrero, extends to the hind margin of the pronotum, the hind angles Mexico. of the pronotum are obtuse, and the inner face of the middle tibia is toward the apex expanded medially. The other spec- Diagnosis.—Euchroa puertogallo is distinguished from imens have the usual condition in the subgenus Dyschromus the other three species of the subgenus Dyschromus in for the pronotal medial impression, that is, it extends poste- Mexico that are uniformly brassy in color dorsally, i.e., riorly to about the midpoint of the basolateral impressions. E. filodecaballo, E. chrysophana, and E. santacatarina, by The median lobe of the aedeagus is smaller, more slender, the sides of the pronotum, which are more sinuate posteri- and differs somewhat in shape (Figs. 44d–f). We are not cer- orly, and by the elytral striae, fewer of which are interrupt- tain whether this specimen is conspecific with the others, ed on the disc, or at least are interrupted fewer times. The and so have omitted it from the type series. median lobe of the aedeagus most resembles that of E. atoyac, but is more slender. Collecting Notes.—We collected the type series in mesic Description.—Body Size. Individuals moderate in size. Apparent body oak-pine forest at about 2000 m elevation on the east length 10.5M–12.5F mm. Length (n=4M,2F): head 0.87–0.91M, pronotum (inland) side of the Sierra de Atoyac. The other specimen 3.26M–3.61F, elytra 5.80M–6.35F mm; width: head 2.08M–2.42F, prono- was also found at about 2000 m elevation, but in cloud for- tum 3.26M–3.61F, elytra 3.80M–4.35F mm. est on the west (seaward) side of the Sierra de Atoyac. Body Proportions. Head 2.37M–2.71F× wider than long. Pronotum 1.28M–1.41F× wider than long. Elytra 1.46F–1.53M× longer than wide. Geographical Distribution.—This species is known only Color. Dorsum brassy, with slight greenish tint (as in Fig. 3e). Microsculpture. Dorsum of head and pronotum smooth in spots, from the Sierra de Atoyac in southern Guerrero (Fig. 42b). sculpticells effaced. Elytra with sculpticells on disc isodiametric to slightly transverse, finely impressed, flat; portions of interval 9 and apex of ely- Phylogenetic Relationships.—We postulate that E. puer- tra beaded (sculpticells convex). togallo occupies the second most basal position in a line- Head. Postocular transverse groove shallow to nearly obsolete dorso- age that also includes E. atoyac, E. filodecaballo, E. medially. Submentum slightly prominent, posterior face sloped evenly to transgular groove; latter moderately deep posterad submentum. chrysophana, E. yucuyacua, and E. santacatarina, with E. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in atoyac having the basal position. For details, see phyloge- males narrowly triangular and females narrowly subtriangular, in males netic analyses, below. 1.43–1.59 and females 2.6–3.0× longer than wide. Mentum with paramedian pits moderate in size. Chorological Affinities.—Euchroa filocaballo also lives Prothorax. Pronotum with sides moderately sinuate posteriorly; hind on the inland side of the Sierra de Atoyac at the same ele- angles slightly prominent; basolateral impressions moderately long, extended to posterior margin or nearly so as shallow groove. Prosternal vations as E. puertogallo, although the two species have intercoxal process with apex not or indistinctly margined. not been collected together (see description of that species Pterothorax. Elytra with basal punctures; stria 1 in most specimens for details). Euchroa chrysophana lives at the same eleva- interrupted 2–6 times on disc, in a few specimens interrupted just once or

Fig. 45.—Median lobe of aedeagus of subgenus Dyschromus: B and E, left lateral aspect, other figures in dorsal aspect; A–C, E. filodecaballo n. sp.; D–F, E. chrysophana Bates. dmg, dorsomedial groove. Scale bar = 0.8 mm.

Fig. 46.—Maps of central Mexico, showing geographical distributions of some species of the subgenus Dyschromus belonging to the dimidiata subgroup.

tions as E. puertogallo, but on the next mountain range to bles that of E. chrysophana, except that the preapex is uni- the east. formly thick throughout, and the apical margin is subtruncate, as in E. puertogallo.

28. Euchroa (Dyschromus) filodecaballo, new species Description.—Body Size. Individuals moderate in size. Apparent body Figs. 45a–c, 46a length 11.0M–12.5F mm. Length (n=4M,4F): head 0.79–0.99F, pronotum 2.26M–2.70F, elytra 5.55F–6.70F mm; width: head 2.00M–2.44F, prono- Type Material.—Eight specimens, labeled as follows. HOLOTYPE tum 3.00M–3.91F, elytra 3.70M–4.65F mm. male, “MEXICO Guerrero/ 39.4 km nw Filo de/ Caballo; oak-alder-/ pine; Body Proportions. Head 2.35M–2.60F× wider than long. Pronotum litter; 1730m/ Aug. 10, 1992[,] 40-92”; “MEXICAN EXP. 1992/ G.E. Ball 1.32–1.45F× wider than long. Elytra in males 1.51–1.58 and females and/ H.E. Frania colls.”; “KEPT ALIVE FOR REARING/ M31, died on or 1.42–1.47× longer than wide. near/ 05.iv.1993,” (USNM). PARATYPES, same labels as holotype except Color. Dorsum brassy, with greenish tint that is most evident on head, for longevity and egg laying data, 1F (UASM); the same except, “12.6 km least on elytra (as in Fig. 3e). nw Filo de/ Caballo; oak-pine-/ fir-alder; 2550 m./ Aug. 11, 1992, 46-92,” Microsculpture. Dorsum of head smooth in spots, sculpticells effaced. no rearing data, 1M (UASM). “MEXICO, GUERRERO/ 138.1 km ne Elytra with most sculpticells on disc isodiametric, some slightly trans- Atoyac/ de Alvarez [same locality as holotype], oak-/ alder, leaf litter,/ verse, finely impressed, flat; in most individuals portions of interval 9 and ravine, 1737 m./ 18.VIII.1986[,] 86-76”; “MEXICO EXPED. 1986/ G.E. apex of elytra with surface beaded (sculpticells convex), a few individuals Ball,/ H.E. Frania, &/ D.S. Mulyk colls.,” 1M, 1F (UASM). “MEXICO: with all sculpticells at sides and apex flat. Guerrero, 15.0 km./ S.W. Filo de Caballo, 2500 m/ 92-011, 16.VII.1992, Head. Postocular transverse groove obsolete dorsomedially R.S./ Anderson, oak forest (wet)/ litter[,] berlese,” 1M (UASM). “MEXI- or nearly so. Submentum slightly prominent, posterior face sloped CO Guerrero/ 10.3 km sw Filo de/ Caballo[,] oak-pine-fir/ 2774 m.[,] evenly to transgular groove; latter moderately deep posterad of 8.VII.1987/ R.S. Anderson,” 2F (UASM). submentum. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in Type Locality.—39.4 km northwest of Filo de Caballo, state of Guerrero, males narrowly triangular and females subtriangular, in males 1.63–1.96 Mexico. and females 2.09–2.54× longer than wide. Mentum with paramedian pits moderate in size. Diagnosis.—Of the four species belonging to the sub- Prothorax. Pronotum with sides slightly sinuate posteriorly; hind angles slightly prominent; basolateral impressions moderately long, genus Dyschromus in Mexico that are uniformly brassy in extended to hind margin or nearly so as shallow groove. Prosternal inter- color dorsally, E. filodecaballo has fewer striae interrupt- coxal process with apex indistinctly or not margined. ed on the disc of the elytra than E. santacatarina, while Pterothorax. Elytra with basal punctures; striae 1 to 2, 3, 4, or 5 inter- those same striae are interrupted more times than in E. rupted on disc, stria 1 interrupted 16–30 times, in most individuals 2 interrupted 5–20, 3 interrupted 3–15, 4 interrupted 1–7 times, in a few individ- puertogallo. It differs further from E. santacatarina in that uals 2–4 interrupted less times on one elytron, 5 interrupted 0–3 times; the outer portion of elytral interval 8 is not beaded, the intervals flat on disc, slightly convex posterolaterally; apical declivity sculpticells being flat rather than convex. Unlike in E. with gentle slope. Mesepisternum usually coarsely punctate, nearly chrysophana, the hind angles of the pronotum are slightly impunctate (n=1). prominent. The median lobe of the aedeagus most resem- Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous to coarse; basal sulcus of sternum VII without median fovea.

Fig. 47.—Median lobe of aedeagus of subgenus Dyschromus, B and E, left lateral aspect, all other figures in dorsal aspect; A–C, E. yucuyacua n. sp.; D–F, E. santacatarina n. sp. pdr, preapical dorsolateral ridge; pvb, periostial ventrolateral bulge. Scale bar = 0.8 mm.

Male. Middle tibia with inner face faintly to distinctly expanded near pronotum. The preapex of the median lobe of the aedeagus apex. Hind tibia with most of medial ridge represented by row of promi- most resembles that of E. yucuyacua, but is shorter and the nent subangular tubercles. Aedeagus with intermediate symmetric type 4 median lobe of usual proportions (Fig. 45a), except middle part of medi- middle part of the median lobe distal to the prebasal bend an lobe in profile unusually stout distad prebasal bend (Fig. 45b); preapex is quite stout, as in E. filodecaballo. moderately long, broad, moderately thick and evenly sclerotized through- Description.—Body Size. Individuals small to moderate in size. Apparent out, and with apex subtruncate (Figs. 45a, 45c), preapex with one large M F M F M F centrally located primary haemolymph channel (Fig. 45c); left wall of body length 9.0 –12.0 mm. Length (n=8 ,4 ): head 0.79 –0.95 , pronotum in males 2.12–2.62 and females 2.54–2.74, elytra in males periostial part of median lobe more extensive than, and entirely conceal- M F ing, right wall in lateral view, periostial ventrolateral bulge slightly 5.50–5.90 and females 6.20–6.50 mm; width: head 2.04 –2.36 , prono- prominent in profile (Fig. 45b) and subangulate in cross-section (as in tum in males 3.08–3.41 and females 3.51–3.97, elytra in males 3.70–4.05 Fig. 26c); middle part of median lobe with dorsomedial groove short and females 4.20–4.60 mm. Body Proportions. Head 2.41M–2.59F× wider than long. Pronotum (Fig. 45a). Internal sac with large bulge on right side at middle; apical M M F M band of large microtrichia interrupted dorsally on right side, basal band 1.29 –1.53 × wider than long. Elytra 1.36 –1.49 × longer than wide. not evident. Color. Dorsum brassy with a slight greenish tint (n=7) (Fig. 3e), or head (n=3) and pronotum (n=1) with more pronounced greenish tint. Material Examined.—The type series. Microsculpture. Dorsum of head in some individuals smooth in spots, with sculpticells effaced. Elytra with sculpticells on disc isodiametric to slightly transverse, finely impressed, flat; interval 9 and apex of Collecting Notes.—Euchroa filodecaballo has been col- elytra with surface not beaded, sculpticells flat (n=1), apex of elytra with lected in forest of mesic to temperate aspect consisting var- surface beaded, sculpticells convex (n=4), interval 9 also with surface iously of alder, oak, pine, and fir, at elevations of about beaded (n=2). 1700–2800 m. Head. Postocular transverse groove obsolete dorsomedially or nearly so. Submentum slightly prominent, posterior face sloped evenly to trans- Geographical Distribution.—This species lives in the gular groove; latter moderately deep posterad submentum. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in Sierra Madre del Sur in Guerrero, where we and others males broadly and females narrowly subtriangular, in males 1.51–2.15 and have collected it on the inland side of the Sierra de Atoyac females 2.42–2.61× longer than wide. Mentum with paramedian pits mod- (Fig. 46a). erate in size. Prothorax. Pronotum with sides entire posteriorly and hind angles Phylogenetic Relationships.—The closest relatives of rounded (n=3F) or sides slightly sinuate posteriorly and hind angles obtuse (n=7M,1F); basolateral impressions long. Prosternal intercoxal E. filodecaballo could comprise a group consisting of process with apex not or indistinctly margined (n=10), distinctly mar- E. chrysophana, E. yucuyacua, and E. santacatarina, but gined (n=1). some evidence indicates that it is most closely related to E. Pterothorax. Elytra with basal puncture present on both sides chrysophana (see phylogenetic analyses below for (n=3), one side (n=5), or absent (n=3); striae 1 to 2, 3, 4, or 5 interrupt- details). ed on disc, in most individuals stria 1 interrupted 10–30, 2 interrupted 5–26, 3 interrupted 1–9, 4 and 5 interrupted 0–4 times, in some individuals striae on one elytron interrupted fewer times; intervals flat on Chorological Affinities.—We have collected E. filodeca- disc, slightly convex or flat posterolaterally; apical declivity with gen- ballo at several places along the road between the towns tle slope. Mesepisternum moderately coarsely punctate in males, more of Filo de Caballo and Puerto Gallo, and have also found so in females. E. puertogallo along the same road at a site situated Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous in most specimens, coarse in two females; basal sulcus of sternum VII with- between two of the localities for E. filodecaballo. The near- out median fovea. est site is only eight kilometers away, is at about the same Male. Middle tibia with inner face not or just slightly expanded near elevation, and the forest is of similar aspect, so the two apex. Hind tibia with most of medial ridge represented by row of promi- species probably occur together at some sites. nent subangular tubercles (Fig. 16e). Aedeagus with intermediate asymmetric type 3 median lobe of usual proportions (Fig. 45d), except middle part in profile unusually stout distad prebasal bend (Fig. 45e); preapex 29. Euchroa (Dyschromus) chrysophana Bates moderately long, broadly tapered to a rounded point (Fig. 45d), area to left of midline somewhat thinner and less darkly sclerotized than area to right Figs. 3e, 16e, 42b, 45d–f (Fig. 45f), right side of preapex with flat preapical dorsolateral ridge (Fig. 45d), two primary haemolymph channels, longest with origin at extreme Euchroa chrysophana Bates 1891:249. Type material. LECTOTYPE right side of ostium, broadest with origin to left of midline (Fig. 45f); left female, labeled: “Type/ HT circular, ringed with red]; “Sp fig- wall of periostial part of median lobe more extensive than, and entirely ured”; “Omilteme/ Guerrero/ 8000 ft./ July H.H. Smith”; Tr. Ent. concealing, right wall in lateral view, periostial ventrolateral bulge slight- S.L., 1891/ Euchroa. chrysophana,/ Bates”; “Euchroa/ chrysoph- ly prominent in profile (Fig. 45e) and subangulate in cross-section (as in na/ Bates” [handwritten] (BMNH). Tschitschérine, 1898:60. Fig. 26c); middle part of median lobe with dorsomedial groove short (Fig. Dyschromus chrysophanus (Bates): Darlington, 1935:180. 45d). Internal sac with large bulge on right side at base and on left side at middle; apical band of large microtrichia extending completely around Type Locality.—Omilteme, state of Guerrero, Mexico. sac, basal band not evident. Diagnosis.—Besides E. chrysophana, three other species Remarks.—Females tend to be larger than males, and as of the subgenus Dyschromus in Mexico are uniformly well, compared to males, the pronotum of females tends to brassy in color dorsally. Of these, E. chrysophana differs be broader posteriorly, and the mesepisternum and basal sul- externally from E. santacatarina and E. puertogallo in the cus of abdominal sterna V–VII tends to be more coarsely same ways as does E. filodecaballo. It is distinguished from punctate, although only two females exhibit all four traits. E. filodecaballo by the obtuse to rounded hind angles of the

Material Examined.—Eleven specimens. GUERRERO: Omiltemi, Head. Frontal furrows with deeply and sharply impressed portion 8000 ft., July., H.H. Smith, F holotype, 2M and 1F syntypes, (BMNH); 1M unusually short, not extended posteriorly to level of anterior supraorbital syntype, (UASM); same data as holotype, identified as Euchroa dimidia- setigerous punctures (as in Fig. 8a). Postocular transverse groove shallow- ta Chaud., 1F (BMNH); Omiltemi, 7,300', VII.14–15.1966, Ball- ly to moderately deeply impressed dorsomedially. Submentum slightly Whitehead, 5M (UASM); the same except, 0.9 mi. e. Omiltemi, prominent, posterior face sloped evenly to transgular groove; latter mod- VII.19.1966, 1F (UASM). erately deep posterad submentum. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in Collecting Notes.—Euchroa chrysophana has been found males narrowly triangular and female subtriangular, in males 1.60–1.90 in cloud forest at about 2200 m elevation. and female 2.07× longer than wide. Mentum with paramedian pits small. Prothorax. Pronotum with sides moderately sinuate posteriorly; hind angles prominent; basolateral impressions short. Prosternal intercoxal Geographical Distribution.—This species is known only process not or indistinctly margined (n=1) posteriorly. from the western versant of the Sierra Madre del Sur in Pterothorax. Elytra without basal punctures; striae 1–7 barely or not central Guerrero, west of Chilpancingo (Fig. 42b). evident, 8 more deeply impressed or not, 1–7 interrupted numerous times; intervals flat; apical declivity moderately steep to nearly perpendicular. Phylogenetic Relationships.—Euchroa yucuyacua, either Mesepisternum with few shallow punctures. Abdomen. Sterna V–VII with foveae in basal sulcus coarse; basal sul- by itself or together with E. santacatarina, could be the cus of sternum VII with median fovea. closest relative of E. chrysophana, or perhaps the nearest Male. Middle tibia with inner face not expanded near apex. Hind tibia relative is E. filodecaballo (see phylogenetic analyses with most of medial ridge represented by row of prominent subangular below for details). tubercles. Aedeagus with intermediate asymmetric type 3 median lobe of usual proportions (Figs. 47a–b); preapex long, broadly tapered to a round- Chorological Affinities.—Euchroa filodecaballo, and also ed point, area to left of midline somewhat thinner and less darkly sclerotized than area to right (Fig. 47c), preapical dorsolateral ridge present (Fig. E. puertogallo, both occur in the next range of mountains 47a) and flat in profile (Fig. 47b), two primary haemolymph channels, to the west, at the same elevation as E. chrysophana. longest with origin at extreme right side of ostium, broadest with origin to left of midline (Fig. 47c); left wall of periostial part of median lobe more extensive than and entirely concealing right wall in lateral view, periostial 30. Euchroa (Dyschromus) yucuyacua, new species ventrolateral bulge slightly prominent (Fig. 47b) and subangulate in cross- Figs. 3f, 42b, 47a–c section (as in Fig. 26c); dorsomedial groove extended entire length of middle part of median lobe (Fig. 47a). Internal sac without bulges; apical Type Material.—Five specimens labeled as follows. HOLOTYPE male and basal bands of large microtrichia extended completely around sac. “MEX. Oaxaca/ Cerro Yucuyacua/ 11,175'[,] summit/ (e. Nundaco)/ August 17, 1972”; “B.S. Heming,/ G.E. Ball/ COLLECTORS,” (USNM). Material Examined.—The type series. PARATYPES, same labels as holotype, 1M, 1F (UASM); the same, except “10300–10900',” 2M (UASM). Collecting Notes.—We (Ball) collected the type series of E. yucuyacua in pine forest at elevations of about 3100–3400 Type Locality.—Summit of Cerro Yucuyacua, east of Nundaco, state of m, at and close to the summit of Cerro Yucuyacua. Oaxaca, Mexico. Geographical Distribution.—This species is known only Diagnosis.—No other species of the subgenus Dyschromus from Cerro Yucuyacua, in westernmost Oaxaca (Fig. 42b). has the combination of coppery-colored head and pronotum, and bronze-colored elytra. Also, the surface of the Phylogenetic Relationships.—The closest relative of body has a smoother appearance than in any other member E. yucuyacua is either E. chrysophana or E. santacatarina. of the subgenus because the sculpticells are very finely For details, see phylogenetic analyses, below. impressed, and the elytral striae are barely or not evident. The median lobe of the aedeagus most resembles that of E. Chorological Affinities.—This is the only species of the chrysophana, but is not as stout, and the preapex is longer. subgenus Dyschromus known from Cerro Yucuyacua.

Description.—Body Size. Individuals moderate in size. Apparent body length: 10.0M–11.0F mm. Length (n=4M,1F): head 0.79M–0.91F, prono- 31. Euchroa (Dyschromus) santacatarina, new species tum 2.20M–2.40, elytra 5.35M–5.80F mm; width: head 2.00M– 2.20F, pronotum 3.14M–3.49F, elytra 3.75M–4.20F mm. Figs. 46a, 47d–f Body Proportions. Head 2.40M–2.50F× wider than long. Pronotum 1.36M–1.45F× wider than long. Elytra 1.38F–1.45M× longer than wide. Type Material.—Five specimens, labeled as follows. HOLOTYPE male, Color. Dorsum of head and pronotum coppery with indistinct light “MEXICO. OAXACA./ 5800'. 22.2 mi./ s. Juchatengo./ VII.21– green reflection, elytra dark bronze (Fig. 3f), suture and outer edge green- 22.1966; “George E. Ball/ D.R. Whitehead/ collectors,” (USNM). PARATYPES, “MEX. Oaxaca/ 6.6 mi. e. Sta./ Catarina Juquila/ pine-oak ish (n=1), head and pronotum with green reflection more evident (n=1), M F head and pronotum light metallic green with coppery reflection (n=1), the forest/ July 14, 1972”; “P.A. Meyer, G.E. Ball COLLECTORS,” 1 , 1 (UASM); the same except, “7.3 mi. e. Santa/ Catarina Juquila/ July 12,” same, except copper reflection not evident without magnification (n=1), F head and pronotum light metallic green with coppery reflection, elytra 2 (UASM). with bluish green reflection (n=1). Microsculpture. Dorsum of head and pronotum smooth in spots, Type Locality.—Ca. 35.5 km south of Juchatengo, state of Oaxaca, sculpticells effaced. Elytra with sculpticells on disc isodiametric to slight- Mexico. ly transverse, very finely impressed, flat; interval 9 with sculpticells only slightly more deeply impressed than on disc, small portions of apex of ely- Diagnosis.—Euchroa santacatarina differs from the other tra with surface slightly beaded (sculpticells somewhat convex). three species of the subgenus Dyschromus in Mexico that

Fig. 48.—Median lobe of aedeagus of subgenus Dyschromus, B and E, left lateral aspect, all other figures in dorsal aspect; A–C, E. zempoaltepetl n. sp.; D–F, E. carbonera n. sp. Scale bar = 0.8 mm.

are uniformly brassy in color dorsally in that stria 6, or 6 E. santacatarina could be E. yucuyacua. Another possibil- and 7, as well as 1–5, are interrupted in one or more places ity is that it is most closely related to E. miahuatlan (see on the disc of the elytra, and the entire surface of elytral phylogenetic analyses below for details). interval 9 and the outer part of 8 is distinctly beaded, the sculpticells being markedly convex. The median lobe of the Chorological Affinities.—Euchroa juchatengo (soladeve- aedeagus most resembles that of E. suchixtepec and E. car- ga subgroup) is known to occur at some of the same sites bonera, but the preapical dorsolateral ridge is flat and the as E. santacatarina (see description of that species). periostial ventrolateral bulge is inconspicuous.

