Biosystematics of the Genus Merobruchus of Continental North America and the West Indies (Coleóptera: Bruchidae)

Home , Acanthoscelides, Merobruchus, Pithecellobium, Pseudosamanea

'^^. United States |L|cJ|i Department of ^%^ Agriculture Biosystematics of the Agricultural Research Service Genus Merobruchus Technical Bulletin Number 1744 of Continental North America and the West Indies (Coleóptera: Bruchidae) Abstract

Kingsolver, John M. 1988. Biosystematics of the genus Merobruchus of continental North America and the West Indies (Coleóptera: Bruchidae). U.S. Department of Agri- culture, Technical Bulletin No. 1744, 63 pp.

A diagnosis of the genus Merobruchus is presented, including a key to species, synonymical names, geographical distribution, and host plant associations detailed for the 22 species now assigned to this genus for the United States, Mexico, Central America, and the West Indies. The following seven of these species are new to science: chetumalae, cristoensis, iysilomae, politus, porphyreus, triacanthus, and xanthopygus. Pseudopachymerus steinbachi Pic is a NEW SYNONYM of Merobruchus pickeli (Pic) (NEW COMBINA- TION), and Pachymerus subuniformis Pic is a NEW SYN- ONYM of Merobruchus bicoloripes (Pic). Bruchus flexicaulis Schaeffer is shown to be an available name and is provi- sionally synonymized with Merobruchus major (Fall) (NEW SYNONYMY). Bruchus limpidus Sharp is synonymized with Merobruchus placidus (Horn) (NEW SYNONYMY). Illustra- tions of salient characters are provided for each species. All known host associations are with seeds of leguminous trees and shrubs in the subfamily Mimosoideae, mostly in the genera Acacia, Lysiloma, and Pithecellobium. None of the species affect major agricultural crops, but they reduce the potential for regeneration of trees used for fuel, furniture, vegetable gums, tanbark, honey sources, and ornamental plantings.

This is part of a series of studies on bruchid genera contrib- uting to a comprehensive database for this important seed- feeding beetle family of North America. It provides scientific names for taxonomists and ecologists conducting studies in rangeland, pasture, and forest management in the South- western United States, Mexico, and Central America.

KEYWORDS: Acacia, Albizia, Enterolobium, Leguminosae, Leucaena, Lysiloma, Merobruchus, Mimosoideae, Pithecello- bium, seed beetles.

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Issued December 1988 United States Department of Agriculture Biosystematics of the Agricultural Research Genus Merobruchus Service Technical of Continental North Bulletin Nunnber 1744 America and the West indies (Ckileoptera: Bruchidae)

By John M. Kingsolver

Acknowledgments Contents

I thank the following individuals and institutions for loans of Materials and methods, 1 specimens used in this study: Annerican Museum of Natu- History of the genus, 2 ral History, New York; Blatchley Collection, Purdue Univer- Host plants, 3 sity, W. Lafayette, Indiana; British Museum (Natural His- Genus Merobruchus Bridwell, 4 tory), London; California Academy of Sciences, San Relationships of Merobruchus to other genera, 4 Francisco; California Insect Survey, Berkeley; Canadian Na- Key to species of Merobruchus, 5 tional Collection, Ottawa, Ontario; Chamela Biological Re- Major group, 7 serve, Jalisco, Mexico; Florida Department of Agriculture, major (Fall), 7 Gainesville; H.F and AT. Howden, Ottawa, Ontario, Can- Columbinus group, 9 ada; D.H. Janzen, University of Pennsylvania, Philadelphia; columbinus (Sharp), 9 CD. Johnson, Flagstaff, Arizona; Louisiana State Univer- Julianus group, 11 sity, Baton Rouge; M.V. Mâcedo, Rio de Janeiro, Brazil; Mu- julianus (Horn), 11 seum of Comparative Zoology, Cambridge, Massachusetts; triacanthus, new species, 12 Museum G. Frey, Tutzing, W. Germany; Muséum National politus, new species, 13 Histoire Naturelle, Paris, France; National Museum of Natu- Vacillator group, 15 ral History, Washington, DC; Natural History Museum of Los vac/V/afor (Sharp), 15 Angeles County, California; Ohio State University, Colum- porphyreus, new species, 15 bus; Snow Museum, University of Kansas, Lawrence; R. cristoensis, new species, 16 Turnbow, Ft. Rucker, Alabama; and J. Wappes, Chadds lineaticollis (Sharp), 17 Ford, Pennsylvania. /cní7///(White), 18 xanthopygus, new species, 19 Special thanks to CD. Johnson and D.H. Janzen for the Terani group, 20 many host records from field-collected and reared speci- terani Kingsolver, 20 mens. My appreciation to C Feller and L.H. Lawrence, Sys- Insolitus group, 21 tematic Entomology Laboratory, Agricultural Research Serv- Insolitus subgroup, 21 ice, for the many excellent habitus illustrations. insolitus (Sharp), 21 hastatus Kingsolver, 23 paquetae Kingsolver, 23 Sonorensis subgroup, 24 sonorensis Kingsolver, 24 santarosae Kingsolver, 24 iysiiomae, new species, 25 chetumalae, new species, 26 Placidus subgroup, 27 placidus (Horn), 27 solitarius (Sharp), 28 Boucheri subgroup, 29 boucheri Kingsolver, 29 Literature cited, 30 Appendix 1. Morphological terms, 33 Appendix 2. Synonymical list of Merobruchus species, 34 Appendix 3. Merobruchus species and associated host plants, 35 Appendix 4. Host plants attacked by Merobruchus species, 36

Trade names are used in this publication solely for the purpose of providing specific information. Mention of a trade name does not constitute a guarantee or war- ranty of the product by the U.S. Department of Agri- culture or an endorsement by the Department over other products not mentioned.

Biosystematics of the Genus Merobruchus Materials and Methods of Continental North America and the West Indies (Coleóptera: Bruchidae) John M. Kingsolver

During the 1970's and 1980's, nearly all the 27 Bruchidae Approximately 6,000 specimens were examined during this genera now known from continental North America and the study. Many were without host associations, but most of the West Indies have been monographed or revised, so that specimens provided by CD. Johnson and D.H. Janzen this fauna is now one of the most thoroughly documented were reared from seed pods collected in the field. in the world. It would be imprudent, however, to infer that all the Bruchidae species in this area are well known or Since male genitalia provide a major source of distinguish- even that all of them are described. The rich leguminous ing characters for species, careful preparation is necessary. flora in the subtropical and tropical regions is still poorly The following procedure should produce satisfactory sampled despite the years of extensive collecting and rear- results. Relax the specimen thoroughly either by soaking in ing by workers such as CD. Johnson in Mexico and Central hot water or alcohol or by using a relaxing chamber. Hold America and D.H. Janzen in Costa Rica. A taxonomic base the specimen between thumb and forefinger, and gently lift is being established, however, for subsequent workers to the apex of the pygidium with fine forceps. Sever the dorsal build upon. Generic limits are rather nebulous in several and ventral membranes surrounding the genital apparatus, sections of the family, and although additional species un- and remove the parts from the body. The eighth tergum and doubtedly await discovery, most of the known species now the Y-shaped eighth sternum are usually extracted with the have been well characterized. This study and one in median and lateral lobes. Extracted genitalia are cleared of progress will complement already published monographs tissue by treatment in 10 percent sodium hydroxide solution and revisions to provide workers with means of identifying and subsequently washed in acidified ethanol. Muscle fi- most of the bruchids attacking the seeds of desert, range- bers are then gently pressed out of the genitalia by using land, and pasture plants of North America and the Carib- blunt forceps or probes. The parts can be mounted in glyc- bean area. erin on slides for examination and illustration. Permanent storage is in glycerin in microvials attached to the specimen Larvae of Merobruchus species develop in the seeds of le- pin. guminous trees and shrubs that are characteristic of the more xeric areas of the tropical and subtropical regions. For each species, the following illustrations are given: Ha- Adults feed on nectar and pollen of available flowers. Host bitus, pygidium, median and lateral lobes of the male geni- plant genera include Acacia, Albizia, Leucaena, Lysiloma, talia, hindleg, and for some species the antenna. A few and Pithecellobium north of Panama and Pitheceííobium, species are further illustrated by drawings or photographs Pseudosamanea, and Enterolobium south of Panama; the of the head, female fifth sternum, and other isolated last genus is a host for Merobruchus bicoloripes (Pic). Al- characters. though members of this genus are not of primary economic significance to man and his activities, they do affect the Several techniques were used for the illustrations. Habitus potential regeneration of trees and shrubs that locally pro- and pygidial drawings are on coquille board with either ne- vide fuel, wood for furniture, vegetable gums, and tanbark. gro or carbon pencils; line drawings were made on bristol Several of the plants are used in ornamental plantings, and board using a microprojector; photographs were taken with a number of the shrubby species are good honey plants. either a Wild M-400 Photomakroscop or a scanning elec- tron microscope.