Description.—Body Size. Individuals moderate in size. Apparent body 32. Euchroa (Dyschromus) zempoaltepetl, new species length 10.0M–11.0F mm. Length (n=2M,3F): head 0.83F–0.95F, prono- Figs. 13b, 46b, 48a–c tum 2.34F–2.58F, elytra 5.60–6.20F mm; width: head 2.02F– 2.30F, pronotum 3.22M–3.55F, elytra 3.80M–4.35F mm. Type Material.—Six specimens labeled as follows. HOLOTYPE male, Body proportions. Head 2.41M–2.45F× wider than long. Pronotum “MEX. OAXACA 10.5 km. s./ San Pedro y San Pablo/ Ayutla[,] Rte. 179/ 1.28M–1.38F× wider than long. Elytra 1.43F–1.52M× longer than wide. oak-pine forest (dry)/ in litter[,] 1870 m./ June 16, 1979 79-40”; “MEX- Color. Dorsum uniformly brassy, with pronounced greenish tint, espe- ICAN EXP. 1979/ J.S. Ashe, G.E. Ball,/ & D. Shpeley/ collectors,” cially on head and pronotum (as in Fig. 3e). (USNM). PARATYPES, same labels as holotype, 1M (UASM); the same Microsculpture. Dorsum of head in some individuals smooth in spots, except, “...14.2 km. s./ San Pedro y San Pablo/ Ayutla, 1720m June 18, sculpticells effaced. Elytra with sculpticells on disc isodiametric, finely 1979 79-41,” 1M (UASM); the same except, “Cerro Zempoaltepetl[,] impressed, flat; outer half of interval 8, all of 9, and apex of elytra with 8500'[,] (n[ea]r. Tlahuitol-/ tepec)[,] pine-oak for[est]. & vic[inity], Aug. surface distinctly beaded, sculpticells markedly convex. 19, 1972”; “B.S. Heming,/ G.E. Ball/ COLLECTORS,” 1M, 2F (UASM). Head. Postocular transverse groove obsolete dorsomedially or nearly so. Submentum slightly prominent, posterior face sloped evenly to trans- Type Locality.—10.5 km south of San Pedro y San Pablo Ayutla, state of gular groove; latter moderately deep posterad submentum. Oaxaca, Mexico. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in males narrowly triangular and females subtriangular, in males 1.49–1.53 Diagnosis.—Six other species of the subgenus and females 2.15–2.26× longer than wide. Mentum with paramedian pits Dyschromus from Mexico besides E. zempoaltepetl have a moderate in size. Prothorax. Pronotum with sides slightly sinuate (n=4) or entire pos- metallic green head and pronotum, and brassy-colored ely- teriorly (n=1); hind angles obtuse; basolateral impressions short. tra. Of these, E. zempoaltepetl is the most nondescript. It is Prosternal intercoxal process not (n=3) or indistinctly margined (n=2) separable from most specimens of E. dimidiata by the rel- posteriorly. atively gentle slope of the apical declivity of the elytra, and Pterothorax. Elytra with basal punctures; striae 1 to 6 or 7 interrupted on disc, stria 1 interrupted 12–24, 2 interrupted 11–15, 3–5 each interrupt- is further distinguished from E. dimidiata, and also E. ixta- ed 4–14, 6 interrupted 0 (one side only) –8, and 7 interrupted 0–4 times, pa, by the elytral striae, which are not so interrupted on the middle section of stria 8 nearly obliterated by microsculpture; intervals disc, from E. tenancingo by the basal sulcus of abdominal flat; apical declivity moderately steep. Mesepisternum with few to many sterna V–VI, which does not extend to the lateral margins punctures, these fine or coarse. of the abdomen, from E. carbonera by the less prominent Abdomen. Sterna V–VII with foveae in basal sulcus inconspicuous (n=4) or coarse (n=1); basal sulcus of sternum VII without median fovea. hind angles of the pronotum, from E. miahuatlan by the Male. Middle tibia with inner face distinctly expanded near apex. sides of the pronotum, which are sinuate posteriorly, and Hind tibia with most of medial ridge represented by row of prominent sub- from E. flohri by the presence of patches of beaded angular tubercles. Aedeagus with extreme asymmetric type 3 median lobe microsculpture along the entire length of elytral interval 9, of usual proportions (Figs. 47d–e); preapex long, narrow, developed only to right of midline of median lobe except at very base (Fig. 47d), stout and and by the prosternal intercoxal process, which is at least darkly sclerotized throughout (Fig. 47f), preapical dorsolateral ridge flat margined laterally. The median lobe of the aedeagus is in profile (Fig. 47e), one primary haemolymph channel, with origin at nearly identical in appearance to that of E. carbonera (see extreme right side of ostium (Fig. 47f); right wall of periostial part of description of that species). median lobe more extensive than left wall, ostium visible in left lateral view, periostial ventrolateral bulge barely evident in profile (Fig. 47e) and Description.—Body Size. Individuals moderate in size. Apparent body subangulate in cross-section (as in Fig. 26c); dorsomedial groove extend- length 10.0M–12.0F mm. Length (n=4M,2F): head 0.79M–0.97M, prono- ed entire length of middle part of median lobe (Fig. 47d). Internal sac tum 2.36M–2.78F, elytra 5.60M–6.50 mm; width: head 1.99M–2.42M, without bulges; apical and basal band of large microtrichia both extended pronotum 3.20M–3.89M, elytra 3.65M–4.50F mm. entirely around sac. Body Proportions. Head 2.44M–2.56M× wider than long. Pronotum 1.35M–1.44M× wider than long. Elytra in males 1.48–1.53 and females Material Examined.—The type series. 1.43–1.44× longer than wide. Color. Dorsum of head and pronotum metallic green, with or without Collecting Notes.—Euchroa santacatarina has been found bluish reflection; elytra brassy, with or without greenish tint. in forest of tropical aspect at about 1800 m elevation (see Microsculpture. Dorsum without smooth spots, sculpticells evident collecting notes for E. juchatengo for more details). over entire surface. Elytra with sculpticells on disc isodiametric to slightly transverse, moderately deeply impressed, flat; portions of interval 9 and apex of elytra with surface beaded (sculpticells convex). Geographical Distribution.—This species is known from Head. Postocular transverse groove shallow but distinct dorsomedial- a small area on the seaward side of the western part of the ly. Submentum prominent, posterior face sloped evenly to transgular Sierra de Miahuatlan, Oaxaca (Fig. 46a). groove; latter deep posterad submentum. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in Phylogenetic Relationships.—The closest relative of males broadly and females narrowly triangular, in males 1.28–1.70 and

Fig. 49.—Median lobe of aedeagus of E. (Dyschromus) miahuautlan n. sp.: A and C, dorsal aspect; B, left lateral aspect. bu, bulla. Scale bar = 0.8 mm.

females 1.87–1.93× longer than wide. Mentum with paramedian pits Geographical Distribution.—This species is known only small. from the Sierra de Zempoaltepetl in eastern Oaxaca (Fig. Prothorax. Pronotum with sides slightly sinuate posteriorly; hind angles obtuse to slightly prominent; basolateral impressions long or mod- 46b). erately so (Fig. 13b). Prosternal intercoxal process with apical third margined all around or just laterally. Phylogenetic Relationships.—All indications are that Pterothorax. Elytra with basal punctures; all striae entire on disc E. zempoaltepetl occupies the basal position in a lineage (n=3), or stria 1, 1–2, or 1–5 interrupted on disc (n=3), stria 1 interrupted that also contains E. carbonera, E. miahuatlan, E. suchix- 1–7, 2–3 interrupted 1–3 times, 4–5 interrupted once; intervals slightly tepec, and possibly E. santacatarina (see phylogenetic convex; apical declivity moderately steep. Mesepisternum with few shallow punctures. analyses below for details). Abdomen. Sterna V–VII with foveae in basal sulcus coarse; basal sulcus of sternum VII with median fovea. Chorological Affinities.—This is the only species of the Male. Middle tibia with inner face distinctly expanded near apex. subgenus Dyschromus known from eastern Oaxaca, south Hind tibia with most of medial ridge represented by row of prominent sub- of the Rio Santo Domingo, except for E. dimidiata. Its angular tubercles. Aedeagus with intermediate asymmetric type 2 median lobe of usual proportions (Figs. 48a–b); preapex long, broadly tapered to nearest relatives are in western Oaxaca. apex, area to left of midline very thin compared to area to left and colorless (Fig. 48c), preapical dorsolateral ridge markedly convex in profile (Figs. 48a–b), one primary haemolymph channel with origin at extreme 33. Euchroa (Dyschromus) carbonera, new species right side of ostium (Fig. 48c); right wall of periostial part of median lobe Figs. 14a, 46b, 48d–f more extensive than left wall so ostium visible in left lateral view, periostial ventrolateral bulge very prominent in profile (Fig. 48b) and rounded Type Material.—Fourteen specimens labeled as follows. HOLOTYPE in cross-section (as in Fig. 26b); dorsomedial groove extended entire male, “MEX. OAXACA 7.9 km./ n.w. La Carbonera/ Rte. 190[,] oak- length of middle part of median lobe (Fig. 48a). Internal sac with apical pine/ for[est]. (dry)[,] 2250 m./ in litter[,] 79-29/ June 11, 1979”; “MEX- and basal band of large microtrichia both extended completely around sac. ICAN EXP. 1979/ J.S. Ashe, G.E. Ball,/ & D. Shpeley/ collectors,” (USNM). PARATYPES, same labels as holotype, 1M, 2F (UASM); the Material Examined.—The type series. same except “...15.7 km s./ Rte. 190[,] rd. to Ojo de/ Agua[,] oak-pine zone/ Alnus near stream/ in litter[,] 2320 m./ June 12, 1979,” 3M, 2F Collecting Notes.—Euchroa zempoaltepetl has been col- (UASM). “MEX: Oaxaca[,] Hwy./ 190[,] 6.4 km. S. La/ Carbonera[,] 11 lected in dry oak-pine forest at elevations of about Aug[.]/ 1988[,] el. 2450 m./ pine-oak for[est]. litter”; “J.K. Liebherr/ & D.A. Yager,” 3M (CUIC). “MEX. Oaxaca/ Rte. 190, 33.0 mi/ n.w. 1700–2600 m. Oaxaca/ oak forest/ Sept. 4-5, 1967”; “Ball, T.L. Erwin/ R.E. Leech COLLECTORS,” 1M, 1F (UASM).

Type Locality.—7.9 km. northwest of La Carbonera, state of Oaxaca, Prothorax. Pronotum with sides moderately to markedly sinuate pos- Mexico. teriorly; hind angles prominent; basolateral impressions moderately long to long. Prosternal intercoxal process with apical third margined all Diagnosis.—Among species of the subgenus Dyschromus around or just laterally. with metallic green head and pronotum and brassy-colored Pterothorax. Elytra without basal punctures; striae 1–5 or 1–6 interrupted on disc, stria 1 interrupted 4–6 (n=1) or 6–22 times (n=10), 2 inter- elytra, E. carbonera is recognizable by its small body size rupted 0–12, 3–5 interrupted 0–6, 6 interrupted 0–3 times; intervals slight- and prominent hind angles of the pronotum. We have seen ly convex (n=4) to nearly flat (n=1); apical declivity moderately steep to only a few individuals of E. dimidiata and E. zempoalteptl nearly perpendicular. that are as small. The median lobe of the aedeagus differs Abdomen. Sterna V–VII with foveae in basal sulcus coarse; basal sulcus of sternum VII without median fovea (n=2) or with small median only slightly from that of E. zempoaltepetl. fovea (n=3). Male. Middle tibia with inner face distinctly expanded near apex. Description.—Body Size. Individuals relatively small. Apparent body M F M Hind tibia with most of medial ridge represented by row of prominent sub- length 9.0–10.5 mm. Length (n=6 ,5 ): head 0.81(0.73 –0.85), prono- angular tubercles. Median lobe of aedeagus as in E. zempoaltepetl, except tum 2.29(2.04M–2.54F), elytra 5.25(4.7M–5.7) mm; width: head M F somewhat broader in dorsal aspect (Fig. 48d), preapex with area to left of 2.01(1.81 –2.12 ), pronotum in males 3.06(2.80–3.20) and females midline more extensive (Figs. 48d, 48f), and middle part of median lobe 3.26(3.14–3.33), elytra in males 3.45(2.80–3.70) and females shorter distad of prebasal bend (Figs. 48d–e). 3.85(3.65–4.05) mm. M M Body Proportions. Head 2.47(2.39 –2.57 )× wider than long. Material Examined.—The type series. Pronotum 1.39(1.29M–1.43M)× wider than long. Elytra in males 1.46(1.41–1.49) and females 1.40(1.39–1.42)× longer than wide. Color. Dorsum of head and pronotum metallic green, elytra brassy and Collecting Notes.—Euchroa carbonera has been collected with or without greenish hue. in dry oak-pine forest at elevations of about 2200–2300 m. Microsculpture. Dorsum without smooth spots, sculpticells evident over entire surface. Elytra with sculpticells on disc isodiametric to slight- Geographical Distribution.—This species is known only ly transverse, finely impressed, flat; portions of apex of elytra beaded, from the northernmost part of the Sierra de Quatro Venados sculpticells convex. Head. Postocular transverse groove shallow but distinct dorsomedial- in western Oaxaca (Fig. 46b). ly. Submentum moderately prominent, posterior face sloping evenly to transgular groove; latter moderately deep posterad submentum. Phylogenetic Relationships.—Euchroa carbonera occu- Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in pies the second most basal position in a lineage that also males broadly and females narrowly triangular, in males 1.31(1.10–1.44) contains E. miahuatlan, E. suchixtepec, and possibly E. and females 1.92(1.73–2.00)× longer than wide. Mentum with paramedi- santacatarina, with E. zempoaltepetl occupying the basal an pits small. position (see phylogenetic analyses below for details). Chorological Affinities.—This species has been found together with E. dimidiata at several localities. Its nearest relatives, E. miahuatlan and E. suchixtepec, occur further south, in the Sierra de Miahuatlan.

34. Euchroa (Dyschromus) miahuatlan, new species Figs. 13d, 46b, 49a–c

Type Material.—Nine specimens, labeled as follows. HOLOTYPE male, “MEX. OAXACA 27.2 km./ s. Miahuatlan[,] oak-/ pine zone; Alnus/ litter;/ 2440 m./ June 14, 1979[,] 79-34”; “MEXICAN EXP. 1979/ J.S. Ashe, G.E. Ball,/ & D. Shpeley/ collectors,” (USNM). PARATYPES, same labels as holotype, 3M, 2F (UASM); the same except, “April 30, 1977”; “MEXICO EXPED. 1977/ J.S. Ashe/ H.E. Frania/ D. Shpeley coll.,” 1M, 1F (UASM). “MEXICO OAXACA Rte[.] 175/ 3.2 km s. San Jose del/ Pacifico; 2440 m/ pine-oak; under logs/ leaf litter/ July 22, l992[,] 26-92”; “MEXICAN EXP. 1992/ J.S. Ashe,/ H.E. Frania,/ D. Shpeley colls.,” 1M (UASM).

Type Locality.—27.2 km south of Miahuatlan, state of Oaxaca, Mexico.

Diagnosis.—The sides of the pronotum of E. miahuatlan are oblique posteriorly and the hind angles are obtuse. These traits in combination distinguish E. miahuatlan from the six other species of the subgenus Dyschromus in Mexico that have a metallic green head and pronotum, and brassy colored elytra. The median lobe of the aedea- Fig. 50.—Median lobe of aedeagus of E. (Dyschromus) suchixtepec n. gus of E. miahuatlan is nearly identical in appearance to sp.: dorsal aspect. Scale bar = 0.8 mm. that of E. suchixtepec; in particular, there is a ventrally-

directed bulla on the left side of the median lobe, at the Chorological Affinities.—Euchroa suchixtepec occurs in base of the prepex (Fig. 49b), a feature unique to these the same part of the Sierra Miahuatlan as E. miahuatlan, two species. but the latter lives further inland and at higher elevations, in forest of more temperate aspect. Description.—Body Size. Apparent body length 10.0M–11.5F mm. Length (n=6M, 3F): head 0.85M–0.95F, pronotum 2.42M–2.74F, elytra M F M F M F 5.55 –6.20 mm; width: head 2.06 –2.36 , pronotum 3.20 –3.61 , ely- 35. Euchroa (Dyschromus) suchixtepec, new species tra 3.70M–4.40F mm. Body proportions. Head 2.39M–2.55F× wider than long. Pronotum Figs. 3b, 46a, 50a–b 1.30F–1.39F× wider than long. Elytra 1.38F–1.51M× longer than wide. Color. Head and pronotum metallic green, elytra brassy with greenish Type Material.—Four specimens labeled as follows. HOLOTYPE male, tint. “MEX. Oaxaca/ Rte. 175, 26.1/ mi. s. Suchix-/ tepec/ July 8–9, '[19]72”; Microsculpture. Dorsum without smooth spots, sculpticells evident “P.A. Meyer,/ G.E. Ball/ COLLECTORS,” (USNM). PARATYPES, same M F over entire surface. Elytra with sculpticells on disc isodiametric to slight- labels as holotype, 1 , 1 (UASM). “MEXICO Oaxaca/ 22.4 km. s. ly transverse, moderately deeply impressed, flat; parts of extreme outer Suchix-/ tepec, Rte. 175/ oak-pine forest/ ca. 1700 m./ April 30, 1977”; M portion of interval 8, nearly all of interval 9, and apex of elytra distinctly “MEXICAN EXP. 1977/ J.S. Ashe,/ H.E. Frania/ D. Shpeley coll.,” 1 beaded, sculpticells markedly convex. (UASM). Head. Postocular transverse groove shallow but distinct dorsomedially. Submentum moderately prominent, posterior face sloped evenly to Type Locality.—About 42 km south of Suchixtepec, state of Oaxaca, transgular groove; latter moderately deep posterad submentum. Mexico. Mouthparts. Dorsal mandibular grooves long. Labial palpomere 3 in males broadly triangular and females subtriangular, in males 1.39–1.64 and Diagnosis.—Euchroa atoyac and E. jalisco are the only females 1.93–2.24× longer than wide. Mentum with paramedian pits small. species of the subgenus Dyschromus besides E. suchixte- Prothorax. Pronotum with sides entire posteriorly; hind angles obtuse pec, that are entirely metallic green in color dorsally. or rounded; basolateral impressions moderately long (Fig. 13d). Prosternal intercoxal process with apical third deeply margined all around, or just However, the sides of the elytra of some specimens of shallowly margined laterally. E. suchixtepec have such a pronounced reddish purple or Pterothorax. Elytra without basal punctures; striae moderately coppery reflection, that at first glance these individuals impressed, stria 1 entire on disc, 2 and 4 interrupted once on one elytron appear to be bicolored. Recognition is never difficult (n=1), stria 1 interrupted once, other striae entire (n=2), stria 1 inter- because the first four to six elytral striae are not as deeply rupted 1–14, 2 interrupted 0–8, 3–5 interrupted 0–3, 6 interrupted 0–1 times (n=5); intervals slightly convex to nearly flat; apical declivity impressed as the others, and elytral intervals 1–7 are gentle. smooth on the disc; among species of Dyschromus, both Abdomen. Sterna V–VII with foveae in basal sulcus moderately coarse traits are unique to E. suchixtepec. This species differs fur- or relatively inconspicuous; basal sulcus of sternum VII without median ther from E. atoyac and E. jalisco in that elytral stria 1–3 fovea. Male. Middle tibia with inner face distinctly expanded near apex. are interrupted numerous times on the disc. The median Hind tibia with most of medial ridge represented by row of prominent sub- lobe of the aedeagus of E. suchixtepec is virtually identical angular tubercles. Aedeagus with extreme asymmetric type 2 median lobe in appearance to that of E. miahuatlan. of usual proportions (Figs. 49a, 49c); preapex long, narrow, developed only to right of midline of median lobe and stout and darkly sclerotized Description.—Body Size. Individuals moderate in size. Apparent body throughout, except at very base (Figs. 49a, 49c), preapical dorsolateral length 11.0M–12.5M mm. Length (n=3M, 1F): head 0.91F–1.07M, prono- ridge present (Fig. 49a), moderately convex in profile (Fig. 49b), one pri- tum 2.52F–2.90M, elytra 6.00M–6.95M mm; width: head 2.20M–2.50M, mary haemolymph channel present, with origin at extreme right side of pronotum 3.43M–3.93M, elytra 4.00M–4.45M mm. ostium (Fig. 49c); left margin of median lobe with ventrally-directed bulla Body Proportions. Head 2.29M–2.46F× wider than long. Pronotum at base of preapex (Fig. 49b); right wall of periostial part of median lobe 1.28M–1.39F× wider than long. Elytra 1.50M–1.56M× longer than wide. more extensive than left wall, ostium visible in left lateral view, periostial Color. Dorsum dark metallic green, head and margins of pronotum ventrolateral bulge very prominent in profile (Fig. 49a) and rounded in with reddish purple reflection (2M), elytra with faint reddish purple reflec- cross-section (as in Fig. 26b); dorsomedial groove extended entire length tion (1F), elytra with pronounced coppery reflection (1M, Fig. 3b). of middle part of median lobe (Fig. 49a). Internal sac with apical and basal Microsculpture. Dorsum of head and pronotum mostly smooth or band of large microtrichia both extended completely around sac. sculpticells very finely impressed, pronotum also with network of wavy, more deeply impressed microlines. Elytral intervals 1–7 largely smooth, Material Examined.—The type series. interval 8 with sculpticells finely impressed, flat; interval 9 and apex of elytra with sculpticells more deeply impressed, portions with surface Collecting Notes.—We have found specimens of E. miahu- beaded, sculpticells convex. atlan in mesic oak-pine forest at about 2400 m elevation. Head. Postocular transverse groove shallow but distinct dorsomedially. Dorsal mandibular grooves long. Submentum moderately prominent, posterior face sloping evenly to transgular groove; latter moderately deep Geographical Distribution.—Euchroa miahuatlan is posterad submentum. known only from a small area on the inland side of the Mouthparts. Labial palpomere 3 in males broadly triangular and in eastern part of the Sierra de Miahuatlan, Oaxaca (Fig. female subtriangular, in males 1.32–1.45 and female 2.0× longer than 46b). wide. Mentum with paramedian pits small. Prothorax. Pronotum with sides entire or slightly sinuate posteriorly; hind angles obtuse or rounded; basolateral impressions moderately long Phylogenetic Relationships.—The nearest relative of (Fig. 3b). Prosternal intercoxal process with entire apical third shallowly E. miahuatlan is either E. suchixtepec or E. santacatarina. to deeply margined. For details, see phylogenetic analyses, below. Pterothorax. Elytra with basal punctures present on both sides, or one side only; striae 1–4, 1–5, or 1–6 shallowly impressed, other striae mod-

Fig. 51.—Second instar larva of E. (Dyschromus) dimidiata Chaudoir, dorsal aspect. A, head, pronotum and mesonotum; B, abdominal tergum I; C, tergum IX and urogomphi. Scale bar = 0.5 mm.

erately deeply impressed, stria 1 interrupted on disc 16–25 times, 2 inter- Some differences between larvae of Dyschromus and rupted 6–16, 3 interrupted 2–13, 4 interrupted 0–7, 5 interrupted 0–3 those of other pterostichines are with respect to the posi- times; intervals flat; apical declivity with gentle slope. Mesepisternum with many moderately coarse punctures. tion of certain setae and pores, while chaetotaxic differ- Abdomen. Sterna V–VII with foveae in basal sulcus relatively incon- ences between larvae of species of Dyschromus involve spicuous to moderately coarse; entire basal sulcus of sternum VII foveate. only size or presence of various setae. We classify setae Male. Middle tibia with inner face distinctly expanded near apex. according to size as follows: minute setae are not notice- Hind tibia with most of medial ridge represented by row of prominent sub- ably longer than the width of the setal socket (Fig. 52b, angular tubercles. Median lobe of aedeagus as in E. miahuatlan except preapex somewhat more slender (Figs. 50a–b). Internal sac with apical seta FR4); short setae are at least 1.5 times longer than the and basal band of large microtrichia both extended completely around sac. width of the setal socket but not more than about 0.2 times the length of the longest setae on the body (Fig. 53a, seta Material Examined.—The type series. 10); we classify all the rest as long. If a seta is minute, then we only illustrate the socket because the seta itself cannot Collecting Notes.—Euchroa suchixtepec has been collect- be seen except in slide preparations viewed at 250 times ed in oak-pine forest of tropical aspect at about 1700 m ele- magnification or more. Many setae exhibit considerable vation. variation for size, even from one side of the body to the other (Fig. 53a). Geographical Distribution.—This species is only known Two subprimary setae present in second and third instar from a small area on the Pacific side of the eastern part of larvae of some species of Dyschromus have not been the Sierra de Miahuatlan, Oaxaca (Fig. 46a). reported before in the literature on pterostichine larvae (Y. Bionomics.—One male collected on July 8–9, 1972, has a Bousquet, pers. comm.). These are designated here as lightly tanned aedeagus, indicating that when found, this pronotal seta alpha and pronotal, mesonotal and metanotal individual was still somewhat teneral. seta beta. When present, pronotal seta alpha is located just anterad of pore g and so is close to seta 9 (Fig. 51a). Phylogenetic Relationships.—Euchroa miahuatlan, Pronotal, mesonotal, and metanotal seta beta is located either by itself, or together with E. santacatarina, has a ventral to seta 11 and seta 9, respectively, well off the main sister group relationship with E. suchixtepec (see phyloge- part of the sclerite (Figs. 51a, 53c). netic analyses below for details). Larvae of different species and instars of the subgenus Dyschromus also vary with respect to microsculpture, Chorological Affinities.—Euchroa miahuatlan occurs in which is of the type referred to by some carabid workers the same part of the Sierra de Miahuatlan as E. suchixtepec, as pointed or multipointed (e.g. Bousquet and Liebherr but the latter lives at lower elevations in forest of more 1994). We refer to it as spiculate microsculpture, a term in tropical aspect. wide use (Stehr 1991).