Descriptions vary in their extent. Species previously but in- adequately characterized are fully described as though they were new species, whereas for those I described in 1980, the diagnosis includes little more than color and salient morphological characters.

A glossary of morphological terms and definitions is in Ap- pendix 1. Many of these terms have been used in previous publications on the Bruchidae by various authors without explanation.

The geographical distribution for each species is divided Research entomologist, Systematic Entomology into two sections: A list of countries and States to provide a Laboratory, Agricultural Research Service, U.S. quick summary for mapping and for automated data proc- Department of Agriculture, c/o National Museum essing, and a more detailed list of localities given in the of Natural History, Washington, DC 20560. literature as well as those for which I have examined speci- History of the Genus

mens. For those species I described in 1980, the detailed Merobruchus was proposed by Bridwell in 1946 based on section includes records since 1979, the submittal date of Bruchus julianus Horn. He did not assign any additional the manuscript for my 1980 publication. species to Merobruchus in this or any subsequent publication, but Bradley later that year transferred Bruchus major Scientific names of plants are listed in Appendix 4 with Fall to the newly proposed genus. Species subsequently their authors' names and the associated species of Mero- assigned to Merobruchus were Bruchus placidus Horn bruchus. Botanical references consulted for authenticity of (Johnson, 1967), B. knulli VJhWe (Johnson, 1968), B. vacilla- names were Standley, 1922; Isley, 1973; Polhill and Raven, tor Sharp (Bottimer, 1968), B. columbinus Sharp (Kingsolver, 1981; Hastings, Turner, and Warren, 1972; and various 1975), B. limpidus Sharps and B. solitarius Sharp (Johnson books and files in the Botanical Library, National Museum and Kingsolver, 1977), B. insolitus Sharp (Johnson, 1979), of Natural History, Washington, DC. and B. lineaticollis Sharp (Johnson and Kingsolver, 1982). Some of these names had been placed in as many as four genera by various workers. In 1980, Kingsolver described six new species in Merobruchus from Mexico and Central America. An additional 7 new taxa are described in this publication, bringing the total to 22 species from the United States, Mexico, Central America, and the West In- dies. Merobruchus apparently has its greatest development of species in Central America.

Two species have been described from South America, Me- robruchus pickeli (Pic), 1927 (NEW COMBINATION) ( = steinbachi Pic, 1934, NEW SYNONYMY), and M. bicoloripes (Pic), ^927( = subuniformis Pic, 1938, NEW SYNON- YMY). Several South American species remain to be described.

Except for the few characters given in Bridwell's generic key (1946) and the brief, tentative diagnosis by Kingsolver (1980), no formal generic description of Merobruchus ap- peared in the literature until Borowiec's (1987) treatment of the bruchid genera of the world. His description is reason- ably accurate, but some important characters were over- looked, probably because he based it largely on the literature. The present publication is the first to bring together the described species placed in Merobruchus by various authors, with additional descriptions of new species, to include a thorough generic diagnosis based on 22 species with keys, redescriptions, synonymies, lists of host plants, and geographical distribution for continental North America and the West Indies. Characters for delimiting the genus as well as those for species and for species groups are found in external morphology, especially in the male genitalia. Other genera and species, described and undescribed, were examined to delimit the genus and to attempt to deter- mine its position within the family; however, no attempt has been made to devise a phylogenetic scheme for the genus, since only two South American species are considered and several more await description.