LARVAE Description of Larvae of the Subgenus Dyschromus Chaudoir Morphological Traits of Larvae of the Subgenus Dyschromus Chaudoir Diagnosis.—Larvae of the subgenus Dyschromus are separable from those of most other Pterostichini by the short, The structural features that distinguish larvae of the sub- markedly transverse head, absence of a coronal ecdysial genus Dyschromus from larvae of other Pterostichini and line, presence on the nasale of a pair of large denticles that those that vary from species to species are the kind of traits bear seta FR10, and by the large number of primary setae that have been employed by many workers dealing with that are minute, including parietal setae 8 and 14, pronotal larvae of other groups of Carabidae, especially other seta 13, and mesonotal and metanotal setae 1 and 2. Only pterostichines; thus we do not provide a detailed introduc- the last mentioned trait, i.e., the very small size of certain tion to the morphological features of Dyschromus larvae. primary setae, distinguish larvae of Dyschromus from all Most of the traits involve chaetotaxy, and the notation for instars of Abaris Dejean and Orthomus Chaudoir. primary setae and pores developed by Bousquet and Goulet (1984) is quite suitable for our purposes, although for ease Description of First Instar.—Color. Head, tergum IX and urogomphi of reading the text, the name of the seta is often written out yellowish to brownish orange, other sclerites paler; parietale with darker macula mesad of cervical groove; urogomphi darker than tergum IX or in full, e.g., parietal seta 6 instead of PR6. Bousquet (1984, not. 1985, 1989) also named some setae that appear initially in Head. Head short, about 1.6 times wider than long (as in Fig. 51a). the second instar and have fixed positions, referring to Nasale not projected, with pair of large denticles (as in Figs. 51a, 52a–b); these as secondary setae. We call them subprimary setae, seta FR10 located laterally on denticle, seta FR11 mesad denticle. Adnasale with additional seta laterad seta FR8 (as in Fig. 51a); seta FR4 reserving the term secondary setae for setae that do not short or minute (as in Figs. 52a–b). Egg bursters consisting of row of have fixed positions, in conformity with the broader litera- about 20 spinules, these barely extended to level of seta FR2, all spinules ture on immature insects (Stehr 1991; Wheeler 1990). separate. Coronal ecdysial line absent, frontale in broad contact with hind

Fig. 52.—Larvae of subgenus Dyschromus, third instar. A–B, frontale, anterior portion, dorsal aspect; A, E. harrisoni n. sp.; B, E. dimidiata Chaudoir, seta FR4 shown at 1,250 times magnification; C–D, head of E. dimidiata; C, vicinity of eye, dorsal aspect; D, lateral aspect; E–G, parietal region of head around eye, lateral aspect; E, E. harrisoni; F, E. dimidiata; G, E. atoyac n. sp.; H, E. harrisoni, right maxilla, ventral aspect. No setae or sensilla shown except for setae 5, 6. cg, cervical groove; ls, longitudinal sulcus; te, transverse extensions of lateral and medial membranous areas of stipes; vs, ventral sclerite of stipes. Scale bar: A–B, E–G = 0.2 mm; C = 0.3 mm; D = 0.4 mm; H = 0.32 mm.

margin of head (as in Fig. 51a). Parietal region with 5 to 6 stemmata (as contraction of that portion of the parietale region between in Figs. 52d–g); no ocular groove; no longitudinal sulcus; cervical groove the cervical groove and the stemmata (Fig. 52d). This also extended dorsad to seta PA3 (as in Fig. 51a), ventral portion not prolonged anteriorly (as in Fig. 52d); distance between stemmata and cervi- resulted in anterior displacement of setae 6, 7 and 8, and cal groove comparatively short (as in Fig. 52d), seta PA8 level with pos- pore c (Fig. 51a), and possibly also the longitudinal sulcus terior row of stemmata, seta PA7 nearly so (as in Figs. 51a, 52c–d); pore (Figs. 52c–d) if it is homologous with the longitudinal sul- PAb mesad of seta PA4 (as in Fig. 51a); seta PA8 minute (as in Figs. 51a, cus of Poecilus Bonelli (Bousquet 1989, fig. 10). The 52c), seta PA6 long or minute, seta PA14 minute (as in Fig. 52d). region of the head posterad the frontale also shortened, Antennal proportions typical for a pterostichine; antennomere 2 without setae. Mandibles moderately curved; retinaculum wide; medial edge of resulting in loss of the coronal ecdysial line (Fig. 51a). terebra smooth. Maxillary galea with seta 7 about as long as segment 2. The only other pterostichine larvae known to us that Lacinia with apex acuminate, not prolonged (as in Fig. 52h); seta MX6 lack the coronal ecdysial suture are the first and second about as long as stipital seta MX5. Stipes with setal group gMX with instars of Abaris and Orthomus; in the third instar this about 20 setae, lateral and medial membranous areas not extended onto ventral sclerite. Prementum with ligula short. suture is present, but very short (Bousquet and Liebherr Thorax. Pronotum with setae 6, 9 and 11 long (as in Fig. 51a), setae 1994). These authors also indicated that in second and 3 and 12 long or minute, seta 10 short or minute, other setae minute; third instars of Abaris and Orthomus the lateral membra- mesonotum and metanotum with setae 9, 11 and 12 long (as in Fig. 51a), nous area of the stipes narrowly extends across the entire seta 13 long or minute, other setae minute. Mesonotum and metanotum ventral surface of the stipes. They then proposed that both anteriorly and laterally with numerous transverse rows of spicules, as many as 3 spicules per row; notal carina extended posteriorly to seta 11, traits are derived, and went on to suggest that Abaris is sharp for entire distance (as in Fig. 53b). closely related to Orthomus. Since all larval instars of the Abdomen. Tergum I with seta 9 long (as in Fig. 51b), seta 7 long, subgenus Dyschromus not only lack the coronal ecdysial short, or minute, seta 10 long or minute, setae 1 and 6 short or minute, suture, but also in the second and third instars the ventral other setae minute. Tergum IX with seta UR2 much longer than UR3 (as in Fig. 51c). Urogomphi with 5 long primary setae (setae 4, 5, 6, 7, 8) (as surface of the stipes is slightly divided by extensions of in Fig. 51c). Terga with numerous transverse rows of spicules, 2–5 both the lateral and medial membranous areas (Fig. 52h), spicules per row, these fine to coarse, tergum I with fewer rows and it follows that Euchroa is related, at least at the subtribal spicules finer; urogomphi with spicules, these more or less uniformly dis- level, to Abaris and Orthomus, as recently proposed by tributed; tergal carina extended posteriorly to seta 7, sharp for entire dis- Will (2000) on the basis of adult characters. Other poten- tance (as in Fig. 51b). tial synapomorphies concern the presence in second and Description of Second and Third Instars.—As in the first instar, except third instar larvae of certain species of Dyschromus of as follows. pronotal seta alpha, and pronotal, mesonotal, and metano- Color. Nota and terga I–VIII tending to be as dark as head and tergum tal seta beta. Both are also present in Abaris, but not in IX and to have some dark maculae, especially third instar. Urogomphi as pale as tergum IX. Pterostichus (Y. Bousquet, pers. com.). Head. Parietale with longitudinal sulcus in form of lobe over eye A derived trait shown by larvae of Dyschromus, but not (Figs. 52c–g). Labium with several secondary setae laterally. Stipes with Abaris is the large number of primary setae that are minute one seta (MXalpha) posterad seta MX2; ventral sclerite of stipes notched in size. Each of these setae (see diagnosis) is large or near- at sides by narrow transverse extensions of lateral and medial membra- ly so in illustrations of larvae of other Pterostichini nous areas (Fig. 52h). Thorax. Nota with numerous subprimary sensilla (Fig. 51a). (Bousquet 1984, 1985, 1989; Bousquet and Liebherr Pronotum with seta alpha short or not evident, seta beta long or not evi- 1994), various Platynini (Liebherr 1991), and a general- dent. Mesonotum with seta 8 short or minute, seta beta long or not evi- ized carabid larva showing the distribution of ancestral dent. Mesonotum and metanotum with fine spicules present anteriorly, setae and pores (Bousquet and Goulet 1984). Among spicules more or less uniformly distributed; notal carina extended posteriorly beyond seta 11 or not. species of the subgenus Dyschromus for which the larva is Abdomen. Terga with seta alpha long or absent; terga I-V with known, those of E. atoyac and E. filodecaballo have the spicules only present laterally, terga V-VIII with 1-3 spicules per row of fewest long primary setae. spicules, these very fine, tergum IX with spicules somewhat coarser; tergal carina extended posteriorly to seta 9, on tergum I sharp to seta 7, becoming sharp posteriorly nearly to seta 9 on successive segments. Larva of Euchroa (Dyschromus) perote Urogomphi with up to 4 subprimary setae (Fig. 51c); spicules confined to basal half of urogomphi or absent. Diagnosis.—Second and third instars of E. perote are dis- Remarks.—Larvae of the subgenus Dyschromus have all tinguishable from E. soladevega and E. dimidiata by hav- of the traits given by Bousquet (1985) for the Pterostichini, ing fewer subprimary setae on the urogromphi, and from and Pterostichus Bonelli in particular, except for those E. harrisoni by mesonotal and metanotal seta beta which noted above. One fundamental difference is the shortness is always long or nearly so. First instars of E. perote are of the head in Dyschromus larvae. From descriptions of separable from E. soladevega and E. dimidiata by the less larvae of other Pterostichini (e.g. Bousquet 1984, 1985, deeply impressed microsculpture on abdominal tergum IX. 1989; Bousquet and Liebherr 1994), and of Platynini (e.g., All instars are readily distinguished from E. atoyac and Liebherr 1991), a group which according to Arndt (1993) E. filodecaballo by the larger number of short to long pri- “may come close to the ground plan of the Harpalinae” to mary setae, including pronotal setae 3 and 12. First instars which the Pterostichini belong, we suggest that the condi- are not separable from E. harrisoni. tion in Dyschromus is derived, and was achieved in part by

Fig. 53.—A, second instar larva of E. (Dyschromus) harrisoni n. sp., prothorax in vicinity of seta 10, dorsal aspect, right and left sides of one individual; B–C, third instar, mesonotum, right lateral aspect; B, E. harrisoni; C, E. dimidiata Chaudoir; D, third instar larva of E. dimidiata, abdominal tergum I, dorsal aspect. nc, notal carina. Scale bar: A = 0.2 mm; B–D = 0.4 mm.

Description of First Instar.—Microsculpture. Terga II–VIII with spicu- Head. Parietale with lobe formed by longitudinal sulcus rounded, not late microsculpture very fine, that on tergum IX as fine as on terga as prominent as in Fig. 52e. II–VIII. Thorax. Pronotum with setae alpha and beta short or minute. Head. Frontale with seta FR4 minute; seta FR5 short (as in Fig. 52b). Mesonotum and metanotum with seta beta long or nearly so (as in Fig. Parietale with 6 stemmata; seta PA6 long (as in Fig. 52d). 53c); notal carina sharp to seta 11, not extended beyond it (as in Fig. 53b). Thorax. Pronotum with setae 3 and 12 long; seta 10 short or minute. Abdomen. Terga I–VIII with seta alpha long (as in Fig. 51b). Mesonotum and metanotum with seta 13 long (as in Fig. 51a); seta 8 usu- Urogomphi with 6 long setae including 1 subprimary seta (epsilon), seta ally minute, rarely short. gamma usually minute or not evident, rarely short, no other subprimary Abdomen. Terga with setae 7 and 10 long; tergum I with setae 1 and setae evident. 6 minute. Description of Third Instar.—As in second instar, except as follows. Description of Second Instar.—As in first instar, except as follows. Head. Frontale with seta FR4 short (as in Fig. 52a). Parietale with lobe Microsculpture. Urogomphi without spiculate microsculpture. formed by longitudinal sulcus moderately prominent (as in Fig. 52e).

Fig. 54.—Map of Mexico, showing numbers of species of the subgenus Dyschromus, except for E. dimidiata Chaudoir, in each area where this subgenus is known to occur. RSD, Rio Santo Domingo.

Thorax. Pronotum with seta 10 short; seta alpha short to long; seta than minute, while third instars are distinguishable from E. beta long or nearly so. Mesonotum and metanotum with seta 8 usually perote by notal seta beta, which is short or minute. First short, rarely minute; seta beta long. Abdomen. Tergum I with setae 1 and 6 usually short, rarely minute instars of E. harrisoni are distinguishable from the other (as in Fig. 53d), becoming minute on successive segments. Urogomphi species in the same ways as E. perote, but are not separa- with two subprimary setae—epsilon and gamma, former long, latter long, ble from E. perote. short or minute. Description of First Instar.—Microsculpture. Terga II–VIII with spicu- Material Examined.—Nine first, ten second, and six third instar larvae, late microsculpture very fine, that on tergum IX as fine as on terga and exuviae of the first two instars (CNCI, USNM), all reared from eggs II–VIII. laid by two adult females from Mexico, Tlaxcala, 6.8 km. n. Tlaxco, 9.vii Head. Frontale with seta FR4 short or minute; seta FR5 short. Parietale and 22.vii.1992 (UASM). Most of the larvae are stored in alcohol; a few with 6 stemmata; seta PA6 long (as in Fig. 52d). are mounted on slides. Thorax. Pronotum with setae 3 and 12 long; seta 10 short or minute. Mesonotum and metanotum with seta 13 long (as in Fig. 51a); seta 8 short or minute. Larva of Euchroa (Dyschromus) harrisoni Abdomen. Terga with setae 7 and 10 long; tergum I with setae 1 and 6 Figs. 52a, 52e, 52h, 53a–b short or minute. Description of Second Instar.—As in first instar except as follows. Diagnosis.—Second and third instars of E. harrisoni are Microsculpture. Urogomphi without spiculate microsculpture. separable from the other species, except for third instars of Head. Frontale with seta FR4 short (Fig. 52a). Parietale with lobe E. perote, by seta FR4 on the frontale which is short rather formed by longitudinal sulcus rounded, moderately prominent (Fig. 52e). Thorax. Pronotum with seta alpha minute; seta beta short or minute.

Mesonotum and metanotum with seta beta short or minute; notal carina ed on slides. sharp to seta 11, not extended beyond it (Fig. 53b). Abdomen. Terga I–VIII with seta alpha long (as in Fig. 51b). Urogomphi with 6 long setae including 1 subprimary seta (epsilon), no Larva of Euchroa (Dyschromus) dimidiata other subprimary setae evident. Figs. 51a–c, 52b–d, 52f, 53c–d Description of Third Instar.—As in second instar except as follows. Thorax. Pronotum with seta alpha short or minute. Diagnosis.—Second and third instars of E. dimidiata are Abdomen. Tergum I with setae 1 and 6 short (as in Fig. 53d), becom- distinguishable from the other species, except for ing minute on successive segments. E. soladevega, by the urogomphi, which have four instead of one or two long subprimary setae, and they are Material Examined.—Twenty-one first, two second, and eight third instar larvae, and exuviae of the first two instars (CNCI, USNM), all separable from E. soladevega by the mesonotal and reared from eggs laid by three adult females from Mexico, Queretaro, 29 metanotal carina, which extends posteriorly to seta 12, km. e. Landa de Matamoros, 24–25.vii.1992 (UASM). Most are stored in and the third instar also by the longer length of setae 1 alcohol; a few are mounted on slides. and 6 on abdominal tergum I. The first instar of E. dimidiata is unique because of the coarseness of the Larva of Euchroa (Dyschromus) soladevega spiculate microsculpture on abdominal terga II–VIII, which is even coarser than on tergum IX. Diagnosis.—Second and third instars of E. soladevega Description of First Instar.—Microsculpture. Terga II–IX with spicules are distinguishable from the other species, except for coarse, more so on terga II–VIII. E. dimidiata, by the urogomphi which have four rather Head. Frontale with seta FR4 minute, seta FR5 short (as in Fig. 52b). than one or two long subprimary setae, and they are sep- Parietale with 6 stemmata; seta PA6 long (as in Fig. 52d). arable from E. dimidiata by the mesonotal and metanotal Thorax. Pronotum with setae 3 and 12 long, seta 10 short or minute. Mesonotum and metanotum with seta 13 long (as in Fig. 51a); seta 8 carina, which extends posteriorly just to seta 11, and minute. the third instar also by the minute size of setae 1 and 6 Abdomen. Terga with setae 7 and 10 long, setae 1 and 6 minute (as in on abdominal tergum I. The first instar larva is Fig. 51b). separable from E. dimidiata, E. perote, and E. harrisoni by the spiculate microsculpture on tergum IX, which is Description of Second Instar.—As in first instar except as follows. Microsculpture. Urogomphi with spicules confined to basal third. coarser than on the other terga, and from E. atoyac and Head. Parietale with lobe formed by longitudinal sulcus sharp-edged E. filodecaballo by the larger number of short to long pri- and almost as prominent as in Fig. 52d. mary setae. Thorax. Pronotum with seta alpha short or minute, seta beta long (as in Fig. 51a). Mesonotum and metanotum with seta 8 short to minute, seta Description of First Instar.—Microsculpture. Terga II–VIII with spicu- beta long; notal carina extended to seta 12, sharp to seta 11. late microsculpture relatively fine, slightly coarser on posterior segments, Abdomen. Terga I–VIII with seta alpha long (Fig. 51b). Urogomphi coarsest on tergum IX. with 9 long setae including 4 subprimary setae (beta, gamma, delta and Head. Frontale with seta FR4 minute, seta FR5 short (as in Fig. 52b). epsilon), no other subprimary setae evident (Fig. 51c). Parietale with 6 stemmata; seta PA6 long (as in Fig. 52d). Thorax. Pronotum with setae 3 and 12 long, seta 10 short or minute. Description of Third Instar.—As in second instar except as follows. Mesonotum and metanotum with seta 13 long (as in Fig. 51a); seta 8 Head. Parietale with lobe formed by longitudinal sulcus sharp- minute. edged and prominent (Figs. 52c–d, 52f). Abdomen. Terga with setae 7 and 10 long, setae 1 and 6 minute. Thorax. Mesonotum and metanotum with seta 8 short to long; notal carina sharp to seta 12 (Fig. 53c). Description of Second Instar.—As in the first instar, except as follows. Abdomen. Tergum I with setae 1 and 6 short (Fig. 53d), becom- Microsculpture. Urogomphi with some spicules at base. ing minute on successive segments. Head. Parietale with lobe formed by longitudinal sulcus rounded, not as prominent as in Fig. 52e. Material Examined.—Forty-six first, four second, and 12 third Thorax. Pronotum with seta alpha short or minute, seta beta long (as instar larvae, and exuviae of the first two instars (CNCI, USNM), all in Fig. 51a). Mesonotum and metanotum with seta 8 short to minute, seta reared from eggs laid by two adult females, one from Mexico, beta long; notal carina sharp to seta 11, not extending beyond it (as in Fig. Oaxaca, 24.2 km. s.w. Yolomecatl, 20.viii.1992, and the other from 53b). 38.4 km. N. Telixlahuaca, 19.vii.1992 (UASM). Most are stored in Abdomen. Terga I–VIII with seta alpha long (as in Fig. 51b). alcohol; a few are mounted on slides. Urogomphi with 9 long setae including 4 subprimary setae (beta, gamma, delta and epsilon), no other subprimary setae evident (as in Fig. 51c). Larva of Euchroa (Dyschromus) atoyac Description of Third Instar.—As in second instar except as follows. Fig. 52g Head. Parietale with lobe formed by longitudinal sulcus sharp-edged and prominent (as in Figs. 52c–d, 52f). Thorax. Mesonotum and metanotum with seta 8 short to long (as in Diagnosis.—Larvae of E. atoyac have more minute Fig. 53c). primary setae than do larvae of the other species except for E. filodecaballo, including parietal seta 6, pronotal setae 3 Material Examined.—Fifteen first, nine second, ten third instar larvae, and 12, mesonotal and metanotal seta 13, and tergal seta one pupa, and exuviae of all three instars (CNCI, USNM), all reared from 10. We are unable to distinguish the first instar larva of eggs laid by two adult females from Mexico, Oaxaca, 14.9 km. n. Sola de Vega, 20.vii.1992 (UASM). Most are stored in alcohol; a few are mount- E. atoyac from that of E. filodecaballo.

TABLE 1. Adult characters of species of the subgenus Dyschromus. For explanation of characters see Appendix 2.

Characters

1 2 3 4 5 6 7 8 9 10 11 12 13 14

E. tiburonica aabcababab baaa E. opaca cabcbcaecb ba aa E. centralis ddaDabaaecbbbaa E. independencia ceacacadcb bbaa E. cupripennis ceacacacbbbbaa E. perezi ceacacabba bbaa E. pedernales beacacabaabbaa E. nitidipennis cgaaabbabaacbb E. sallei agaaabbabaacbb E. huautla ahaaaba?ca acbb E. cuiyachapa cgaDaabbdcaaccb E. zongolica dgaaabbc-aaccb E. lasvigas cgababbcbaaccb E. flohri beababbccaaccb E. teotitlan bgababbcbaaccb E. citlaltepetl bgacabacbaacdb E. perote agacabbcbaacdb E. harrisoni abababbbbaacdb E. soladevega abbcacabbaacca E. juchatengo cbabacaddaacba E. nizavaguiti cbabacaccaacca E. jalisco acaaaaabcabcbb E. tenancingo beaaabacdaabbb E. ixtapa beaaabacdaacbb E. dimidiata aeacabacba acdb E. atoyac bcaaabaddaacbb E. puertogallo bdaaabaddabbbb E. filodecaballo bdaaabaddabbbb E. chrysophana bdaaabaddabbba E. yucuyacua cfaDaabbdcbacbb E. santacatarina cdaaacadcabcba E. zempoaltepetl beababacbaaccb E. carbonera ceaaabacba acbb E. miahuatlan ceababadca acba E. suchixtepec bcaeabacca acba

TABLE 1. CONT.

Characters

15 16 17 18 19 20 21 22 23 24 25 26 27 28

E. tiburonica ababaacbaaaDaaa E. opaca ababaeedacabaa E. centralis ababaeedacaDaba E. independencia ababaeedaaaaaa E. cupripennis ababaeedaaaabb E. perezi abcbaeedaaaabb E. pedernales abbbaeedaaaabc E. nitidipennis bbcaaabbabacab E. sallei bbaaaaaaaaacac E. huautla bbcaaabaaaac?? E. cuiyachapa babaaacbaaacad E. zongolica babaaabbaaacac E. lasvigas baaaaabbacacac E. flohri bbbaaabbaaacad E. teotitlan baaaaabbabacad E. citlaltepetl baaaaabbabacad E. perote bbaaaabbabacad E. harrisoni bbaaaacbacacac E. soladevega abaaaadcaaacad E. juchatengo bbaaaabbaaacac E. nizavaguiti abaaaabbaaacad E. jalisco aacaabccab acad E. tenancingo bacaacdcbabDaad E. ixtapa bacaaedcbaacad E. dimidiata bbbaadecbaacad E. atoyac bbcaabdcaaacad E. puertogallo bacaacecaaacbd E. filodecaballo bacaadecaaacad E. chrysophana aacabdecaa acbd E. yucuyacua bbcaceedbaabbd E. santacatarina abcaaddcaaacad E. zempoaltepetl babaacccaaabad E. carbonera babac ddc aa aba d E. miahuatlan abbaccdcaaacad E. suchixtepec abaabdecaaaDaad

TABLE 1. CONT.

Characters

29 30 31 32 33 34 35 36 37 38 39 40 41 42

E. tiburonica baaaaaaaaaaaba E. opaca baaaaabaaaaaba E. centralis aaaaaaaaaaaabb E. independencia aaaaaabaaaaaab E. cupripennis aabaaabaabaaab E. perezi aacabbbaabaaab E. pedernales abcabbbaabaaab E. nitidipennis abaabcbaad aaaa E. sallei acabbcbaadacCaa E. huautla a????????????? E. cuiyachapa acabbccaadaaCaa E. zongolica acaccccaadacCa?a E. lasvigas acaecccaadacCaa E. flohri acacbccaadacCaa E. teotitlan acacbccaadacCaa E. citlaltepetl acadbccabdacCaa E. perote acaebccabdacCaa E. harrisoni acaebccabdacCaa E. soladevega acaabccaadacCaa E. juchatengo abaaDaaDbaaCaacCaa E. nizavaguiti acaaDaaDbaaCaabCaa E. jalisco acaaDaaDbaaCaaaCaa E. tenancingo acaaDaaDbaaCaabCaa E. ixtapa acaaDaaDbaaCaabCaa E. dimidiata acaaDaaDbaaCaabCaa E. atoyac abaaDaaDbaaCaaaCaa E. puertogallo abaaDaaDbaaCaaaCaa E. filodecaballo abaaDaaDbaaCabbCaa E. chrysophana a b a a b b b a a Ca b b Ca a E. yucuyacua acaabbbaaCaacCaa E. santacatarina acaebbcabCaacCaa E. zempoaltepetl acacbccabdacCaa E. carbonera acacbccabdacCaa E. miahuatlan acaebccbbd acCaa E. suchixtepec acaebccbbd acCaa

Fig. 55.—Phylogenetic relationships of species groups and subgroups of the subgenus Dyschromus, and species belonging to the opaca group and soladevega subgroup, as indicated by mental analysis, and showing distribution of derived character states for selected adult characters. Underlining indicates a reversal to a more plesiomorphic condition. Description of First Instar.—Microsculpture. Terga II–VIII with spic- carina extending nearly to seta 12, sharp to seta 11; metanotal carina ulate microsculpture relatively fine, slightly coarser on posterior seg- sharp to seta 11, not extended beyond it. ments, coarsest on tergum IX. Abdomen. Terga with seta alpha not evident. Urogomphi with 6 Head. Frontale with setae FR4 and FR5 minute. Parietale with 5 long setae including 1 subprimary seta (epsilon), no other subprimary stemmata; seta PA6 minute (as in Fig. 52g). setae evident. Thorax. Pronotum with setae 3, 10 and 12 minute. Mesonotum and metanotum with setae 8 and 13 minute. Description of Third Instar.—As in second instar except as follows. Abdomen. Tergum I with seta 7 short or minute, II–VIII with seta 7 Thorax. Mesonotal and metanotal carina extending to seta 12, sharp minute or not evident, I–VIII with other setae including seta 10 minute or for entire distance (as in Fig. 53d). not evident. Abdomen. Urogomphi with 7 long setae including 2 subprimary setae (gamma and epsilon), no other subprimary setae evident (as in Fig. 51c). Description of Second Instar.—As in first instar except as follows. Microsculpture. Urogomphi without spiculate microsculpture. Material Examined.—Two first and two third instar larvae, and exu- Head. Frontale with seta FR5 short. Parietale with lobe formed by viae of the first two instars (CNCI, USNM), all reared from eggs laid longitudinal sulcus not prominent, rounded (Fig. 52g). by one adult female from Mexico, Guerrero, 71 km. n.e. Atoyac de Thorax. Pronotum with setae alpha and beta not evident. Alvarez, 26.vii.92 (UASM). Most are stored in alcohol; a few are mount- Mesonotum and metanotum with seta beta not evident; mesonotal ed on slides.

Fig. 56.—Phylogenetic relationships of species belonging to the nitidipennis subgroup of the subgenus Dyschromus as indicated by mental analysis, showing distribution of derived character states for selected adult characters (see also Figs. 55, 57–58). Underlining indicates a reversal to a more plesiomorphic condition. L, species living at elevations up to 1400 m; M, species living at at elevations up to 2300 m; H, species living at elevations up to 3000 m. 1, Sierra Madre Oriental. 2, Cofre de Perote and vicinity. 3, Volcan Citlaltepetl and environs. 4, Sierra Madre de Oaxaca, north of Rio Santo Domingo.

Larva of Euchroa (Dyschromus) filodecaballo goal being the recognition of monophyletic groups through the discovery of derived characters shared by all Diagnosis.—We are unable to distinguish the first instar members of each group (synapomorphies). Three different larva of this species (the only larval stage at hand) from that kinds of analyses were done, one employing a qualitative of E. atoyac. Hennigian approach, termed Mental Analysis by Ball and Maddison (1987), in which by visual inspection of the Material Examined.—One first instar larva (CNCI), found dead in the rearing chamber that contained the adult female, which is from the type- data, first a search is made to identify broad groupings locality for this species (UASM). The specimen is stored in alcohol. based on correlations between morphological traits, as well as ecological attributes such as altitudinal range, and also geographical distribution; then intra-group relation- ANALYSIS OF THE PHYLOGENETIC ships are established largely through the interpretation of RELATIONSHIPS AND CLASSIFICATION OF morphoclines, and finally inter-group relations are deter- SPECIES OF THE SUBGENUS DYSCHROMUS mined based upon the characters initially used to identify CHAUDOIR the broader groupings (Ball 1975; Shpeley and Ball 1993). As most species of the subgenus Dyschromus are known The other two kinds of analyses were strictly quantitative, only from adults, we present first a phylogenetic analysis and concerned only the morphological data, one being par- based upon adult characters (Table 1), and then an analy- simony analysis, in which a search is made to find clado- sis based upon larval characters (Table 2). Phylogenetic grams that minimize the total number of character state analyses were done according to Hennigian principles, the changes over the entire tree, irrespective of whether any

particular change from one condition to another is a derivation or a reversal. We also did character compatibility TABLE 2. Larval characters of some species of the analysis, in which the arrangements of the taxa are found subgenus Dyschromus. For explanation of characters see that are supported by the maximum number of uniquely Appendix 3. derived and unreversed character states, homoplasious characters being excluded from consideration. In doing Characters three different kinds of analyses, our intention was to determine what are the robust groupings. Rather than 12 34567 selecting a preferred cladogram, we considered it more important in this first comprehensive study of the sub- E. perote ba a caaa genus Dyschromus, to identify the more plausible major E. harrisoni aa bdaaa alternative arrangements of the taxa that should be consid- E. soladevega ca aaaba ered in future studies. E. dimidiata ca aaacb E. atoyac cb cbbac E. filodecaballo –b ––b–– Character State Data Adults of all 35 species of the subgenus Dyschromus were scored for the condition of 42 qualitative morphological and RATCHET subprograms within the WINCLADA characters, including 19 characters with two states, eight computer program (Nixon 2000). Compatibility analyses with three states, nine with four states, five with five were done using the CLINCH computer program (Fiala states, and one character with eight states (Appendix 2). 1984). All details about options used when running these Larvae of six species were also examined for the condition programs are given in Appendix 4. Both kinds of numeri- of seven morphological characters, two having two states, cal analyses were done once with all ordered characters four with three states, and one with four states (Appendix (i.e. they were additively binary coded), except for adult 3). Characters are referred to in the text by a number, and body color (Appendix 2, character 2), which was treated as character states by a lower case letter (e.g., Appendix 2, an unordered character. The parsimony analysis was then character 2a). For character state trees with two branches, repeated with all of the characters unordered. The parsi- all of the character states on one branch were assigned let- mony analyses were initially done with all of the characters sequentially, e.g., characters 4a, 4b, 4c. The first char- ters weighted equally, but because of the large number of acter state on the second branch was assigned the next let- minimum length trees obtained, the analyses were done a ter in the sequence, but in upper case, and followed by a second time using the successive approximations weight- lower case letter indicating the character immediately ing procedure of Farris. This weighting procedure was ancestral to it, e.g. character 4Da. The remaining character done once as implemented in NONA, the weight of each states on the second branch were assigned a lower case let- character being based on its consistency index, and then ter, e.g., character 4e. Distribution of character states again, this time as implemented in HENNIG86 (Farris among taxa is shown in Table 1. 1988), the character weights being based upon the product Character states were ordered according to degree of of their consistency and retention indices. In all of the similarity between the various states, as implicit in the def- numerical analyses, trees were rooted by including a hypo- inition of a morphocline—an intergrading series of steps thetical outgroup in the data set having just plesiomorphic from one extreme to another (Maslin 1952). Adult charac- character states. Euchroa huautla was left out of the numer- ters were polarized by outgroup comparison with other ical analyses because it is known only from females. euchroines, especially other members of the Euchroa- Parsimony analyses of the additively binary coded data Microcephalus clade, including Lobobrachus and Bothy- set (i.e., with ordered character states) using the RATCHET noproctus, although we also examined Haplobothynus, program in conjunction with NONA yielded 12 trees having Meropalpus, Setalis, and Setalimorphus. For most charac- a length of 291 steps, consistency index (CI) of 0.30, and ters, we accepted the condition in the subgenus Euchroa as retention index of 0.68. The strict consensus tree of these 12 decisive, there being little interspecific variation in that trees was largely resolved (for details see Fig. 59). subgenus for nearly all of the characters. Larval characters Successive approximations weighting by CI, as implement- were polarized by outgroup comparison with carabid lar- ed in NONA, starting with the above 12 trees, after two iter- vae in general, and Abaris in particular. ations yielded one tree with a length of 293 when the character states were reweighted back to one (Fig. 59), while successive approximations weighting by CI X RI, as imple- Analysis of the Adult Data, mented in HENNIG86, starting with these same 12 trees, Procedures and Results after three iterations yielded four trees, one of which had a minimum length of 298 when the character states were The qualitative Hennigian analysis was done as described reweighted back to one (Fig. 60). Using the RATCHET pro- above. Parsimony analyses were done using the NONA gram in conjunction with NONA, and with all character

states unordered, produced 328 trees having a length of 244 nitidipennis group also exhibit this condition (Table 1). steps, CI of 0.36, and RI of 0.65. The strict consensus tree A synapomorphy for the nitidipennis group is the at least of these 328 trees had just six resolved nodes (for details see slight prominence of the submentum, and the at least mod- Fig. 61). Successive approximations weighting as imple- erate depth of the transgular groove posterior to the submen- mented in NONA, starting with these 328 trees, after three tum (Fig. 55, character 13b). Another synapomorphy is that iterations produced one tree having a length of 245 when the the periostial ventrolateral bulge of the median lobe has a character states were reweighted back to one (Fig. 61). more apical position (character 38Ca) than in the opaca Character compatibility analysis revealed that the major- group. A third possible synapomorphy for the nitidipennis ity of the adult characters conflict with one another, there group is presence of a narrow subapical band, and often also being only 20 out of a possible 90 character states in the a narrow basal band of large microtrichia that extend at least largest sets of mutually compatible characters. There were partially around the internal sac of the aedeagus (character 12 such sets; the trees based on them were only partly 41Ca). However, E. nitidipennis has a single broad band resolved, and significant portions of the 12 arrangements extending around the internal sac (character 41a), as do did not reflect any of those obtained by either mental analy- some members of the opaca group (Table 1), and we are sis or parsimony analysis of the adult or larval data, nor did uncertain as to which is the ancestral condition of this trait there appear to be any geographical or ecological correlates (Appendix 2). in support of the compatibility analysis results, and so we do The opaca and nitidipennis groups emerged in some of not consider them further. our parsimony analyses as sister groups (Fig. 60), but in oth- Mental analysis of the ordered characters yielded a tree ers E. tiburonica of the opaca group was positioned by itself 315 steps in length, with CI of 0.27, and RI of 0.64 (Figs. at the very base of the cladogram (Fig. 59) because this 55–58). Although the mental analysis tree is substantially species has so many plesiomorphic traits. None are confined longer than any of the trees obtained using parsimony analy- to it, and so we do not consider this alternative further. sis, we note that in parsimony analyses, very different arrangements were obtained depending on whether the Relationships Within the Opaca Group.—What are the characters were ordered or not, and if they were weighted, relationships of species belonging to the opaca group and and how that weighting was done (Figs. 59–61). Moreover, how should they be classified? The answers to these ques- preliminary parsimony analyses conducted while refining tions depend in large part on how the three uniformly dark- the adult characters showed that addition or deletion or colored species of Dyschromus from Hispaniola are relat- recoding of what appeared to us to be minor or highly ed to one another, and to the remaining four species on that homoplasious characters invariably led to substantial island, all of which have a metallic green head and prono- changes in tree topology. tum and brassy to coppery-colored elytra. We place the We discuss below the phylogenetic relationships and former in the opaca subgroup, and the latter in the classification of the species of Dyschromus largely in terms cupripennis subgroup. of the arrangement obtained by mental analysis of the adult We are reasonably certain that the cupripennis sub- data, and contrast certain of the results with those obtained group is monophyletic, in part because the combination of in our parsimony analyses. metallic green head and pronotum and brassy or coppery colored elytra is undoubtedly a comparatively derived trait Basal Dichotomy.—We consider the subgenus Dyschromus within the genus (Figs. 55–60, character 2e). As well, the to comprise the opaca and nitidipennis groups, the former four species in the subgroup can be arranged according to containing all of the species on the island of Hispaniola, the the form of the distal part of the median lobe of the aedea- latter all of those in Mexico. A synapomorphy for the opaca gus, starting with E. independencia, which has a general- group is the rounded, as opposed to prominent, shoulders of ized asymmetric type of median lobe, and ending with E. the elytra (Fig. 55, character 18b), a condition only perezi and E. pedernales, both of which have a median approached in the nitidipennis subgroup by some specimens lobe of extreme asymmetric type 1, with E. cupripennis of E. lasvigas. Another derived trait that is shared by most exhibiting an intermediate stage (Fig. 55: characters 31b, members of the opaca group is that the frontal furrows on 31c, 34b, 38b). The third reason is that the above morpho- the head are sharply impressed for only a short distance pos- cline in genitalic structure is paralleled by at least one teriorly, if at all (character 10b), but in E. perezi, E. peder- change in external structure: reorganization of the medial nales, and some specimens of E. cupripennis the sharply ridge of the male hind tibia into a row of prominent tuber- impressed portion is as long as in many members of the cles (characters 28b, 28c), although this trend is also evi- nitidipennis group (character 10a), while in one species dent in the nitidipennis group. belonging to the nitidipennis group, E. yucuyacua, the In contrast to the status of the cupripennis subgroup, sharply impressed portion is short. Moreover, the polarity of we are not convinced about monophyly of the opaca sub- this trait is doubtful (Appendix 2). Yet another derived trait group. It did emerge in one of our parsimony analyses shared by all members of the opaca group is that the postoc- (Fig. 61), but the only derived trait shared by all three ular transverse groove is obsolete dorsomedially or nearly species is that the males have just one narrow band of large so (character 11b), but some species belonging to the microtrichia extended around the base of the internal sac

of the aedeagus (character 41b), and we are uncertain exception, which conflicts with the second one (character about the polarity of this character (Appendix 2). 42b), is the slenderness of labial palpomere 3 in males of One alternative is that E. tiburonica, E. opaca, and E. E. opaca and E. centralis (character 8e). A fourth excep- centralis split off in sequence on the phyletic line leading tion, which conflicts with the third one (character 8e) is the to the cupripennis subgroup, so that E. tiburonica has the two rows of medial flexile setae on the hind tibia (charac- basal position within the opaca group, and E. centralis is ter 29b), a condition found only in E. opaca and the sister of the cupripennis subgroup (Figs. 55, 58). E. tiburonica. Evidence for monophyly of the opaca group, exclusive of We regard as unresolved the relationships of the three E. tiburonica, is that the other species are metallic colored uniformly dark-colored species on Hispaniola, since we (characters 2d, 2e), except for E. opaca (character 2a), lack corroborative evidence from the larval stages, nor do labial palpomere 3 is narrower (Fig. 55, characters 8c, 8d, there appear to be geographical or ecological correlates or 8e, 9b or 9c), except for E. perezi (character 8b) and E. that might allow us to choose among the various alterna- pedernales (characters 8b, 9a), the elytral striae are only tives (Figs. 55, 59–61). As for how to classify these three shallow impressed and are interrupted numerous times species, each one is quite distinctive, as shown by the large (Fig. 55, character 20e), the elytral intervals are flat (Fig. number of morphological differences that separate it from 55, characters 21e, 22d), the sculpticells along the sides as its sister taxon in any of the above arrangements well as at the apex of the elytra are distinctly convex (Fig. (Appendix 4). On this basis, it would be reasonable to 55, character 6b), except for E. centralis (character 6a), place each species in a subgroup by itself, but as we show and the preapical part of median lobe has just two rather below, if we were to apply this criterion to the nitidipennis than four primary haemolymph channels (Fig. 55, charac- group, we would have to recognize an inordinate number ter 35b), again except for E. centralis. of monospecific subgroups. We choose to place all three of Evidence for a sister group relationship between the uniformly dark-colored species of the subgenus E. centralis and the cupripennis subgroup is the brassy (as Dyschromus on the island of Hispaniola together in the opposed to black) color of the dorsum of E. centralis opaca subgroup, which, based on most of our evidence is, (character 2d), which is presumably intermediate to the at worst, a paraphyletic group. bicolored condition in the cupripennis subgroup and the all black condition in E. tiburonica. Other synapomorphic Subgroups of the Nitidipennis Group in Mexico.—How traits are the short, straight spermatheca (character 42b), many subgroups of Dyschromus are there in Mexico, and middle tibia of males only slightly expanded or not at all what are their limits? We have recognized three subgroups (character 27b), and the moderate rather than large size of based largely upon body color, degree of interruption of the mental paramedian pits (character 12b), except for the elytral striae, geographical distribution, and to a lesser E. perezi and E. pedernales (character 12a). However, in extent, the condition of the median lobe of the aedeagus certain parsimony analyses E. centralis was paired with (Fig. 58). As constituted, two of the three subgroups are E. opaca (Figs. 59–60) because in both species labial heterogenous for type of median lobe of the aedeagus (see palpomere 3 is especially narrow, particularly in males Appendix 1 for description of each type of median lobe), (characters 8e, 9c), the sculpticells along the sides of the one being the soladevega subgroup, which contains two elytra are elongate (character 5b), and the mesepisternum species with an intermediate or extreme symmetric type 4 is smooth (character 24c); in some of these analyses E. median lobe (Figs. 36d–g), and one species with an inter- independencia was grouped with E. opaca and E. centralis mediate asymmetric type 2 median lobe (Figs. 36a–c). The (Fig. 59) because labial palpomere 3 is also quite narrow dimidiata subgroup is even more heterogenous, as it con- in this species (characters 8d, 9c). In still other parsimony tains four species with an intermediate or extreme asym- analyses, E. opaca was paired with E. tiburonica (Fig. 61) metric type 2 median lobe (Figs. 48, 49, 50), three species because the hind tibia has two rows of flexile setae medi- with an intermediate or extreme asymmetric type 3 medially (character 29b), and the sculpticells on the disc of the an lobe (Figs. 45d–f, 47), and seven species with an inter- elytra are elongate (character 3b). mediate or extreme symmetric type 4 median lobe (Figs. Much of the evidence for each of the above arrange- 26c, 38–40, 43–44, 45a–c). The nitidipennis subgroup is ments of species in the opaca group is equivocal because more homogeneous than the others for aedeagal type the derived condition of many of the characters is also evi- because it contains only species that exhibit either an inter- dent among species of the nitidipennis group from mediate (Figs. 25, 26a, 28, 29a–c, 30, 32–33) or extreme Mexico. One exception pertains to the characters involved asymmetric type 2 median lobe (Figs. 26b, 34–35), or a in the morphocline leading to extreme asymmetric type 1 variant of this type of median lobe (Figs. 29d–f). median lobe (characters 31b, 31c, 34b, 38b), of which all The substantial amount of heterogeneity for structure but character 34b are confined to members of the of the median lobe within subgroups of the nitidipennis cupripennis subgroup. A second exception is the short group contrasts with the situation in the opaca group. straight spermatheca (character 42b), a condition restrict- Moreover, the external characters upon which we base our ed to E. centralis and the cupripennis subgroup. A third recognition of the dimidiata and soladevega subgroups are all homoplasious (see below). In all parsimony analyses,

Fig. 57.—Phylogenetic relationships of species belonging to the dimidiata subgroup of the subgenus Dyschromus as indicated by mental analysis, showing distribution of derived character states for selected adult characters. Underlining indicates a reversal to a more plesiomorphic condition (see also Figs. 55–56, 58).