^ Now a synonym of B. placidus. Host Plants

Group characterizations rely largely on details of the male All known host plants with which Merobruchus is associated genitalia. These organs show a remarkable consistency of belong to the subfamily Mimosoideae in the family Faba- form within each species in this genus (except M. placidus, ceae. Members of this subfamily are, for the most part, q.v.), and at the same time they afford a source of charac- trees or shrubs found in tropical or subtropical regions, with ters for grouping species. some genera penetrating into temperate areas (Pohill and Raven, 1981). Fruit types vary from indéhiscent to partially In nearly all species of Merobruchus, the internal sac of the dehiscent to fully dehiscent. Many species are well adapted genitalic median lobe bears a Y-shaped sclerite, sometimes to a xeric climate. termed a "wishbone" wherever the resemblance is appro- priate. The median sclerite, however, may be slender, sub- Seven genera of mimosoids in 3 tribes are known to be triangular, or thornlike. Merobruchus major is the only spe- hosts for Merobruchus—Acacia in the Acaciae with 8 bru- cies in the genus without a median sclerite. chid species; Leucaena in the Mimoseae with 1 bruchid species; and Albizia, Enterolobium, Lysiloma, Pithecellobium, and Pseudosamanea in the Ingeae with 16 bruchid species feeding on seeds of 1 or more genera in this tribe. The five genera in the tribe Ingeae are closely related, with similar pod structure, and generic assignment of the plant species is often a matter of opinion. The common denominator for this assemblage of genera in the three tribes is probably the partial dehiscence of the seed pod valves.

A summary of host plant genera (see Appendix 3) shows that the major group, the columbinus group, and boucheri in the insolitus group are associated only with seeds of Pi- thecellobium, four of the six species in the vacillator group with Lysiloma, and the sole species in the terani group with Acacia; whereas the julianus group is associated with Aca- cia, Lysiloma, Leucaena, and Pithecellobium, and the insolitus group with Acacia, Albizia, Lysiloma, Pithecellobium, and Pseudosamanea. The association of Enterolobium with Merobruchus bicoloripes occurs only in South America. Genus Merobruchus Bridwell Relationships of Merobruchus to Other Genera

Merobruchus Bridwell, 1946:54; Bradley, 1946:41; Bottimer, No one yet has attempted a phylogenetic study of the Bru- 1962:955; Johnson, 1967:264, 1968:1269; Bottimer, chidae at the generic level. Some indications of affinities, 1968:1023; Johnson, 1969a:55, 1969b:676; Kingsolver, however, were suggested by Whitehead and Kingsolver 1970:374; Janzen, 1975:180; Kingsolver, 1975:60; Jan- (1975b, p. 212 et seq.), who advanced the tentative hypoth- zen, 1977:164; Johnson, 1979:122; Kingsolver, 1980:230; esis that Merobruchus belongs to a group of three "assem- Janzen, 1980:948; Johnson, 1981a:999; Johnson and blages" of acanthoscelidine genera corresponding roughly Kingsolver, 1982:418; Janzen, 1982:1274; Borowiec, to their host plants in the leguminous subfamilies Mimosoi- 1987:86. deae, Caesalpinioideae, and Papilionoideae. They are, re- Type-species: Bruchus julianus Horn, 1894:410, monotypic. spectively, the Merobruchus assemblage (including only Merobruchus), the Gibbobruchus assemblage (Gibbobru- Small to medium-sized beetles (1.8-8.8 mm). Body subde- chus, Penthobruchus, and Pygiopachymerus), and the Car- pressed dorsally or with elytra slightly concave. Head with yedes assemblage (Caryedes, Meibomeus, and perhaps frontal carina usually prominent (fig. 3); ocular sinus about Ctenocolum, although the affinities of the last two genera one-half length of eye (fig. 3); antenna gradually clávate are yet somewhat obscure). (fig. 4), segments of club ovate or elliptical. Lateral carina of pronotum with distinct sinuation at middle in lateral aspect. In most characters, Merobruchus would fall into the defini- Prosternum Y-shaped, acutely triangular between coxae, tion of the genus Acanthoscelides presented by Johnson sometimes separating coxal apices; cervical boss always (1983:5). He stated, however, that Acanthoscelides does not present and bisetose. Elytron with striae 3 and 4 usually appear to be a natural group; it is composed of many dis- arising basally from denticles on summit of basal gibbosity parate entities. Merobruchus could, with a slight expansion (except columbinus and major). Pygidium of male usually of his definition of Acanthoscelides, be considered a spe- reflexed apically to fit sternal emargination. Foreleg and cies group within Acanthoscelides, but I believe that the fol- midleg not especially modified; metafemur strongly swollen lowing characters are sufficient to segregate the species dorsoventrally (fig. 2); ventral margin proximad of pectén here assigned to Merobruchus as a distinct genus: The dis- carínate, lacking fine denticles; pectén with three to five tinctive combination of elytral striae 3 and 4 arising basally denticles, first denticle sometimes separated from those from bidentate basal gibbosities (except columbinus and following by gap of varying width (fig. 38); metatibia usually major), anal notch of female deeply emarginate and usually straight but with base bent at insertion into apex of femur, with lateral flanges, ventral valve of the male genitalia some species with tibia slightly arcuate (fig. 2). Abdomen broadly rounded or truncate, and armature of the internal with first sternum as long as or longer than remaining four sac of most species with a recognizably common theme of sterna combined; fifth sternum in both sexes usually with form and placement of sclerites, especially a Y-shaped me- carínate flange flanking emargination of posterior margin dian sclerite. Each of these characters, however, can be (fig. 8, extreme development), some males with slight found individually in one or another of the species now flange. Male genitalia with ventral valve nearly as broad as placed in Acanthoscelides. apex of median lobe (fig. 10); internal sac usually with wishbone-shaped median sclerite (figs. 10, 36) or modifica- Perhaps the species group in Acanthoscelides most closely tion thereof, sclerites of internal sac characteristic for spe- approaching Merobruchus is the mexicanus group (Johnson, cies, lateral lobes with sensory pores along lateral basal 1983, p. 6, No. 9). It differs principally in lacking the deep margin, ventral strut flat, not carínate. anal notch and accompanying flanges, in a distinctive pattern of internal sac sclerites, and in the shape of the ventral valve. In addition, nearly all 20 species tentatively assigned to the mexicanus group, like Merobruchus, are associated with plant genera in the subfamily Mimosoideae.