two clusters of species emerged within the nitidipennis analysis, and which constitutes the basis for our classifica- group (Figs. 59–61), one containing all of the species with tion of species in the nitidipennis group, is discussed in an intermediate or extreme asymmetric type 2 median detail below. Aspects of it are contrasted with the results of lobe, as well as the one species with an extreme asymmet- our parsimony analyses. ric type 3 median lobe, and the other cluster containing some of the species with an intermediate asymmetric type Nitidipennis subgroup.—We have included 11 species in 3 or intermediate symmetric type 4 median lobe, and all of the nitidipennis subgroup, based on five lines of evidence. those with an extreme symmetric type 4 median lobe. 1) Nine of the species have a metallic bluish-purple or cop- Certain species having an intermediate asymmetric type 3 pery colored head and pronotum, and reddish-purple ely- or intermediate symmetric type 4 median lobe were posi- tra, a combination of colors otherwise found among tioned in one cluster or the other, depending on how the euchroines only in Meropalpus, so this is clearly a derived parsimony analysis was done (e.g., E. yucuyacua, cf. Figs. trait (Figs. 55–56, 58, characters 2g, 2h). The other four 60, 61). The arrangement that we obtained by mental lines of evidence also apply to the other two differently

colored species that we include in the subgroup: E. flohri, preapex (character 32b), and at least the base of the medi- which has a metallic green head and pronotum and brassy- al ridge of the male hind tibia is represented by a row of colored elytra (character 2e); and E. harrisoni, which is tubercles (characters 28b, 28c). The second most basal uniformly metallic bluish-purple dorsally (character 2b). position within the nitidipennis subgroup is occupied by E. 2) All but two of the ten species for which the male is sallei because in males of all other species of the subgroup known, can be arranged more or less according to the for which that sex is known, except for E. nitidipennis, the structure of the distal part of the median lobe of the aedea- primary haemolymph channel on the left side of the gus, starting with E. nitidipennis, which has a relatively preapex of the median lobe is lacking (character 35c), and plesiomorphic intermediate asymmetric type 2 median labial palpomere 3 is not so broad (characters 8b, 8c, 8d). lobe (Figs. 25a–c, 26a), and ending with E. perote and E. As well, the submentum is more prominent and the adjoin- harrisoni, both of which have an extreme asymmetric type ing portion of the transgular groove is deeper (character 2 median lobe (Figs. 26b, 34–35), with the transitional 13c), again except for E. nitidipennis and also E. huautla, stages exhibited by E. sallei (Figs. 25d–f), E. cuiyachapa which is only known from two females. (Fig. 28), E. flohri (Fig. 30), E. teotitlan (Fig. 32), and E. We consider E. huautla to be most closely related to E. citaltepetl (Fig. 33). As for the other two species, E. lasvi- sallei because only a few other species of Dyschromus gas, exhibits what appears to us to be a variant of an attain such a large size (Appendix 2, Table 1, character 1a), extreme asymmetric type 2 median lobe (Figs. 29d–f) that and no other members of the subgenus have the elytral is more directly derivable from the intermediate asymmet- intervals so convex posteriorly (character 22a). Moreover, ric type 2 median lobe shown by E. cuiyachapa (Fig. 28), as far as known, they occur in adjoining mountain ranges with E. zongolica exhibiting a transitional condition (Figs. (Sierra Zongolica and Sierra de Huautla, respectively), and 29a–c). 3) The third line of evidence is that the above mor- at lower elevations in forest of more tropical aspect than phocline in male genitalic structure is more or less paral- other species belonging to the nitidipennis subgroup. With leled (see below) by changes in two external morphologi- regards to size of the body, convexity of the elytral inter- cal traits (Fig. 56): sculpticells on disc of elytra becoming vals, and altitudinal distribution, E. sallei and E. huautla more deeply impressed (character 4); and increasing are in fact more primitive than E. nitidipennis, which com- prominence of submentum and depth of adjoining portion pared to those two species, is smaller in size (character of transgular groove (character 13). 4) The fourth line of 1b), has flatter elytral intervals (character 22b), and lives evidence is that the altitudinal distributions of the 11 at higher elevations. Hence these data are in conflict with species, and also the type of forest inhabited by each, both the evidence from males that E. nitidipennis is the most partially reflect the above arrangement (Fig. 56). The primitive member of the nitidipennis subgroup. sequence of species, starting with the one found at the low- In our parsimony analyses, E. nitidipennis and E. sallei est elevations in forest of most tropical aspect, and ending were positioned on the cladograms either near the base of with the one at the highest elevations in forest of most tem- the branch containing all of the species with an intermedi- perate aspect is: E. sallei, E. huautla (male unknown), ate or extreme type 2 median lobe (Fig. 60), or at the apex E. nitidipennis, E. lasvigas, E. cuiyachapa, E. zongolica, of that branch (Figs. 59, 61). We consider that the available E. lasvigas, E. teotitlan, E. harrisoni, E. flohri, E. citlal- data best support placement of E. nitidipennis and E. sallei teptel, and E. perote. This is significant because all mem- (together with E. huautla) at the base of the nitidipennis bers of the subgenus Euchroa live at comparatively low subgroup (cf. Figs. 56, 58, 60), but reserve judgement on elevations in wet tropical forest, indicating that these their exact relationships. aspects of their way of life are plesiomorphic for the The next most derived member of the nitidipennis subgenus. 5) The last line of evidence for grouping the 11 group according to the morphocline leading from an inter- species together is that all of them are not only confined to, mediate to an extreme asymmetric type 2 median lobe is E. but are the only members of, the subgenus Dyschromus in cuiyachapa (Fig. 56), followed by E. flohri and E. teotit- eastern Mexico, south to the Rio Santo Domingo (Fig. 54), lan, both of which exhibit more extensive thinning of the with the possible exception of E. dimidiata, suggesting left margin of the preapex of the median lobe than E. cuiy- that they evolved in that area as a group. We consider that achapa (character 32c), and also have the sculpticells on the above five lines of evidence, taken together, are suffi- the disc of the elytra slightly more deeply impressed (char- cient reason for placing all 11 species in one subgroup. acter 4b) than do E. nitidipennis, E. sallei, E. huautla or E. Within the nitidipennis subgroup, based on the mor- cuiyachapa (character 4a or 4Da). The sequence toward phocline leading from an intermediate to extreme asym- attainment of an extreme asymmetric type 2 median lobe metric type 2 median lobe, E. nitidipennis occupies the continues with E. citlaltepetl since the entire area of the basal position within the subgroup (Fig. 56). A derived preapex to the left of the midline of the median lobe is very trait shared by all other members of the subgroup for thin and somewhat reduced in extent (character 32d), and which the male is known is the long preapex of the medi- the left wall of the periostial part of the median lobe is an lobe (character 30c); also some portion of the left mar- more extensively developed than the right wall (character gin of the preapex is markedly thinner than the rest of the 37b). The sequence ends with E. perote and E. harrisoni,

Fig. 58.—Phylogenetic relationships of species of the subgenus Dyschromus as indicated by mental analysis, showing distribution of selected adult traits. See also Figs. 55–57. A, color of head and pronotum/color of elytra: Bl, black; Br, brassy; Bz, bronze; C, coppery; G, metallic green; P, metallic bluish purple; R, reddish purple. B, interruption of elytral striae, character 20. C, type of median lobe of aedeagus: E1-3, extreme asymmetric types 1, 2, 3. E4, extreme symmetric type 4. G, generalized asymmetric type. i1-3, intermediate asymmetric types 1, 2, 3. i4, intermediate symmetric type 4. csg, dsg, nsg, osg, ssg: cupripennis, dimidiata, nitidipennis, opaca, and soladevega subgroups.

the preapex being developed only to the right of the mid- laltepetl and E. perote (character 4c), but contrary to line of the median lobe (character 32e). expectation, the sculpticells on the disc of the elytra of The morphocline pertaining to the contours of the sub- E. harrisoni are no more deeply impressed than in E. teoti- mentum and adjoining portion of the transgular groove tlan (character 4b). also finishes with E. perote and E. harrisoni, as well as E. Placement of E. lasvigas in the above sequence poses a citlaltepetl, because in all three species the submentum is problem because while the preapex of the median lobe is very prominent and concave posteriorly, and the transgular stout throughout, as in E. harrisoni and E. perote, it is con- groove is very deep posterior to the submentum (character siderably broader basally, as in E. zongolica (character 13d). The trend toward more deeply impressed sculpticells 32Fb). Also, as in E. zongolica, but not E. harrisoni or on the disc of the elytra also comes to an end with E. cit- E. perote, the right wall of the periostial part of the medi-

an lobe is unmodified (character 37a). On this basis, we formly metallic bluish-purple in color dorsally (character consider E. lasvigas to be most closely related to E. zon- 2b), have the elytral striae uninterrupted on the disc (char- golica. In turn, the median lobe of E. zongolica seems to acter 20a), and with the microsculpture all along the sides, us to most resemble that of E. cuiyachapa, and so we sug- as well as at the apex of the elytra, consisting of sculpti- gest that the three species constitute a monophyletic group cells that are markedly convex (character 6c). This is not (Fig. 56). However, the only derived external morpholog- particularly strong evidence for relationship because the ical trait known to us that links E. cuiyachapa to the other “all purple” condition is probably plesiomorphic relative two species is their comparatively small body size (char- to all other color combinations except for the “all black” acter 1b), a feature they share with some populations of state, while the uninterrupted condition of the elytral stri- E. nitidipennis. The only other evidence for grouping the ae is undoubtedly plesiomorphic (Figs. 58–61). Only the three species together is that they all occur at middle ele- third trait (character 6c) is clearly apomorphic, but it is vations, but this is actually a difficulty because the geo- also exhibited by E. santacatarina, and to a lesser extent graphical ranges of E. lasvigas and E. zongolica are dis- by E. miahuatlan of the dimidiata subgroup, and also by junct, the former being found on the Cofre de Perote, the some of the more derived members of the opaca group latter in the northern part of the Sierra Madre de Oaxaca, (Table 1). Otherwise, E. soladevega does not resemble and E. cuiyachapa in the intervening area on Volcan much the other two members of the soladevega subgroup, Citlaltepetl (Fig. 27b), so that E. cuiyachapa cannot have E. juchatengo and E. nizavaguiti, which in all parsimony diverged first if these three species evolved in situ. analyses were split off together or one after another, and While it appears that E. flohri and E. teotitlan are most separate from E. soladevega (Figs. 59–61). closely related to the E. citlaltepetl-perote-harrisoni stock We place in the dimidiata subgroup (Figs. 55, 57) all of (see above), we are uncertain about the exact relationships the species in Mexico that are uniformly metallic green, or of these two species. Based on geographical distribution, it brassy in color dorsally (characters 2c, 2d), or have a seems unlikely that they are sister species, and it is most metallic green head and pronotum and brassy-colored ely- parsimonious to position E. teotitlan rather than E. flohri tra (character 2e) or a coppery head and bronze elytra next to E. citlaltepetl, etc. (Fig. 56). (character 2f), and with one or more striae interrupted on As already noted, the arrangement based on our mental the disc of the elytra (character 20b), and at least some- analysis is roughly congruent with the altitudinal distribu- what flat elytral intervals (characters 21c, 22c). This evi- tion and kind of forest inhabited by each species in the dence of relationship is not especially convincing because nitidipennis subgroup, with the morphologically most ple- although all of the traits are apomorphic, they are all also siomorphic species at the lowest elevations in forest of exhibited by the more derived members of the opaca most tropical aspect, and the most derived ones at the group (Fig. 55). Moreover, E. flohri of the nitidipennis highest elevations and in forest of the most temperate subgroup also has a metallic green head and pronotum, aspect for the subgroup (Fig. 56). Euchroa harrisoni is an and brassy-colored elytra, while E. soladevega of the exception as it is found at lower elevations than either E. soladevega subgroup, and E. cuiyachapa and E. harrisoni perote or E. citlaltepetl, but that may be because it occurs of the nitidipennis subgroup, all have the elytral intervals at the northern end of the range of the subgroup. While as flat as certain members of the dimidiata subgroup, at these correlations are not perfect, they do provide a basis least on the disc (Table 1). Finally, in some individuals of for recognition of the nitidipennis subgroup. In our parsi- a few species belonging to the dimidiata subgroup, i.e., mony analyses, the species that we have included in the E. atoyac, E. tenancingo, and E. zempoaltepetl, none of the nitidipennis subgroup always emerged together, along striae are interrupted on the disc of the elytra, although in with certain species that we have placed in the soladevega such individuals, stria 1 or 2 is, in most specimens, clear- or dimidiata subgroups, i.e., E. soladevega, E. zempoalte- ly interrupted several times towards the apex of the elytra. petl, E. carbonera, E. suchixtepec, E. miahuatlan, and Other kinds of evidence for monophyly of the dimidia- E. santacatarina, although the exact positioning of the ta subgroup are also equivocal. Many of the species live in species differed greatly depending on how the parsimony comparatively dry oak-pine forest, but so do two of the analysis was done (Figs. 59–61). The possibility that all of three members of the soladevega subgroup. Although the these additional species actually do belong in the geographical range of the dimidiata subgroup does not nitidipennis subgroup is considered next, in the context of overlap with that of the nitidipennis subgroup, except pos- the limits of the soladevega and dimidiata subgroups. sibly for E. dimidiata, the dimidiata and soladevega subgroups are both present in southwestern Oaxaca, and in Soladevega and Dimidiata Subgroups.—Species belong- one instance, members of the two subgroups have been ing to the soladevega and dimidiata subgroups cannot be collected at the same locality. arranged in any sequence according to the structure of the We also suggest that the soladevega and dimidiata sub- median lobe of the aedeagus, that also correlates well with groups have a sister group relationship, but the only mor- an external morphological feature, or any other attribute of phological evidence for this comes from larvae (see these species. We include in the soladevega subgroup (Fig. below). As we have already noted, the soladevega and 55) the three species from western Oaxaca that are uni-

Fig. 59.—One of 12 minimum length trees for adults of species of the subgenus Dyschromus obtained using the RATCHET and NONA programs within the WINCLADA computer program, with most characters ordered and all of them equally weighted. Tree also obtained using successive approximations character weighting, as implemented in NONA, except E. santacatarina n. sp. is split off just before E. suchixtepec n. sp. and E. miahuatlan n. sp. Arrows indicate portions of strict consensus tree of the above 12 trees that are completely unresolved. Selected adult traits are shown. A, color of head and pronotum/color of elytra: Bl, black; Br, brassy; Bz, bronze; C, coppery; G, metallic green; P, metallic bluish purple; R, reddish purple. B, interruption of elytral striae, character 20. C, type of median lobe of aedeagus: E1-3, extreme asymmetric types 1, 2, 3. E4, extreme symmetric type 4. G, generalized asymmetric type. i1-3, intermediate asymmetric types 1, 2, 3. i4, intermediate symmetric type 4. csg, dsg, nsg, osg, ssg: cupripennis, dimidiata, nitidipennis, opaca, and soladevega subgroups.

dimidiata subgroups are heterogenous for type of median groups with an intermediate or extreme asymmetric type 2 lobe of the aedeagus, and so our arrangement requires median lobe emerged together in one cluster, along with some remarkable instances of convergence for characters E. santacatarina with its extreme asymmetric type 3 medi- of the median lobe, i.e., independent origin of an extreme an lobe, while all members of the dimidiata subgroup with asymmetric type 2 median lobe in the nitidipennis and an intermediate or extreme symmetric type 4 median lobe dimidiata subgroups, and of an extreme symmetric type 4 were grouped in a second cluster, with E. soladevega split median lobe in the soladevega and dimidiata subgroups off at the base of the first cluster, and E. juchatengo and (Fig. 58). Our parsimony analyses suggest otherwise, E. nizavaguiti split off together or one after the other at the because all members of the nitidipennis and dimidiata sub- base of the second cluster (Figs. 59–60) or before

Fig. 60.—Minimum length tree for adults of the subgenus Dyschromus obtained using successive approximations character weighting, as implemented in the HENNIG86 computer program, starting with the 12 trees obtained using the RATCHET and NONA programs in WINCLADA, and with most characters ordered. Selected adult traits are shown. A, color of head and pronotum/color of elytra: Bl, black; Br, brassy; Bz, bronze; C, coppery; G, metallic green; P, metallic bluish purple; R, reddish purple. B, interruption of elytral striae, character 20. C, type of median lobe of aedeagus: E1-3, extreme asymmetric types 1, 2, 3; E4, extreme symmetric type 4; G, generalized asymmetric type; i1-3, intermediate asymmetric types 1, 2, 3; i4, intermediate symmetric type 4. csg, dsg, nsg, osg, ssg: cupripennis, dimidiata, nitidipennis, opaca, and soladevega subgroups.

E. soladevega on the branch leading to the first cluster group related to one another? Based upon our mental (Fig. 61). We are skeptical because in all of our analyses of analysis, there are three lineages (Fig. 57). One lineage the larval data (see below), E. soladevega was grouped comprises E. jalisco, E. tenancingo, E. ixtapa, and with E. dimidiata, E. filodecaballo, and E. atoyac, the E. dimidiata, all of which live in uplands that border upon adult males of which have an intermediate or extreme the Rio Balsas Basin, the first two species being known symmetric type 4 median lobe, rather than with E. perote only from the central and western parts of the Trans- and E. harrisoni, which, in the adult male, have an extreme Volcanic Sierra, respectively, and E. ixtapa from the west- asymmetric type 2 median lobe. ernmost part of the Sierra Madre del Sur (Figs. 37b, 42a). How are the species that belong to the dimidiata sub- Besides geographic distribution, other evidence for rela-

tionship is that all four species have an extreme symmetric the elytral intervals on the disc (character 21e), and the type 4 median lobe (e.g., Figs. 40a–c), the preapex being fact that four of the five species occur at higher elevations longer than in species with an intermediate symmetric type in forest of more mesic or temperate aspect than E. atoy- 4 median lobe (Fig. 57, character 30c). Relationship of the ac, the exception being E. santacatarina. In many respects last three species is further supported by their coloration, E. atoyac resembles E. jalisco, a member of the first line- the head and pronotum being metallic green and elytra age, but besides its geographical distribution, and the con- brassy (character 2e), by the considerable extent to which dition of the labial palps, relationship of E. atoyac to the the elytral striae are interrupted (characters 20d, 20e), by other species in the second lineage is supported by evi- the condition of the apical declivity of elytra, which is dence from larvae that it is more closely related to E. puer- nearly perpendicular in most specimens (character 23b), togallo than to E. dimidiata. and by their presence in drier forest formations than In regards to the most derived members of the second E. jalisco, which is uniformly metallic green in color dor- lineage, the long preapex of the median lobe is evidence sally (character 2c), has nearly entire elytral striae (charac- that E. yucuyacua is most closely related to E. santacata- ter 20b), and comparatively gentle apical declivity of the rina (character 30c). Alternatively, the stoutness in lateral elytra (character 23a), and lives in mesophytic forest. aspect of the middle part of the median lobe distal to the Close relationship of E. ixtapa to E. dimidiata is indicated basal bend (character 39b), and their similar external by the greatly interrupted elytra striae (character 20e), or appearance, indicates that E. filodecaballo and E. chryso- of E. ixtapa to E. tenancingo by the slenderness of labial phana are sister species, but absence of at least one of the palpomere 3 of the female (character 9d). basal punctures of the elytra (character 19b), and slender- According to the mental analysis, the second lineage in ness of the middle tibia in at least some males (character the dimidiata subgroup includes, in order of relationship, 27b), provides evidence that E. yucuyacua is most closely E. atoyac, E. puertogallo, E. filodecaballo, E. chryso- related E. chrysophana. phana, E. yucuyacua, and E. santacatarina (Fig. 57), of Based on our mental analysis, the third lineage in the which the first four species are known only from the Sierra dimidiata subgroup contains the four species from Oaxaca de Atoyac and vicinity in Guerrero, and the other two from that have an intermediate (E. zempoaltepetl, E. carbonera) the adjoining portion of the Sierra Madre del Sur in north- or extreme (E. miahuatlan, E. suchixtepec) asymmetric western Oaxaca (Figs. 42, 46a). Besides geographical dis- type 2 median lobe, and a metallic green head and prono- tribution, other evidence for relationship of these species is tum and brassy-colored elytra, except for E. suchixtepec, that in both sexes labial palpomere 3 is comparatively which is all green dorsally (Figs. 57–58, characters 2e, slender (characters 8d, 9c, 9d). However the palps are also 32d, 32e, 33b, 34c, 35c, 37b, 38d, 40c). A sister group rela- this slender in some other members of the dimidiata sub- tionship between E. miahuatlan and E. suchixtepec is indi- group. The species in the second lineage can be arranged cated not only by their having an extreme asymmetric type according to the structure of the male genitalia (Figs. 2 median lobe (character 32e), but also by the relative 57–58, characters 30c, 33b, 34b), starting with E. atoyac, slenderness of labial palpomere 3 in the female (character E. puertogallo and E. filodecaballo, all of which have an 9c), short pronotal basolateral impressions (character 16b), intermediate symmetric type 4 median lobe (Figs. 43–44, and the presence of a ventrally-directed bulla along the left 45a–c), continuing with E. chrysophana and E. yucuy- margin of median lobe at the base of the preapex (charac- acua, both of which have an intermediate type 3 median ter 36b). Also, in both species the sides of the pronotum lobe (Figs. 45d–f, 47a–c), and ending with E. santacatari- are oblique posteriorly, and the hind angles are rounded; na, which has an extreme asymmetric type 3 median lobe although both conditions are plesiomorphic for the sub- (Figs. 47d–f). This morphocline is paralleled by another genus (characters 14a, 15a), in the dimidiata subgroup involving increasing interruption of the eltytral striae, each is restricted to the more derived species. Closer rela- beginning with E. atoyac, which has nearly entire striae tionship of E. carbonera to E. miahuatlan and E. suchixte- (character 20b) and ending with E. yucuyacua, which has pec than to E. zempoalteptl is indicated by absence of the greatly interrupted striae (character 20e), with the other basal elytral punctures (character 19c), and the compara- species exhibiting an intermediate condition (character tively flat elytral intervals (character 21d), as well as geo- 20d). A third morphocline is with respect to body color, graphic proximity, with only E. zempoalteptl being found E. atoyac being all green (character 2c), and the others uni- in eastern Oaxaca. formly brassy (character 2d), except for E. yucuyacua, A serious difficulty with division of the dimidiata sub- which has a coppery head and pronotum and bronze col- group into the above three lineages is to establish how the ored elytra (character 2f). The above three morphoclines, third lineage comprising E. zempoaltepetl, E. carbonera, taken together, indicate closer relationship of E. puertogal- E. miahuatlan, and E. suchixtepec is related to the other lo, E. filodecaballo, E. chrysophana, E. yucuyacua, and two lineages. All of the species in the first lineage except E. santacatarina to one another, than to E. atoyac. Further for E. jalisco have a metallic green head and pronotum, evidence is the shallow condition of the postocular trans- and brassy colored elytra, as do all of the species in the verse groove (characters 11b), the entirely flat condition of third lineage except for E. suchixtepec. This would consti-

Fig. 61.—Minimum length tree for adults of the subgenus Dyschromus obtained using successive approximations character weighting, as implemented in the NONA computer program, with all characters unordered. Arrows indicate nodes resolved in strict consensus tree of 328 trees obtained prior to doing successive approximations character weighting. Selected adult traits are shown. A, color of head and pronotum/color of elytra: Bl, black; Br, brassy; Bz, bronze; C, coppery; G, metallic green; P, metallic bluish purple; R, reddish purple. B, interruption of elytral striae, character 20. C, type of median lobe of aedeagus: E1-3, extreme asymmetric types 1, 2, 3. E4, extreme symmetric type 4. G, generalized asymmetric type. i1–3, intermediate asymmetric types 1, 2, 3. i4, intermediate symmetric type 4. csg, dsg, nsg, osg, ssg: cupripennis, dimidiata, nitidipennis, opaca, and soladevega subgroups.

tute evidence of relationship of the first and third lineag- E. carbonera, E. miahuatlan, and E. suchixtepec as a es, provided that E. jalisco has a more basal position in group, are most closely related to one or more members of the dimidiata subgroup, and we note that labial palpomere the second lineage that have an asymmetric type 3 medi- 3 in the male of E. jalisco is broader than in any other an lobe, i.e., E. chrysophana, E. yucuyacua, and/ or E. member of the dimidiata subgroup (character 8b). santacatarina, but there is no evidence for this from However, we cannot envision direct derivation of an external morphology. extreme asymmetric type 2 median lobe from a extreme If E. zempoaltepetl, E. carbonera, E. miahuatlan, and symmetric type 4 median lobe or vice versa. Derivation of E. suchixtepec all actually belong to the nitidipennis suban extreme asymmetric type 2 median lobe from an inter- group instead of the dimidiata subgroup, as is indicated by mediate or extreme asymmetric type 3 median lobe seems our parsimony analyses (Figs. 59–61), then the problem to us more probable, and so perhaps E. zempoaltepetl, disappears of how to derive the extreme type 2 median

Fig. 62.—Phylogenetic relationships for six species belonging to the subgenus Dyschromus as indicated by characters of the larvae. Underlining indicates homoplasy.

lobe from one of the other types, at least for the E. zem- in E. santacatarina is derived from the extreme asymmet- poaltepetl-carbonera-miahuatlan-suchixtepec stock. In ric type 2 median lobe shown by E. miahuatlan and one of our parsimony analyses, these four species, along E. suchixtepec, and all resemblances to the intermediate with E. santacatarina, were positioned as the sister group asymmetric type 3 median lobe of E. chrysophana and of E. flohri plus the rest of the nitidipennis subgroup (Fig. E. yucuyacua are owing to reversion in E. santacatarina to 59), E. flohri being the only member of the nitidipennis a more plesiomorphic condition for those traits, i.e., the subgroup with a metallic green head and pronotum and flat preapical dorsolateral ridge (character 34b) and the brassy colored elytra, but such an arrangement conflicts inconspicuous periostial ventrolateral bulge (character with evidence that E. nitidipennis, E. sallei, E. huautla, 38Ca). and E. cuiyachapa occupy basal positions within the We do not dismiss the possibility that E. zempoaltepetl, nitidipennis subgroup. In another parsimony analysis, no E. carbonera, E. suchixtepec, E. miahuatlan, and perhaps such conflict was evident, the five species being as group E. santacatarina, all actually belong to the nitidipennis paired together with E. citlalteptl, E. perote, and E. har- subgroup, and consistent with this alternative is the fact risoni (Fig. 60) because in all of them, the left wall of the that the least derived species, E. zempoaltepetl, lives in periostial part of the median lobe is not as extensive as the eastern rather than western Oaxaca. We do consider below right wall (character 37b), but evidence from external mor- the biogeographic and evolutionary implications of this phology for this arrangement is lacking. As for E. santa- possibility, but we note that the problem of deriving an catarina, in most parsimony analyses it was paired just asymmetric type 2 median lobe from a symmetric type 4 with E. miahuatlan (Figs. 59–61); this is in part because median lobe, or vice versa, would still remain for the the preapex is only developed to the right of the midline of soladevega subgroup. the median lobe (character 32e). Moreover, E. santacata- Under most or all arrangements that we have consid- rina not only has the features of the pronotum, and of labi- ered, certain species belonging to the nitidipennis group al palpomere 3 in the female that we cite above as evi- stand out as being morphologically very different from dence for a sister group relationship between E. suchixte- their presumed nearest relatives, i.e., E. soladevega, pec and E. miahuautlan (characters 9c, 15a, 16b), but as in E. nitidipennis, E. dimidiata, E. yucuyacua, and E. santa- E. miahuatlan, labial palpomere 3 in the male is compara- catarina. The long branch lengths at the relevant nodes of tively slender (character 8d), and the basal sulcus of the cladograms (Appendix 4) suggest to us that these abdominal segment VII lacks a median fovea (character species do not have any close living relatives, or they have 26c). We also note that the extent to which the surface at not been discovered yet. We could erect a separate group the sides of elytra is beaded in E. santacatarina is or subgroup for each of these divergent species, but the approached by E. miahuatlan. If these two are sister result would be an excessive number of monospecific taxa. species, then the extreme asymmetric type 3 median lobe We believe that recognition of the soladevega and dimidi-

ata subgroups provides a sufficient basis for discussion cladograms, E. soladevega was positioned at the base of the and further investigation of taxonomic problems in the branch that contains E. perote and E. harrisoni. However, nitidipennis group, and choose not to further subdivide in our mental analysis of the adult data, E. soladevega, as a either subgroup. member of the soladevega subgroup, was grouped with the dimidiata subgroup instead of the nitidipennis subgroup (of which E. perote and E. harrisoni are members) based Analysis of the Larval Data, upon the results of the analysis of the larval data. Procedures and Results Moreover, altering the larval tree (Fig. 62) so that it is con- The larval data was analyzed using the same procedures as sistent with cladograms obtained by parsimony analysis of for the adult data, except that it was not necessary to do the adult data (Figs. 59–61), by grouping E. soladevega any character weighting. Parsimony analysis of the addi- with E. perote and E. harrisoni, only increases the length tively binary coded larval data, with all characters weight- of larval tree by one step. This is because the number of ed equally, yielded just two minimum length trees, these character differences between larvae of the six species is having a length of 17 steps, CI of 0.72, and RI of 0.57. small. In fact, they are so similar that we do not know of a These same two trees were recovered using compatibility uniquely derived larval trait for E. perote or for E. solade- analysis, and one of them was also obtained by mental vega. Yet, according to the arrangements obtained from analysis (Fig. 62). With all unordered character states, just our analyses of the adult data (Figs. 58–61), these six one minimum length tree was found; this had a length of species represent some of the most distantly related lineag- 13 steps, CI of 0.94, and RI of 0.83. Only the arrangement es of the subgenus Dyschromus in Mexico. We expect that shown in Fig. 62 proved to be largely or entirely consistent larvae of many of the species for which the immature stages are unknown will prove to be identical to one anoth- with the trees that were obtained either by mental analysis er or nearly so, and that some of the traits discussed above (Fig. 58) or parsimony analysis of the adult data (Figs. will characterize entire lineages. 59–61), so neither the second tree obtained by parsimony Based upon the arrangement in Fig. 62, certain reduc- analysis of the ordered larval characters, nor that obtained tions in size and number of setae that have taken place in with the larval characters unordered are considered further. larvae of the nitidipennis subgroup have also occurred The arrangement obtained both by parsimony and com- within the soladevega + dimidiata subgroups, but not nec- patibility analysis, as well as by mental analysis of the lar- essarily to the same extent (character 4); the converse is val data (Fig. 62), has E. harrisoni and E. perote on one also true (characters 1, 3). branch, and E. soladevega, E. dimidiata, E. atoyac and Of the four larval characters used in this analysis that E. filodecaballo split off in that order on a second branch. show variation from one instar to the next, based on out- Euchroa harrisoni and E. perote were paired together group comparison, we find that two conform to the notion because the second and third instars have fewer long sub- that morphological changes first occur in the later devel- primary setae on the urogomphi than the other four opmental stages, i.e., the change first took place in the last species. The larva of E. harrisoni is more derived in this instar (Appendix 3, characters 6 and 7). We infer the respect because it has only seta epsilon (Appendix 3, char- change to the derived condition of the other two traits ini- acter 4d); larvae of E. perote also have seta gamma, tially took place in the first one or two instars (characters although it is short or minute (character 4c). Euchroa 1, 4). Such contradictions between the evidence soladevega, E. dimidiata, E. atoyac, and E. filodecaballo for polarity from outgroup comparison and ontogeny were grouped together on another branch because in all have been noted by Wheeler (1990) for Agathidium instars of the first three species and at least the first instar (Leiodidae). of E. filodecaballo frontale seta FR4 is minute (character 1c). Closer relationship of E. atoyac to E. filodecaballo than to E. dimidiata or E. soladevega is indicated by the BIOGEOGRAPHIC CONSIDERATIONS minute condition of several primary setae in larvae of these two species (character 2b), and presence of just five Distribution Patterns of the Subgenera stemmata on each side of the head (character 5b), while of Euchroa,Overall and in Mexico, closer relationship of this pair of species to E. dimidiata and Their Interpretation than to E. soladevega is indicated by the condition of the mesonotal carina, which extends posteriorly to seta 12 The previously published locality data for the subgenus (character 7b). Dyschromus consist of a few records from the island of We have already noted that the arrangement based upon Hispaniola, and some additional localities for Mexico, i.e., larval characters shown in Fig. 62 is entirely congruent the eastern part of the Trans-Volcanic Sierra (Las Vigas, with the tree obtained by mental analysis of the adult data Jalapa, Orizaba), the interior of Oaxaca (Sierra Mixteca (Fig. 58), but it is also nearly congruent with the clado- Alta), and the Sierra Madre del Sur in Guerrero (Amula, grams that emerged in our parsimony analyses of the adult Omilteme). We now know that the subgenus occurs at least data (Figs. 59–61), the difference being that in the adult locally throughout the mountains of Oaxaca and Guerrero,

Fig. 63.—Transformation series for types of median lobe of the aedeagus within the subgenus Dyschromus. Solid arrows are based on apriori arrangement of the types, and are consistent with tree obtained by mental analysis of the adult data (Fig. 58). Broken arrows show alternate pathways, as indicated by cladograms obtained in parsimony analyses of the adult data (Figs. 59–61). E1-3, extreme asymmetric types 1, 2, 3. E4, extreme symmetric type 4. G, generalized asymmetric type. i1-3, intermediate asymmetric types 1, 2, 3. i4, intermediate symmetric type 4.

the Trans-Volcanic Sierra, and the southern part of the Euchroa are also evident at lower taxonomic levels in the Sierra Madre Oriental. It remains unknown from the Sierra subgenus Dyschromus. The obvious gap in distribution is Madre Occidental, or south of the Isthmus of Tehuantepec, between that of the nitidipennis group, which comprises or anywhere else except the Dominican Republic and Haiti all of the species of Dyschromus in Mexico, and its sister, (Figs. 4, 54). Although we have increased the number of the opaca group, which contains all of the species on named species of Dyschromus in Mexico from from five to Hispaniola (Fig. 4). Other distributional anomalies are 29, we have not substantially altered the geographical dis- revealed by considering each species group in turn. tribution of the subgenus as it was understood at the beginning of the last century. Similarly, the subgenus Euchroa, Patterns of Distribution in the Nitidipennis Group.—Of while somewhat more widespread than previously indicat- the 28 species of Dyschromus now described from ed, is still only known from the mountains along the Mexico, only E. dimidiata is represented in more than one Atlantic coast of Brazil, and the eastern foothills of the mountain region (Fig. 42a); most species are known from Andes in Ecuador (Fig. 4; Shpeley and Araujo 1997). just a single mountain range. Eleven species are known only from the mountains of eastern Mexico, north of the Patterns of Distribution in the Subgenus Dyschromus.— Rio Santo Domingo (Fig. 54); all belong to the nitidipen- The restricted ranges and marked disjunctions that charac- nis subgroup. Another 13 species are known just from the terize the distribution patterns of the two subgenera of Sierra Madre del Sur or adjoining uplands of the interior of

western Oaxaca and Guerrero (Fig. 54); all of them we suggest that for most, if not all, of its existence, the have placed either in the soladevega or dimidiata sub- nitidipennis subgroup has evolved in isolation in the groups. Only three species, besides E. dimidiata, have humid mountain forests of eastern Mexico, north of the been collected outside of these two regions: E. zempoal- Rio Santo Domingo, and that the entire area as far north as teptl, from the vicinity of Cerro Zempoaltepetl in eastern the Cofre de Perote has been continuously occupied by Oaxaca (Fig. 46b), and E. jalisco and E. tenancingo, from these beetles during that period. the western and central parts of the Trans-Volcanic Sierra, Southwestern Mexico.—The situation pertaining to the respectively (Fig. 37b). The closest relatives of all three distribution of species of the subgenus Dyschromus in the species are in the Sierra Madre del Sur. Thus the distribu- Sierra Madre del Sur and adjoining uplands is more com- tion of the subgenus Dyschromus in central Mexico, while plex than in eastern Mexico. Two areas of the Sierra Madre not strictly disjunct, is evidently centered on two distant del Sur have concentrations of species of Dyschromus: one regions, one in the northeast, the other in the southwest, area comprises the Sierra Miahuautlan, Cerro Yucuyacua, each occupied by different subgroups of Dyschromus. and adjoining uplands in western Oaxaca, from which we There are differences besides taxonomic composition as to have eight species besides E. dimidiata (Fig. 54); the sec- how species of Dyschromus are distributed in each region. ond area is centered on the Sierra de Atoyac in south- Eastern Mexico.—The species of Dyschromus in the central Guerrero just west of Chilpancingo, where we mountains of eastern Mexico, north of the Rio Santo know of at least four species of Dyschromus, in addition to Domingo, are all restricted to the humid forest formations a distinct population of E. dimidiata. From that portion of that cover (or covered) much of the region. Three adjoin- the Sierra Madre del Sur between Cerro Yucuyacua and ing areas have concentrations of species of Dyschromus: Chilpancingo, a distance of about 250 km, we have seen from north to south, the Cofre de Perote and vicinity; only a single specimen of Dyschromus that perhaps repre- Volcan Citlalteptl and environs; and the Sierra Madre de sents yet another distinctive population of E. dimidiata. Oaxaca, north of the Rio Santo Domingo. At least three or Some high points in the area in question attain an elevation four species occur in each area, all of which live at least in of 3000 m, so the question arises whether the gap is real or part at different elevations, e.g., E. nitidipennis and due to a lack of collecting? We can only suggest that if E. lasvigas; or if two species occur at the same elevations, there are any species of Dyschromus other than E. dimidi- then one is found further inland than the other in forest of ata, then they are confined to the highest peaks, as more temperate aspect, e.g., E. flohri and E. perote. Only E. yucuyacua appears to be on Cerro Yucuyacua. With E. nitidipennis and E. sallei are known to occur in more regard to the homogeneity of the Dyschromus fauna, alti- than one of these three areas; along with E. huautla they tudinal zonation of the species, etc., the situation in the live at lower elevations in forest of more tropical aspect vicinity of the Sierra de Atoyac is comparable to that in than other species of the nitidipennis subgroup. eastern Mexico; the situation in western Oaxaca is not. According to the phylogeny for the nitidipennis sub- The Dyschromus fauna of the Sierra de Atoyac and group derived by mental analysis (Fig. 56), the closest rel- vicinity is comparatively homogeneous, since except for ative (or relatives) of a given species is to be found in one E. dimidiata, the species present in those mountains, of the other two areas at about the same elevation in a namely, E. atoyac, E. puertogallo, E. filodecaballo, and comparable type of forest; the most plesiomorphic species E. chrysophana all belong to the same lineage, are restrict- live at the lowest elevations in forest of most tropical ed to the humid forest formations of the area, and appear aspect, while the most derived ones are at higher eleva- to be distributed in a fairly regular manner, the most ple- tions in forest of more temperate aspect. Either of two evo- siomorphic one, E. atoyac, occurring on the western ver- lutionary scenarios are sufficient to account for, in a gen- sant at lower elevations in forest of more tropical aspect eral way, the pattern of distribution shown by the than the others, which are found at higher elevations in nitidipennis subgroup in eastern Mexico: 1) speciation in cloud forest or mesic oak-pine forest, either on the high each of the three areas of species concentration of a wide- ridges that flank Cerro Teotepec (E. puertogallo, E. filode- spread ancestral stock living at comparatively low eleva- caballo), or on mountains nearby (E. chrysophana). The tions, followed by occupation of middle elevation forests Dyschromus fauna of western Oaxaca differs because of by one of those species with subsequent divergence from its composite nature, and because in addition to a humid the lower elevation species, and then spread of the mid- forest element represented by E. juchatengo, E. santacata- elevation stock to and speciation in adjoining areas, and rina, and E. suchixtepec, there is a dry forest element that finally the same sequence of events at the highest eleva- includes E. nizavaguiti, E. soladevega, E. yucuyacua, tions occupied by the subgroup; or 2) fragmentation of the E. carbonera, and E. miahuatlan, all of which live in com- ranges of three formerly widespread species, one living at paratively dry oak or oak-pine forest. All indications are comparatively low elevations, one at middle elevations, that the species of the soladevega subgroup represent an and one at relatively high elevations. The computer old component of the fauna in western Oaxaca (Figs. derived phylogenies (Figs. 59–61) each lead to somewhat 58–61). The status of the other species in western Oaxaca different biogeographic scenarios, but all of our results is problematic; for example, depending upon its phyloge-

netic relationships, the lineage represented by E. carbon- genera of Euchroa, and of the species groups of the sub- era, E. miahuatlan and E. suchixtepec, with or without genus Dyschromus, that these taxa exhibit a relictual dis- E. santacatarina, could be an old component of the fauna tribution. Since, except for the subgenus Dyschromus, all of western Oaxaca (Fig. 58), or a more recent addition other members of the Euchroa-Microcephalus clade are related in one way or another to the nitidipennis subgroup restricted to northern South America, we propose that the (Figs. 59–60). The various components of the Dyschromus subgenus Dyschromus is an old South American element fauna in western Oaxaca overlap widely in range, with of the Middle American fauna (Savage 1982; Halffter actual sympatry of E. juchatengo and E. santacatarina. 1987), so that the ancestor of Dyschromus entered Middle An anomaly is the large gap between the ranges of America from South America in late Cretaceous or early E. juchatengo and its sister species, E. nizavaguiti. Tertiary time. Furthermore, because the species of An additional component of the Dyschromus fauna in Dyschromus on the island of Hispaniola comprise a single the Sierra Madre del Sur comprises two species that also monophyletic group (opaca group), and those in central live in the central part of the Trans-Volcanic Sierra or have Mexico another (nitidipennis group), and all are flightless, a close relative there, i.e., E. dimidiata and E. ixtapa. Their we infer that the two stocks have been isolated since the closest relative is E. tenancingo (Figs. 58–61); all three of late Eocene epoch, at which time, according to some mod- them live in the dry oak or oak-pine woodlands that fringe els of Caribbean plate tectonics, a portion of the island of the Rio Balsas Basin and Valley of Oaxaca. Although Hispaniola was last in close proximity to, or in contact E. dimidiata is comparatively widespread, the populations with, Nuclear Middle America (Rosen 1985). The stock with the largest number of plesiomorphic traits are those in ancestral to the nitidipennis and opaca groups must have the central part of the Trans-Volcanic Sierra. An anomaly resided in Nuclear Middle America, and has since become is that neither E. dimidiata nor any species closely related extinct. to it is known from south of the latitude of Oaxaca City. As far as we know, all members of the subgenus Despite our uncertainty about the phyletic relationships Euchroa live in wet forests of tropical aspect at compara- of the species belonging to the soladevega and dimidiata tively low elevations, and with the exception of E. nizava- subgroups, it is apparent that the humid forest elements of guiti, so do those species of the subgenus Dyschromus the Dyschromus fauna of central Mexico have thrived only with the largest number of plesiomorphic morphological in eastern Mexico north of the Rio Santo Domingo, and in traits, both in eastern and western Mexico: i.e., the vicinity of the Sierra de Atoyac in southwestern E. nitidipennis, E. sallei, and E. huautla on the eastern ver- Mexico. Another humid forest element, represented only sant; and E. juchatengo, E. atoyac, and E. jalisco on the by E. jalisco, has persisted in the extreme western part of western versant. Moreover, outside of eastern Mexico, the Trans-Volcanic Sierra. All three areas have long been north of the Rio Santo Domingo, the more derived taxa recognized as having a considerable endemic element in inhabit drier forest formations, with the exception of their biota (Martin 1958; Rzedowski and McVaugh 1966; E. suchixtepec and E. santacatarina. We conclude that the Halffter 1987; Llorente-Bousquets and Luis-Martinez main features of the distribution pattern were established 1993), including some ancient lineages (Davidson and during the Miocene epoch, when according to Cohn Ball 1998). (1965) and Axelrod (1975), development of dry conditions Elsewhere in central Mexico, with the exception of owing to uplift of the Mexican Plateau, widespread moun- E. juchatengo, E. santacatarina, and E. suchixtepec in the tain building, and climate change, led to the appearance Sierra Miahuatlan, species of Dyschromus are known only and spread in Mexico of dry-adapted vegetation types, and from drier forest formations. Some of these dry-forest taxa at the same time, confinement of humid montane forests of are the most derived members of the lineages to which tropical aspect to the eastern and western versants of they belong, i.e., E. soladevega and E. dimidiata. More- Mexico. We suggest that a widespread species of over, outside of northeastern Mexico, the most widespread Dyschromus living in these humid forests also became stocks of Dyschromus are those that for the most part live confined to the east and west coasts of Mexico. in the drier forest formations. Even so, large areas of Subsequent fragmentation of the humid forests, on the central Mexico are without any species of Dyschromus. western versant, led to confinement of Dyschromus to Even the range of E. dimidiata, the only species with a three areas: the western part of the Trans-Volcanic Sierra, comparatively wide geographical range, appears to be dis- the vicinity of the Sierra de Atoyac, and the vicinity of the continuous, although the scrubby oak and oak-palmetto Sierra Miahuatlan in western Oaxaca. On the east coast, woodlands where it lives are more continuously distrib- the stock there became confined to the mountains north of uted in Mexico than other types of forests inhabited by the Rio Santo Domingo. Only in eastern Mexico, north of species of Dyschromus. the Rio Santo Domingo, and in the vicinity of the Sierra de Atoyac in western Mexico did humid forests remain suffi- Interpretation of the Overall Pattern of Distribution, ciently extensive for stocks of Dyschromus to diversify and that Shown by the Nitidipennis Group in without shifting to drier forest formations, as apparently Mexico.—We infer from the restricted ranges and marked happened elsewhere in central Mexico. disjunctions that characterize the distributions of both sub-