The two described southern South American species, M. pickeli and M. bicoloripes, possess the principal characters outlined in the generic description, i.e., anal notch of female deep and flanged on lateral borders, pronotum without prominent gibbosities, head relatively short, pygidium convex and without gibbosities, male genitalia with ventral valve broad and arcuate, and internal sac with wishbone sclerite and lateral denticles (figs. 137 and 138), but they deviate in the position of the basal elytral denticles. The denticles in M. pickeli are basal to striae 4 to 6 and are sim- Key to Species of Merobruchus

ilar to those of M. major except nearer the basal margin of 1. Elytra without bidentate basal gibbosity on striae 3 and the elytra, whereas M. bicoloripes has a single broad denti- 4 (figs. 19 and 63); size moderate to large (4.7-8.8 cle extending across the bases of striae 4 and 5, with stria mm); metatibial muero length three-fourths width of 4 deflected laterally to meet the denticle. The male genitalia metatibial apex 2 of both South American species are strikingly similar to Elytra with basal bidentate gibbosities; size various; mu- those of M. columbinas (cf. figs. 137 and 138). (See also ero of various lengths 3 Teran and L'Argentier, 1981:74-83, for a description of the 2. Striae 3-6 strongly mueronate basally (fig. 63); striae 6 and 7 not joined basally by earinate loop; 2 fifth ster- morphology and life history of M. bicoloripes.) num truncate or shallowly emarginate and with rounded flange either side of anal notch; J internal sae without selerites (fig. 64); postoeular lobe long (fig. 66) major (Fall) Striae 3-6 finely dentieulate basally, or with 5 and 6 lacking denticles (fig. 19); striae 6 and 7 joined basally by earinate loop; 2 fifth sternum deeply narrowly notehed with small, angulate flanges flanking notch; 5 internal sae with large Y-shaped sclerite (fig. 20); postoeular lobe short columbinus (Sharp) 3. Middle of third elytral interval with elongate, golden stripe; flanks of pronotum with dense gray vestiture; pygidium variegated 4 Middle of third elytral interval with or without elongate, golden stripe; if golden stripe present, flanks of pronotum with yellow vestiture or vestiture lacking; pygidium variously marked 6 4. First and second abdominal sterna each with denuded, polished spot eaudad of middle of metacoxa (fig. 89); pronotal disk with subtriangular, subdepressed, sometimes denuded area; 5 median lobe with median sclerite Y-shaped and pair of slender lateral spines (fig. 87); lateral lobes slightly bowed (fig. 88) politus, new species Abdominal sterna lacking denuded spots; pronotal disk convex with short basal suleus; 5 median lobe with median sclerite Y-shaped or not and with elongated lateral spines (figs. 42 and 122); lateral lobes nearly straight (figs. 43 and 123) 5 5. Pronotum with narrow median line of yellow setae sometimes extending from base to apex but sometimes only in basal one-half of disk (fig. 41); internal sae with median Y-shaped sclerite and paired, elongate lateral spines (fig. 42); size larger (3.2-5.5 mm) julianus (Horn) Pronotal disk with pubescence more uniformly distributed (fig. 121), without distinct median line; internal sae with three subequally elongated spines (fig. 122); smaller (2.3-3.5 mm) triacanthus, new species 6. Pronotum and pygidium reddish brown, with narrow, strongly contrasting, median, yellowish white stripe (fig. 53); median lobe with ventral valve truncate; internal sae with elongated U-shaped sclerite (fig. 54); lateral lobes nearly straight; spatulate (fig. 55) lineaticollis (Sharp) Pronotum and pygidium with or without median stripe, of various colors; pronotum lacking prominent median stripe; median lobe with ventral valve variously