A detailed biogeographic analysis awaits a better shown by Dyschromus in Mexico is of long standing. understanding of the phylogenetic relationships of these Consistent with this, mDNA evidence indicates that many beetles, but one aspect of the pattern is worth pursuing fur- of the speciation events that have taken place among the ther, the apparent absence of humid-forest-adapted species Mexican species of Dyschromus predate the Pliocene of the subgenus Dyschromus from the mountains of east- epoch (Sperling, Frania, and Ball, in prep.). ern Oaxaca, south of the Rio Santo Domingo. We have been reluctant to accept this as fact because at least four members of the nitidipennis subgroup occur in the humid Distribution Patterns of the Opaca Group forests of the mountain ranges of the Sierra Madre de in Haiti and the Dominican Republic Oaxaca north of the Rio Santo Domingo, i.e., E. huautla, and Their Interpretation E. sallei, E. zongolica, and E. teotitlan. Another reason for Just as in Mexico, the species of the subgenus Dyschromus our hesitation is that the only apparent barrier to dispersal, on the island of Hispaniola have very restricted distribu- the breach in the Sierra Madre de Oaxaca created by tions. Interpretation of the pattern of distribution on the downcutting of the Rio Santo Domingo, is considered to island of Hispaniola is relatively straightforward: diver- be of comparatively recent origin, i.e. early Quaternary gence in different parts of southern Hispaniola of the (Brunet 1967). At the opposite end of the range of the tiburonica, opaca, and centralis stocks, and also of the nitidipennis subgroup, a single species, E. harrisoni, does ancestor of the cupripennis subgroup, followed by specia- occur in the Sierra Madre Oriental both north and south of tion within the cupripennis subgroup, which was accom- the canyons of the Rio Panuco watershed, which are con- panied by the eventual spread over much of the western sidered to be a significant barrier to dispersal of plants and part of the island of the most derived phyletic line in animals (Smith 1940; Vega et. al. 1999). We also note that the cupripennis subgroup, the E. cupripennis-perezi- the mountains of eastern Oaxaca are difficult to collect, pedernales stock. Associated geologic events and timing, being very rugged and comparatively inaccessible, with including arrival on southern Hispaniola, were presumably only one road crossing from the interior of Oaxaca over the same as for some species of the carabid genus Platynus the mountains and down to the coastal plain, Hwy. 175, on Hispaniola (Liebherr 1988), except for subsequent which from Ciudad Oaxaca crosses the Mije Highlands extinction in Nuclear Middle America of the common and Sierra Juarez to Valle Nacional. Perhaps E. nitidipen- ancestor of the nitidipennis and opaca groups. nis will eventually be discovered in the Sierra Madre de Oaxaca south of the Rio Santo Domingo, but it now seems to us unlikely that there exists a counterpart to the rich CHARACTER TRENDS IN ADULTS AND Dyschromus fauna of the humid forests of the mountains ECOLOGICAL CORRELATES of eastern Mexico, north of the Rio Santo Domingo. It is useful to contrast the situation in the two subgen- Body Color.—We can now answer to a certain extent the era of Cyrtolaus Bates, another group of flightless pteros- question posed in the introduction to this monograph about tichines of similar size and habits as Dyschromus. Closely the brilliant metallic colors of many of these beetles. The related species belonging to the subgenus Cyrtolaus do combination of a metallic green head and pronotum, and live in the mountain ranges on either side of the Rio Santo brassy to coppery-colored elytra has evolved independent- Domingo, i.e., C. whiteheadi Ball, which is known only ly in the opaca and nitidipennis groups, and several times from the type-locality for E. huautla (Ball 1991), and in the latter group (Figs. 58–61). Both in central Mexico C. newtoni and C. oaxacanus Whitehead and Ball, which and on the island of Hispaniola, the most widespread live further south in the Sierra Juarez and Sierra phyletic lines, and the only comparatively widespread Zempoaltepetl, respectively (Whitehead and Ball 1975). species, all exhibit this combination of colors. In Mexico, Still other species of the subgenus Cyrtolaus live in the most of these taxa occur in the drier forest formations of highlands of Chiapas and Guatemala. The other sub- the interior. The significance of differences in body color genus, Ithytolus, represented only by C. orizabae, is of ground-dwelling carabids is generally unknown (Erwin known only from the vicinity of Volcan Citlaltepetl in 1979:547–548). We surmise that it is in some way advan- Veracruz. This pattern is consistent with the situation in tageous for species of Dyschromus living in dry forests to Dyschromus, and is indicative of a long period of isola- be colored in such a manner. Whether there is a direct tion of the humid montane forest biota of eastern Mexico advantage, or whether the colors merely are a reflection of north of the Rio Santo Domingo from that further south physiological or developmental differences between in eastern Oaxaca and the highlands of Chiapas and species living in different conditions remains unknown. Guatemala. North of the Rio Santo Domingo, Dyschromus has thrived, Ithytolus has merely persisted, Structures Associated with Flight.—Besides having ves- and the stock represented by C. whiteheadi is a compar- tigial hind wings, adults of all species of both subgenera of atively recent arrival from the south. Euchroa also exhibit at least somewhat vaulted elytra, and We conclude that the overall pattern of distribution reduced metasternum, and at least in some species the elytra are fused. This degree of modification of structures

associated with flight is considered to indicate that the taxa (character 28). Also a trend is apparent both in the opaca in question have been unable to fly for a considerable peri- group and some phyletic lines of the nitidipennis group od of geologic time (Darlington 1943; Kavanaugh 1985). towards increasing asymmetry of the distal part of the This leads to the question as to whether the common median lobe of the aedeagus, although this has occurred in ancestor of the two subgenera was also flightless. Possibly, different ways in the two groups. Far more variation is evi- but if so, the rounding of the shoulders of the elytra, anoth- dent for aedeagal type in the nitidipennis group, including er change associated with loss of the ability to fly, has gone the appearance of a more symmetric type of median lobe further in the opaca group on the island of Hispaniola, as (Fig. 63). well as in the subgenus Euchroa in northern South All evidence (Figs. 58–61) indicates that the extreme America, than in the nitidipennis group in Mexico, since asymmetric type 1 median lobe was derived once within the shoulders of the elytra of species belonging to the lat- the opaca group from the generalized asymmetric type via ter group are prominent, and even dentiform in many of intermediate asymmetric type 1, that intermediate asym- the species. metric type 2 and intermediate symmetric type 4 were derived within the nitidipennis group from a type in com- Other External Traits.—Other features, besides body mon rather than independently from the generalized asym- color, that appear to be correlated with way of life are the metric type, and that the extreme asymmetric type 2 medi- sculpticells on the elytra becoming more deeply impressed an lobe was derived one or more times from intermediate and convex (characters 4c, 6c), and the striae on the disc asymmetric type 2 (Fig. 63). Contrary to the tree obtained of the elytra becoming more interrupted (characters 20b to by mental analysis (Fig. 58), all of the cladograms 20e). Both tendencies are evident in taxa that live in drier obtained in our parsimony analyses of the adult data indi- forest formations, although some species exhibit one ten- cate that extreme asymmetric type 3 was derived from dency and not the other. Still other changes in external fea- intermediate or extreme asymmetric type 2 (Figs. 59–61) tures shown by certain members both of the nitidipennis rather than from intermediate asymmetric type 3 (Fig. 58), and s do not show any correlation with habitat: effacement while one cladogram (Fig. 61) indicates that intermediate of the dorsomedial portion of the postocular transverse asymmetric type 3 was derived from a intermediate asym- groove (character 11); decrease in size or secondary metric type 2, rather than from a intermediate or extreme increase in size of paramedian pits (character 12); loss of symmetric type 4 median lobe. the marginal bead of the prosternal intercoxal process (character 13); shortening of the pronotal basolateral impressions (character 16); decrease in punctation of CONSERVATION OF THE MONTANE BIOTA mesepisternum (character 24). OF MEXICO A few changes in external traits that have taken place more than once apparently have been confined to the Ball and Shpeley (2000: 385–386) drew attention to the nitidipennis group: increase of both the prominence of the need for protecting the montane forests of Mexico, if the submentum and depth of the transgular groove (character incredibly rich indigenous biota that is concentrated in 13); sides of pronotum becoming more sinuate posteriorly, those forests is to survive. The same may be said for the and hind angles becoming more prominent (characters 14, island of Hispaniola. The montane forest-inhabiting sub- 15); loss of the elytral basal punctures (character 19); genus Dyschromus, with its numerous localized species, increase in steepness of the apical declivity of the elytra offers a prime example of a taxonomic group that is vul- (character 23). As well, the species in Mexico have nerable to disturbance caused by habitat destruction, and evolved a greater range of body colors than those on thus subject to extinction. Hispaniola (character 2). No trends are evident that are confined to the opaca group, although a few have gone SUMMATION further, such as the shoulders of elytra becoming more rounded (character 18), and labial palpomere 3 becoming Our impression is that the overall pattern of diversification more slender (characters 8, 9). Only a few derived traits in the subgenus Dyschromus is relatively simple, was are confined to a single species, or to two or three closely established early, and interspecific competition has been related species, such as the explanate sides of pronotum in minimal. Although extinction of humid forest elements in E. tiburonica, and the considerably greater extent of the Mexico has been considerable, there has occurred a mod- basal sulcus of the abdomen in E. tenancingo than in other est radiation of a few lineages into drier types of forest. On species (character 25). the island of Hispaniola only one lineage has become widespread. Male Traits.—Changes in secondary sexual characters of Our study is clearly not definitive. Outstanding taxo- males that have taken place independently in the opaca nomic issues about the nitidipennis group in Mexico con- and nitidipennis groups are: middle tibia of male becom- cern monophyly of the soladevega subgroup, and whether ing as slender as in female (character 27); and replacement E. zempoaltepetl, E. carbonera, E. miahuatlan, E. suchix- of the medial ridge of the hind tibia by a row of tubercles tepec, and E. santacatarina belong in the nitidipennis sub-

group instead of the dimidiata subgroup. Our new knowl- duction to town councils that were of substantial importance in obtaining edge of the larval stages of E. soladevega, etc., indicates the approval necessary to obtain access to those mountains. A foray on the great Volcan Citlaltepetl, in 1975, was rendered successful primarily that obtaining larvae of some of the species in question, due to Eutiquío Vite M., farmer and resident of Cuiyachapa, Veracruz, and also of E. flohri and E. nitidipennis, could prove use- who befriended us, and during his service as guide and protector, took us ful in resolving both problems. The main issue about the to sites productive for collecting Dyschromus specimens. opaca group on the island of Hispaniola concerns the We are grateful to Patrice Stephens-Bourgeault of the Royal Ontario monophyly of the opaca subgroup. Thus far, results of Museum for providing the superb paintings, done in pastels, showing the metallic colors of some of these beetles, and for sharing with the senior mDNA sequencing have been disappointing in terms author, her technical skills in the art of computer scanning and editing of of further establishing relationships between all but the illustrations and photographs, and for doing the task herself, when the most closely related species of Dyschromus, but this does senior author fell short of the mark. B. Boyle, photographer at the Royal not preclude sequencing other regions of the mDNA Ontario Museum, made 35 mm color transparencies of the paintings, which were then digitized for publication by Ms. Stephens-Bourgeault. genome. The large gaps in the geographical range of some Most of the scanning electron photomicrographs were taken by F. Neub pairs of species, such as E. juchatengo and E. nizavaguiti, of the Department of Metallurgy and Materials Science, University of and E. perezi and E. pedernales, shows that more collect- Toronto; the others were taken by G. Braybrook of the University of ing is necessary to fully elucidate how species of the sub- Alberta. J.S. Scott prepared the base map of Mexico, copies of which we genus Dyschromus are distributed in central Mexico used in illustrating the geographical ranges of the species of the nitidipennis group. and on the island of Hispaniola. The doubtful status of cer- We acknowledge with appreciation J. Rawlins, T. Moore and other tain individuals of E. perezi, E. pedernales, E. cuiyachapa, editorial staff at the Carnegie Museum of Natural History for their E. ixtapa, E. dimidiata, and E. puertogallo from some encouragement and considerable practical assistance in producing the localities indicates that we only dimly understand the lim- final version of this monograph.. For use of facilities and resources in the Entomology section of the its of these species, and also that we still do not know the Department of Natural History at the Royal Ontario Museum, the senior actual number of species that exist in this subgenus. author thanks G.B. Wiggins, D.C. Darling, and D.C. Currie. Financial support for this study was provided to the junior author initially by grant GB 3312, from the National Science Foundation, and then by grant OGP ACKNOWLEDGMENTS 1399 from the Natural Sciences and Engineering Research Council of Canada. The latter organization also provided support to the junior author for preparation of the color illustrations, and for publication costs. The authors are indebted to the curators and other individuals noted above in the “Material” section of the paper, for loan of the specimens in their care that were vital to this study. Special thanks to K. Desender LITERATURE CITED (IRSNB), N. Stork and M.J.D. Brendell (BMNH) for loaning certain specimens a second time. As well, the junior author is pleased to ARNDT, E. 1993. Phylogenetische Untersuchungen larvalmorphologisch- acknowledge the cooperation and assistance he received from curators er Merkmale der Carabidae (Insecta: Coleoptera). Stuttgarter and other staff members, when studying types in the Museum National Beitraege zur Naturkunde Serie A (Biologie), No. 488:1–56. d'Histoire Naturelle (Paris) and The Natural History Museum (London). Stuttgart, Germany. Special thanks to J. Rawlins and R. Davidson (CMNH) for the loan AUDOUIN, J.V., AND A. BRULLÉ. 1834. Histoire naturelle des Insectes, trai- of much recently collected material from the Dominican Republic that tant de leur organization et de leurs moeurs en général, et com- markedly increased our knowledge about the subgenus Dyschromus in prenant leur classification et la description des espèces. Volume 4, that country. We also thank Y. Bousquet (CNCI) for comparing some lar- Coléoptères. I.D. Pillot, Paris. vae of Dyschromus with those of Abaris. AXELROD, D.I. 1975. Evolution and biogeography of Madrean-Tethyan The junior author is mindful of the interest in the Mexican carabid sclerophyll vegetation. Annals of the Missouri Botanical Garden, fauna that was stirred when, as an undergraduate more than 50 years ago, 62:280–334. he received from C. Richard Robbins (then, a budding ornithologist, later BALL, G.E. 1959. 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APPENDIX 1. Description and distribution within the subgenus Dyschromus of five extreme types of median lobe of the aedeagus, and intermediate conditions.

As noted in the section on morphology, based on form of the preapex, five distinct types of median lobe are discernable in the subgenus Dyschromus, one of which is characteristic of the subgenus Euchroa. The other four types are confined to the subgenus Dyschromus. These five types appear to linked by intermediate types. Several transformation series are proposed, but there is con- flicting evidence about the exact pathway to each of the extreme types (Fig. 63). See section on phylogeny and discussion about character trends for further details. Generalized asymmetric type (Figs. 18, 20). This kind of median lobe is typical of the subgenus Euchroa, and also a few species belonging to the subgenus Dyschromus. It is characterized by the combination of: preapex short, broad and slightly asymmetrical in outline (Fig. 18a), right third somewhat thicker and more darkly sclerotized than rest of preapex (Fig. 18b), preapical dorsolateral ridge absent (Fig. 18b), at least three primary preapical haemolymph channels present (Fig. 18b), largest channel with origin at far right side of ostium, three or four progressively smaller channels arising at intervals to left of largest one; left wall of periostial part of median lobe as extensive or slightly more so than right wall, so neither right wall, nor ostium visible in left lateral view (as in Fig. 19b), periostial ventrolateral bulge not evident (Fig. 20b) or if present, then comparatively basal in position, and only slightly prominent profile (as in Fig. 19b) and subangulate in cross-section (as in Fig. 26c); middle part of median lobe without dorsomedial groove. E. tiburonica and E. centralis. Also, E. opaca and E. independencia, except preapex with just two primary preapical haemolymph channels, a longer one with origin at far right side of ostium; and a broader one arising to left of midline (Figs. 19c, 22c), and E. opaca with slightly developed preapical dorsolateral ridge (Fig. 19a), and moderately prominent periostial ventrolateral bulge (Fig. 19b). Extreme asymmetric type 1 (Figs. 22f–g, 24). This type of median lobe is uniquely characterized by the preapex, the middle third of which is very thin and colorless (Figs. 22f, 24c). Description: preapex long, right third considerably thicker and more darkly sclerotized than rest of preapex, most of middle third of preapex very thin, even compared to left side of preapex, and colorless (Fig. 22g), preapical dorsolateral ridge evident, more or less flat in profile (Figs. 22f, 24b), two primary haemolymph channels present, larger on right; left wall of periostial part of median lobe at least as extensive as right wall, so neither right wall, nor ostium visible in left lateral view (Fig. 24b), periostial ventrolateral bulge comparatively basal in position, prominent in profile (Fig. 24b) and subangulate in cross-section (as in Fig. 26c); middle part of median lobe without dorsomedial groove. E. perezi and E. pedernales. Intermediate asymmetric type 1 (Fig. 22d–e). As in extreme asymmetric type 1, except preapex short, and very thin and colorless portion of middle third of preapex small in extent, preapical dorsolateral ridge only slightly developed (Fig. 22d–e). E. cupripennis. Extreme asymmetric type 2 (Figs. 34, 35, 49, 50). This kind of median lobe is recognizable by the combination of: preapex developed only to right of midline of median lobe, and stout and darkly sclerotized throughout (Figs. 34a, 34c), preapical dorsolateral ridge markedly convex in profile (Fig. 34b); periostial ventrolateral bulge comparatively distal in position, prominent in profile (Fig. 34b) and rounded in cross-section (Fig. 26b). The combination of convex preapical dorsolateral ridge, and prominent periostial ventrolateral bulge imparts a distinctly sinuous aspect to the distal part of the median lobe when viewed from the side (Fig. 34b). Description: preapex long, developed only to right of midline of median lobe, stout and darkly sclerotized throughout (Figs. 34a, 34c), preapical dorsolateral ridge prominent and markedly convex in profile (Fig. 34b), single primary preapical haemolymph channel present, with origin at far right side of ostium (Fig. 34c); left wall of periostial part of median lobe not as extensive as right wall, so right wall and part of ostium visible in left lateral view (Fig. 34b), periostial ventrolateral bulge comparatively apical in position, prominent in profile (Fig. 34b) and rounded in cross-section (Fig. 26b); dorsomedial groove extending length of central part of median lobe (Fig. 34a). E. perote, E. harrisoni, E. miahuatlan and E. suchixtepec. Intermediate asymmetric type 2 (Figs. 33, 48). As in extreme asymmetric type 2 except preapex extending to the left of midline of the median lobe, portion to left thin and colorless. E. citlaltepetl, E. zempoaltepetl and E. carbonera. Eight other species also have an intermediate asymmetric type 2 median lobe. Of these, the median lobe of E. nitidipennis (Figs. 25a–c, 26a) differs from the generalized asymmetric type only in that the preapex is longer, the preapical dorsolateral ridge is more developed, and the periostial ventrolateral bulge is prominent, more distal in position, and rounded in cross-section. The other seven species—E. sallei, E. cuiyachapa, E. zongolica, E. lasvigas, E. flohri, E. teotitlan and E. soladevega—have median lobes that col- lectively bridge the morphological gap between the median lobe of E. nitidipennis and that of E. citlatltepetl, E. perote, etc. (Figs. 25d–f, 28, 30, 32) or appear to be variants of an asymmetric type 2 median lobe (Figs. 29a–f, 36a–c). Extreme asymmetric type 3 (Fig. 47d–f). This kind of median lobe has the features of extreme asymmetric type 2, except that the preapical dorsolateral ridge is rather flat in profile, and the periostial ventrolateral bulge is only slightly developed, so that the preapex and adjoining portion of the periostial part of the median lobe present a linear rather than sinuous aspect when viewed from the side (Fig. 47e). Another difference is that the periostial ventrolateral bulge is subangulate in cross-section (as in Fig. 26c). E. santacatarina. Intermediate asymmetric type 3 (Figs. 45d–f, 47a–c). As in extreme asymmetric type 3, except preapex developed to left of midline of median lobe, left portion thinner and not as darkly sclerotized as area to right, two primary preapical haemolymph channels present, right one longer, left one broader; left wall of periostial part of median lobe more extensively developed than the right wall; middle part of median lobe with dorsomedial groove short or long (Figs. 45d, 47a). E. chrysophana, E. yucuyacua.

APPENDIX 1 CONT.

Symmetric type 4 (Figs. 36d–g, 38, 39, 40, 43, 44, 45a–c). This kind of median lobe is characterized by greater symmetry of the preapex than other types because the preapex is subrectangular in outline, and is more or less moderately thick and darkly sclerotized throughout, and has just one large, centrally located primary preapical haemolymph channel (Figs. 38b, 40c). Description: preapex long or moderately so, subrectangular, more or less moderately thick and darkly sclerotized throughout (Figs. 26c, 40a, 40c), no preapical dorsolateral ridge, one large, centrally located primary preapical haemolymph channel present (Figs. 36e, 38b, 40c); left wall of periostial part of median lobe about as extensive as right wall (Fig. 40b), periostial ventrolateral bulge comparatively distal in position, moderately prominent in profile (Fig. 40b) and subangulate in cross-section (Fig. 26c); dorsomedial groove of middle part of median lobe present (Figs. 36d, 36f) or absent (Fig. 40a). E. juchatengo, E. nizavaguiti, E. jalisco, E. tenancingo, E. ixtapa, E. dimidiata, E. atoyac, E. puertogallo, E. filodecaballo. In some species with a symmetric type 4 median lobe the preapex is longer than in other species (cf. Figs. 36d–g), which appears to provide the only basis for recognition of an intermediate as opposed to extreme condition for this type of median lobe.

APPENDIX 2. Adult characters of the subgenus Dyschromus.

1. Body size. Various euchroines examined, including all members of the subgenus Euchroa, are large in size, and so attainment of a large body size is considered to be a plesiomorphic trait in the subgenus Dyschromus. As only a few individuals of a few species of Dyschromus actually attain a size comparable to the smallest specimens of the subgenus Euchroa examined, each species was scored based on the apparent body length of the largest individual, as follows: a) largest specimen with apparent body length 13 mm; b) 12.5 mm; c) 11.5 mm; d) 9.5 mm. 2. Body color. Besides two species of the subgenus Dyschromus, some other New World euchroines examined are all black, e.g., E. (Euchroa) okonagare, but many others are metallic colored dorsally, the head and pronotum of many species having a different color than the elytra. Although the all black condition is undoubtedly plesiomorphic for Pterostichini and perhaps carabid beetles in general, because so many euchroines are metallic colored, we were not confident that uniform black is the plesiomorphic condition for this trait in Dyschromus, nor was there much basis for proposing a detailed transformation series. We did think it reasonable that the bicolored condition is apomorphic, and initially recognized a two state transformation series: a) dorsum uniform in color; b) dorsum bicolored, head and pronotum differently colored than elytra. Based on mental analysis using the above transformation series, the character was resolved further as: a) dorsum all black or metallic purple; b) dorsum metallic green or brassy; c) head and pronotum green or coppery, elytra brassy or bronze; Da) head and pronotum purple or coppery, elytra reddish. This coding was used in the compatibility analysis, but parsimony analyses were conducted with all unordered character states, as follows: a) dorsum all black; b) dorsum metallic purple; c) dorsum metallic green; d) dorsum brassy; e) head and pronotum metallic green, elytra coppery or brassy; f) head and pronotum coppery, elytra bronze; g) head and pronotum metallic purple, elytra reddish-purple; h) head and pronotum coppery, elytra reddish-purple. 3–6. Microsculpture on disc and sides of elytra. The microsculpture on the disc of the elytra of many euchroines examined, including the subgenus Euchroa, consists of sculpticells that are markedly transverse—about 2–3 times wider than long, rather finely impressed, and flat. The microsculpture at the sides (outer half of interval 8, all of interval 9) and apex of the elytra does not differ from that on the disc, except that the sculpticells tend to be still more stretched and oriented lengthwise. Species of the subgenus Dyschromus show considerably more interspecific variation for elytral microsculpture, and at least the sculpticells at the sides and apex of the elytra are more deeply impressed, and in most species at least at the apex of the elytra some of the sculpticells are convex so that the surface of the elytra is beaded (morphology section). We recognize four transformation series in Dyschromus having to do with elytral microsculpture. 3. Shape of sculpticells on disc of elytra: a) sculpticells isodiametric to slightly transverse; b) somewhat elongate. 4. Prominence of sculpticells on disc of elytra: a) sculpticells moderately finely impressed, flat; b) moderately deeply impressed, flat; c) deeply impressed, convex or keeled; Da) very finely impressed, effaced in spots, flat; e) effaced, elytral intervals smooth. 5. Shape of sculpticells along sides and at apex of elytra: a) sculpticells isodiametric to slightly elongate; b) very elongate. 6. Convexity of sculpticells along sides and at apex of elytra: a) sculpticells flat; b) apex of elytra with some patches of distinctly convex sculpticells, some portion of interval 9 with sculpticells convex or not; c) outer half of interval 8, all of interval 9, and apex of elytra with sculpticells convex. 7. Dorsal mandibular grooves. Long dorsal mandibular grooves are characteristic of Haplobothynus and the subgenus Euchroa, as well as many species of the subgenus Dyschromus, but are lacking from the mandibles of most other euchroines examined, or are relatively short, as in some species of Dyschromus and most other pterostichines. Provisionally, we consider the transformation series for the dorsal mandibular grooves in Dyschromus to be: a) many, long; b) few, short, in some or all individuals. 8–9. Width of labial palpomere 3. Labial palpomere 3 is very broad in Microcephalus and the subgenus Euchroa, especially in males. Among males, palpomere 3 is narrowest in the undescribed species of Euchroa referred to in the diagnosis of the subgenus Dyschromus. As for the subgenus Dyschromus, palpomere 3 of males of just a few species is as broad as in the above undescribed species of Euchroa; there is more overlap for this trait in females of the two subgenera. We recognize for each sex a single transformation series in Dyschromus leading to the comparatively slender condition seen in E. opaca and a few other species of Dyschromus. [N.B. Categories were established using for each species the median value of the ratio: length/width of labial palpomere 3. The median value was used because the overlap between species for this ratio was too great for us to be able to use the range of values. In Dyschromus, the length of labial palpomere 3 varies as well as the width, and is more difficult to measure, hence the wide range in values for the above ratio. A more suitable ratio appears to be: width of labial palpomere 3/ length of pedicel of antenna, but we found this out too late in our study to do all of the neccesary measurements.] 8. Width in males of labial palpomere 3: a) extremely broad, 0.92–0.94 times longer than wide (median values); b) very broad, 1.0–1.12 times longer than wide; c) broad, 1.16–1.41 times longer than wide; d) somewhat broad, 1.47–1.94 times longer than wide; e) comparatively slender, 2.0–2.13 times longer than wide. 9. Width in females of labial palpomere 3: a) broad, 1.29–1.48 times longer than wide (median values); b) somewhat broad, 1.56–1.96 times longer than wide; c) comparatively slender, 2.0–2.19 times longer than wide; d) quite slender, 2.25–2.53 times longer than wide.

APPENDIX 2 CONT.

10. Frontal furrows. Among euchroines examined, besides certain species of the subgenus Dyschromus, only species belonging to the subgenus Euchroa, and also Setalimorphus, have the frontal furrows deeply and sharply impressed for their entire length and extended posteriorly at least as far as the anterior supraorbital setigerous punctures. The frontal furrows are shallowly impressed in most other pterostichines and the sharply impressed portion (if any) does not attain the level of the anterior supraorbital punctures. Provisionally, based on the condition in the subgenus Euchroa, we consider the long, and deeply and sharply impressed condition to be plesiomorphic in Dyschromus, so the character transformation series for the frontal furrows is: a) deeply and sharply impressed, extended posteriorly at least to level of anterior supraorbital setigerous punctures. b) shallow, sharply impressed portion (if any) not extended to anterior supraorbital punctures. 11. Dorsal postocular transverse groove. The postocular transverse groove is distinctly developed dorsomedially in all euchroines examined, except for Lobobrachus, and some species of the subgenus Dyschromus. We consider the transformation series for this trait to be: a) postocular transverse groove nearly as deep dorsomedially as laterally; b) some or all individuals with postocular transverse groove obsolete dorsomedially. 12. Paramedian pits of mentum. The paramedian pits of the mentum are large in Bothynoproctus, and most species of the subgenus Euchroa, but are of moderate size in the undescribed species of Euchroa referred to for characters 8–9, and are small in Microcephalus. Each of the three states is also exhibited by two or more species of the subgenus Dyschromus. We recognize one transformation series for size of the paramedian pits, with the first condition as ancestral: a) large; b) moderate in size; c) small. 13. Submentum and transgular groove. The submentum of nearly all of the New World Euchroina examined is not prominent, the posterior face sloped gently and evenly to the transgular groove, which is shallow posterior to the submentum. Within the subgenus Dyschromus an apparent transformation series exists from the above condition to one in which the submentum is very prominent and concave posteriorly, and the adjoining part of the transgular groove is very deep. The first condition is usual in Carabidae, including the subgenus Euchroa, so interpretation of this trait is straightforward: a) submentum not prominent, posterior face sloped evenly to transgular groove, latter shallow posterad submentum; b) submentum slightly prominent, posteri- or face sloped evenly to transgular groove, latter moderately deep posterad submentum; c) submentum prominent, posterior face sloped evenly to transgular groove, latter deep posterad submentum; d) submentum very prominent, posterior face concave, transgular groove very deep posterad submentum. 14–16. Pronotum. Most euchroines examined, including the subgenus Euchroa, have in common several features of the pronotum: the sides are entire posteriorly, the hind angles are oblique or rounded, and the inner pair of basolateral impressions are long and attain the hind margins of the pronotum. We describe the variation shown by the subgenus Dyschromus for these traits as three separate character transformation series. 14. Sides of pronotum: a) entire posteriorly, or at most very slightly sinuate at very base in some individuals; b) posterior third distinctly sinuate. 15. Hind angles of pronotum: a) rounded or entire; b) square or acute. 16. Inner pair of basolateral impressions of pronotum: a) at least moderately long; b) short to punctiform. 17. Prosternal intercoxal process. Among euchroines examined, the entire apical third of the prosternal intercoxal process is margined only in some species of both subgenera of Euchroa, and also Setalimorphus, and just at the apex in Bothynoproctus. Tentatively, we designate the fully margined condition as plesiomorphic in the subgenus Dyschromus, and so the character state tree for the prosternal intercoxal process is: a) entire apical third margined; b) process only margined laterally in at least some individuals; c) process not margined in at least some individuals. 18. Shoulders of elytra. In all Euchroina examined, except for some species of the subgenus Dyschromus, the shoulders of the elytra are relatively prominent, and at least Haplobothynus exhibits a humeral tooth. On this basis, the transformation series in Dyschromus for the shoulders of the elytra is: a) shoulders at least somewhat prominent, humeral tooth evident or not; b) entirely rounded, humeral tooth not evident. 19. Elytral basal punctures. Presence of the elytral basal punctures is the usual condition in most species of the subgenus Euchroa and related genera, but these punctures are absent in a few species of the subgenus Dyschromus. Accordingly, the transformation series for the elytral punctures is: a) punctures present; b) absent. 20. Elytral striae. Species belonging to the subgenus Euchroa, related genera, and also some species belonging to the subgenus Dyschromus, have all of the stria intact on the disc of the elytra, but many species of Dyschromus have one or more striae interrupted on the disc. The latter is undoubtedly the derived condition for Dyschromus, and so the character transformation series for breakup of the striae is: a) disc of elytra with all striae entire; b) some specimens with stria 1 interrupted up to 2–3 times on disc, a few of these specimens with stria 2 interrupted once or twice; c) some specimens with stria 1 interrupted up to 5 times on disc, 2 and 3 interrupted up to 3 times, 4 and 5 interrupted once in some specimens; d) striae 1–2, 1–3, 1–4, 1–5, 1–6 or 1–7 each interrupted on disc, 1 interrupted 24–32, 2 interrupted 3–28, 3–4 interrupted 0–19, 5 interrupted 0–12, 6 interrupted 0–10, 7 interrupted 0–1 times; e) striae 1–7 each interrupted at least 20 times on disc.

APPENDIX 2 CONT.

21–22. Convexity of elytral intervals. Most euchroines examined have at least moderately convex elytral intervals, but in the subgenus Dyschromus the intervals tend to be flatter, especially on the disc of the elytra. The condition for this trait on the disc is in many species different from that posterolaterally at and in the vicinity of the junction of stria 5 and 6, and so we recognize two separate characters. 21. Convexity of intervals 1–6 on disc of elytra: a) moderately convex; b) somewhat convex; c) somewhat flat; d) nearly flat; e) flat. 22. Convexity of intervals 5–7 posterolaterally on elytra: a) moderately convex (Fig. 15a); b) somewhat to slightly convex; c) somewhat to nearly flat (Fig. 15b); d) flat. 23. Apical declivity of elytra. The apical declivity of the elytra is relatively gentle to moderately steep in all euchroines examined, except for a few species of the subgenus Dyschromus that have the apical declivity nearly perpendicular. Interpretation of the charcter state tree for slope of the apical declivity is straightforward: a) declivity comparatively gentle; b) nearly perpendicular in some or all specimens. 24. Punctation of mesepisternum. The extent to which the mesepisternum is punctate varies considerably among euchroines examined, including the subgenus Euchroa. Our designation of the plesiomorphic condition in the subgenus Dyschromus for mesepisternal punctation is arbitrary: a) punctate; b) smooth in some specimens; c) smooth. 25. Extent of basal sulcus of abdominal sterna V–VII. The basal sulcus of abdominal sterna V–VII does not extend to the lateral margins of the abdomen in the subgenus Euchroa and most species of Dyschromus. It attains the lateral margins in other euchroine genera such as Bothynoproctus and Microcephalus, and also in E. tenancingo. Based upon the condition in the subgenus Euchroa, we consider the latter condition to be apomorphic in Dyschromus, and the character state tree for the basal sulci to be: a) not attaining lateral margins; b) extended to lateral margins. 26. Punctation of basal sulcus of abdominal sternum VII. In Bothynoproctus, the subgenus Euchroa, and some species of Microcephalus and of the subgenus Dyschromus, the basal sulcus of abdominal sternum VII has in addition to lateral foveae, a coarse median fovea. In a few other species of Dyschromus the entire sulcus is coarsely foveate, but in most species the basal sulcus of sternum VII is only foveate laterally, although in some specimens there is a trace of a median fovea. We consider the presence of a coarse median fovea to be plesiomorphic, and the character transformation series to be: a) basal sulcus of abdominal sternum VII with coarse median fovea, in addition to coarse lateral foveae; b) median foveae present in most specimens, but very shallow; c) no median fovea (or rarely evident); Da) entire sulcus coarsely foveate. 27. Middle tibia of male, expansion of inner face. Males of some species of the subgenus Euchroa have the inner face of the middle tibia distinctly expanded near the apex, others do not. Tentatively, we consider the expanded condition to be plesiomorphic in the subgenus Dyschromus, and so the transformation series is: a) inner face distinctly expanded medially near apex; b) not expanded in at least some males. 28–29. Hind tibia, medial ridge and medial row of flexile setae. With the exception of some species of the subgenus Dyschromus, all euchroines examined have on the hind tibia a single row of medial flexile setae that is bordered by a slight medial ridge. Males of certain species of Dyschromus have a row of tubercles instead of a ridge, and males of other species exhibit an intermediate condition. Both sexes of some species of Dyschromus have two rows of medial flexile setae. We consider it best to treat the latter condition as a separate character, and take the usual expression of each trait in euchroines to be plesimorphic. 28. Hind tibia of males, medial ridge: a) medial row of flexile setae bordered by medial ridge in both sexes; b) basal portion of medial ridge represented by few, slight subangular tubercles in males; c) as in condition “b” except tubercles more pronounced; d) medial ridge, in males, consisting entirely (or nearly so) of line of subangular to subrectangular tubercles, some of which are prominent. 29. Hind tibia, medial row of flexile setae: a) single medial row of 6–16 flexile setae; b) two medial rows of flexile setae, outer one with about 10 setae, inner with 3–5 setae, arising from common groove. 30–40. Distal portion of median lobe of aedeagus. As far as we are aware, euchroines other than the subgenus Dyschromus exhibit comparatively little interspecific variation for traits of the distal part of the median lobe. In contrast, Dyschromus exhibits so much variation interspecifically, that as discussed in Appendix 1, we have found it useful to recognize five distinct types of median lobe. We have not employed these categories as characters in our numerical phylogenetic analyses because it was not obvious to us how to a priori arrange them into a single morphocline, and also the median lobe of various species could not be exactly fitted into these categories. Instead, as each type of median lobe is characterized by a combination of traits, the individual traits were used in the numerical analyses. We recognize eight separate characters having to do with the structure of the median lobe of Dyschromus, and take the usual condition of each trait in euchroines that we examined to be plesiomorphic. 30. Length of preapex of median lobe: a) preapex distinctly shorter than ostium, i.e. not more than 0.75 times length of ostium (Fig. 18a); b) moderately long, about same length as ostium (Figs. 24a, 25a); c) long, at least 1.25 times longer than ostium (Fig. 25d). 31. Condition of apicomedial area of preapex of median lobe: a) apicomedial area not unusually thin or less darkly sclerotized than area to left (Fig. 22c); b) very thin and more or less transparent, but small in extent (Fig. 22e); c) very thin, more or less transparent, and large in extent (Fig. 22g).

APPENDIX 2 CONT.

32. Development of left side of preapex of median lobe: a) left margin not unusually thin or less darkly sclerotized than adjoining area (Fig. 25c); b) left margin very thin and more or less transparent near apex (Fig. 25f); c) entire left margin very thin and more or less transparent (Figs. 30c, 32c); d) entire area to left of very stout portion of preapex very thin, more or less transparent, and in some specimens somewhat reduced in extent (Figs. 29c, 33c, 48c); e) preapex only developed to right of either midline or dorsomedial groove of median lobe, stout and darkly sclerotized throughout (Figs. 29c, 34c). 33. Development of right side of preapex of median lobe: a) right third of preapex somewhat stouter and more darkly sclerotized than rest of preapex (Fig. 18b); b) right third to right half of preapex much stouter and darkly sclerotized than rest of preapex (Fig. 25c) or preapex only developed to right of midline of median lobe (Fig. 34c); c) most or all of preapex very stout, preapex quite broad at base (Figs. 29c, 29f); Da) preapex broad, moderately thick and moderately darkly sclerotized throughout (Figs. 36e, 36g). 34. Preapical dorsolateral ridge of median lobe: a) median lobe with extreme right side not developed dorsally as a ridge (Figs. 18a, 26c); b) preapical dorsolateral ridge flat (Figs. 24a–b, 45d–e); c) ridge convex (Figs. 25b, 26a–b, 29b). 35. Primary preapical haemolymph channels of median lobe: a) preapex with at least four primary haemolymph channels, largest channel with origin at extreme right side of ostium, three or four progressively smaller channels originating at intervals to left of largest channel (Fig. 18b); b) two primary haemolymph channels, longest with origin at extreme right side of ostium, broadest to left of midline (Figs. 19c, 25c, 25f); c) one primary haemolymph channel, with origin at extreme right side of ostium (Fig. 28c); Db) one centrally located primary haemolymph channel (Fig. 36e). 36. Presence of bulla along left margin of median lobe at base of preapex: a) left margin of median lobe unmodified at base of preapex (Fig. 48e); b) with ventrally directed bulla (Fig. 49b). 37. Symmetry of periostial part of median lobe: a) periostial part of median lobe with left wall at least as developed as right wall, entirely concealing right wall and ostium in left lateral view (Figs. 29b, 32e); b) left wall not so extensive as right wall, portion of right wall and ostium visible in left lateral view (Figs. 33b, 34b). 38. Periostial ventrolateral bulge of median lobe: a) bulge absent, or slightly developed and comparatively basal in position (Fig. 22b), subangulate in cross-section (as in Fig. 26c); b) bulge prominent in profile (Fig. 24b), otherwise as in condition “a”; Ca) bulge comparatively apical in position, inconspicuous to somewhat prominent in profile, (Figs. 40b, 43b), subangulate in cross-section (Fig. 26c); d) bulge somewhat to markedly prominent in profile (Figs. 25b, 25e, 29b), rounded in cross-section (Fig. 26a–b),otherwise as in condition “Ca.” 39. Thickness of middle part of median lobe: a) middle part of median lobe distad prebasal bend comparatively slender in lateral aspect (Figs. 44b, 44e); b) middle part stout (Figs. 45b, 45e). 40. Dorsomedial groove of middle part of median lobe: a) absent; b) groove short, extended apically only short distance beyond prebasal bend (Fig. 45a); c) groove extended entire length of middle part of median lobe (Fig. 47a). 41. Microtrichia of internal sac of aedeagus. We were unable to evert the internal sac of any euchroines other than the subgenus Dyschromus, but removed from the median lobe, the uneverted sac of a specimen of the undescribed species of the subgenus Euchroa referred to in the diagnosis of the subgenus Dyschromus, and examined it in that state. The internal sac of Dyschromus has either one narrow band of large microtrichia extended around the base of the sac, or one much broader band, or one subapical and often one basal band, one or both of which may be incomplete. The undescribed species of the subgenus Euchroa which is referred to above, appears to have a single broad band, and so we consider the character state tree for arrangement of the large microtrichia of the internal sac to be: a) extended in one broad band around middle of sac (Fig. 24f); b) one narrow band around base of sac (Fig. 19d); Ca) one narrow subapical, and also, in many species, one basal band (Fig. 41a). 42. Female, shape of spermatheca. The apical portion of the spermatheca is long and straight in most species of the subgenus Euchroa, and in Meropalpus and Setalis, short and straight in a few species of the subgenus Dyschromus, and long and recurved in the other species of Dyschromus, and also in E. (Euchroa) onkonegare. We tentatively designate the latter condition as ancestral, and so the transformation series for spermathecal shape is: a) long, apical portion recurved (Fig. 17b); b) short, apical portion straight (Fig. 17a).

APPENDIX 3. Larval characters of the subgenus Dyschromus.

1. Frontale seta 4. Seta FR4 is minute in at least the first instar of several species of Dyschromus (Fig. 52b); otherwise it is short (Fig. 52a). For each instar we consider the minute condition to be the derived state because this seta is shown to be short rather than minute in all illustrations of the pterostichine larvae described by Bousquet (1984, 1985, 1989), and in particular, Abaris bigenera Bates (Bousquet and Liebherr 1994), and also a drawing of a generalized first instar carabid larva showing the ancestral setae and pores (Bousquet and Goulet 1984, . 1). We propose on this basis that the transformation series for seta FR4 in larvae of the subgenus Dyschromus is: a) first instar with seta FR4 short or minute, second and third instars with FR4 short; b) first and second instars with FR4 minute, third instars with FR4 short; c) all instars with FR4 minute. This is contrary to the notion that derived traits first appear in the later instars (see text concerning phylogenetic analysis of larval data). We consider the evidence from outgroup comparison to be decisive in our study. 2. Some parietal, notal, and tergal primary setae. At least first instar larvae of a few species of Dyschromus have a number of minute primary setae that are long in all instars of other species for which the larva is known, including parietal seta 6 (Fig. 52g), pronotal setae 3 and 12, mesonotal and metanotal seta 13, and tergal seta 10. These setae are long in all illustrations of pterostichine and ancestral carabid larvae cited above for larval character 1 (as in Fig. 51a–b). Hence the evolutionary sequence for the above setae is: a) all instars with parietal seta 6 long, as well as pronotal setae 3 and 12, mesonotal and metanotal seta 13, and tergal seta 10; b) all instars with these setae minute. 3. Mesonotal and metanotal seta beta. In second and third instars of several species of Dyschromus mesonotal and metanotal seta beta is long or nearly so (Figs. 51a, 53c); in some others seta beta is short, minute or not evident (Fig. 53b). This seta is well- developed on the mesonotum and metanotum of Abaris bigenera (Y. Bousquet, pers. com.). We consider the character transformation series to be: a) second and third instars with mesonotal and metanotal seta beta long; b) seta beta short, minute or not evident; c) seta beta not evident. 4. Urogomphi, subprimary setae. Second and third instars of some species of Dyschromus have on the urogomphi four subprimary setae: beta, gamma, delta, and epsilon, all of which are all long (Fig. 51c). In other species one or more of these setae are short, minute or absent. The number of subprimary setae on the urogomphi varies considerably among larvae of Pterostichini (illustrations cited for larval character 1), but the larva of Abaris bigenera has each of the above four subprimary setae, and they are long (Bousquet and Liebherr 1994: Fig. 4). Accordingly, the transformation series for the subprimary setae of the urogomphi in Dyschromus is: a) urogomphi of second and third instars with four subprimary setae, i.e. beta, gamma, delta, and epsilon, all four setae long; b) third instar with two subprimary setae, i.e., gamma and epsilon, second instar just with subprimary seta epsilon, both setae long; c) second and third instars with condition “b” except third instar with seta gamma long, short, or minute; d) second and third instars just with subprimary seta epsilon, this seta long. 5. Number of stemmata. Larvae of some species of Dyschromus have five stemmata on each side of the head (Fig. 52g), larvae of the others have six (Figs. 52e–f). Larvae of most other pterostichines have six and this is presumably the ancestral condition for larvae of Carabidae (illustrations cited above for larval character 1). We conclude that presence of five stemmata in larvae of Dyschromus is a reduction from the ancestral condition, the character state tree being: a) each side of head with six stemmata; b) five stemmata. 6. Lobe formed by longitudinal sulcus on dorsum of head. Third instar larvae of the subgenus Dyschromus have a longitudinal sulcus that forms a lobe over the eye. In third instars of some species this lobe is quite pronounced and the posterior margin has a sharp edge (Figs. 52c–d, 52f); in third instars of the other species it not as pronounced and has a rounded edge (Figs. 52e, 52g). This lobe is even less pronounced in second instar larvae of Dyschromus, and only in some individuals of E. dimidiata does it have a sharp edge. First instar larvae do not have this sulcus. Neither do most other pterostichine larvae (illustrations cited for larval character 1). We infer that presence of a longitudinal sulcus is apomorphic, and also propose that the sharp-edged condition of the lobe formed by this sulcus is more derived than the rounded condition, but only because the lobe is more pronounced when it has a sharp edge. The transformation series for the condition of the lobe formed by the longitudinal sulcus in larvae of Dyschromus is: a) second and third instars with lobe formed by longitudinal sulcus somewhat prominent, rounded posteriorly; b) third instar with lobe formed by longitudinal sulcus very prominent, posterior margin sharp; c) second instar also with condition “b.” 7. Mesonotal and metanotal carina. The mesonotal and metanotal carina extends posteriorly to seta 11 in all known first instar larvae of the subgenus Dyschromus, and also in the second and third instars of some species (Fig. 53b). In second and third instars of other species this carina extends to seta 12, at least on the mesonotum (Fig. 53c). We lack data for other pterostichine larvae, and arbitrarily coded the transformation series for the condition of the mesonotal and metanotal carina as: a) second and third instars with mesonotal and metanotal carina extended to seta 11; b) second instar with mesonotal carina extended to seta 12, third instar with both mesonotal and metanotal carina extended to seta 12; c) second and third instar with mesonotal and metanotal carina extended to seta 12.

APPENDIX 4. Details about options used when doing phylogenetic analyses by computer.

1) Parsimony analyses using the computer programs within WINCLADA. Searches for islands of minimum length trees were done by choosing the RATCHET program from the “Analyze” menu, and submitting the trees obtained to the NONA program for branch swapping. Within the RATCHET program the number of iterations was set to 200, 1 tree held per iteration, and 3 characters were sampled (initially weighted). NONA was run with the hold option set to 10,000, and by invoking the “best” and “max” commands. Successive approximations character weighting, when found to be necessary, was done as implemented within NONA by invoking the command swt.run mult*10. When sucessive approximations character weighting was also done as implemented in HENNIG86, this was accomplished by submitting the trees found by NONA to HENNIG86 and repeating the command sequence, xs w, m*, bb* until there was no further change in weights and tree length. 2) Compatibility analyses using CLINCH. Compatibilty analyses were run with the options HASS and LARG.

APPENDIX 5.

The following is a list of the 12 most distinctive species of the subgenus Dyschromus, as indicated by the total number of character state changes separating it from its sister taxon, whether a particular species, or the common ancestor of a group of species, in Figs. 58–60. Branch lengths were obtained by plotting the character state changes on the cladogram using the Winclada computer program, first with the “fast” (Acctran) and then the “slow” (Deltran) option. For each species, the mean and range for the above statistic is given: E. yucuyacua, 21(18–23); E. tiburonica 17.6(14–21); E. santcatarina 16(13–22); E. soladevega 15.1(12–17); E. centralis 15(14–16); E. dimidiata 15(11–23); E. opaca 13(7–15); E. miahuatlan 12.6(12–13); E. nitidipennis 12.3(9–15); E. chrysophana 11.6(7–18); E. suchixtepec 11(9–13); E. jalisco 10.5(9–12).