shaped 7 7. Pronotum with sparse or variegated yellow or gray vestiture 13 Pronotum with uniformly dense yellow or yellowish gray vestiture 8 8. Fifth sternum with prominent digitate process either side Pronotum without lateral patch; vestiture on disk lightly of anal notch; elytra usually with submarginal, variegated, sometimes condensed into round spot ei- rounded black macula (fig. 99) 2 santarosae Kingsolver ther side of midline near middle of disk (fig. 126); Y- Fifth sternum without prominent process, at most with shaped sclerite flat, middle of sac with two broad, marginal carina; elytral pattern various 9 thornlike spines and U-shaped sclerotized plate (fig. 9. Pygidium with median condensed line of yellow vesti- 127); 2 fifth sternum with prominent processes; anal ture, remainder of pygidium variegated as in figure notch shallow, truncate between processes 104 5 santarosae Kingsolver vacillator (Sharp) Pygidium uniformly pubescent except for small, median 17. Pygidium with narrow median stripe of white dark spot or large cordate spot 10 pubescence, in most specimens reaching nearly to 10. Pygidium with large, cordate, dark brown spot (fig. 52); apex (figs. 39 and 40); lateral margins of elytra suf- elytral vestiture yellowish, diffuse; pronotal vestiture fused with dark brown (fig. 35); pronotum with patchy usually with lateral yellowish patches 2 knulli (White) dark brown areas; 2 genitalia with nipple at apex of Pygidium without large cordate spot, uniformly yellow, or ventral valve; sclerites of internal sac slender, acute with small median spot, or pair of small spots (figs. 51 (fig. 36) insolitus (Sharp) and 135); basal two-thirds of elytra, pronotum, and Pygidium, elytra, and pronotum otherwise; ventral valve pygidium densely clothed with yellow vestiture; apical truncate, emarginate, or obtuse, lacking terminal nip- one-third of elytra black mottled with yellow and white ple; sclerites of internal sac otherwise 18 (fig. 131) 11 18. Pygidium with setal pattern as in figure 14; setae yellow- 11. Fifth abdominal sternum with rounded, flangelike cari- ish gray; elytral setal pattern of transverse bands of nae flanking anal notch (best seen in caudal view) small patches of white setae alternating with bands of (fig. 136) 2 xanthopygus, new species dark brown spots (fig. 9); pronotum with broad, dark Fifth sternum lacking prominent carinae(5) 12 median stripe, lateral margins with dense, gray pubes- 12. Y-shaped sclerite in S genitalia flat (fig. 132) cence; 5 genitalia (fig. 10) with three small, thornlike xanthopygus, new species spines in internal sac; 2 fifth sternum with two promi- Y-shaped sclerite skewed (fig. 48) knulli (White) nent, digitate apical processes (fig. 8) — boucheri Kingsolver 13. Vestiture of pygidium bright yellow, of scutellum gray; Pygidium with median stripe or basal triangle of setae metepisternum with inconspicuous grayish yellow (figs. 75 and 106); setal pattern of elytra various; pro- spots of vestiture; ventral valve broadly arcuate, wish- notum lacking distinct median stripe, usually vaguely bone sclerite large, internal sac with pair of lateral mottled; internal sac of 5 genitalia with median sub- spines (fig. 26) cristoensis, new species triangular sclerite and two lateral thornlike sclerites Combination of vestiture colors on pygidium and scutel- (figs. 79 and 81-85); 2 fifth sternum with slight flange- lum otherwise; ventral valve and internal sac armature like processes or processes lacking 19 otherwise 14 19. Pygidium with basal triangle indistinct (figs. 20 and 21); 14. Vestiture of pygidium and scutellum bright yellow; met- apical one-half of pygidium usually with paired dark episternal pubescent spots distinct; ventral valve brown spots, sometimes with additional paired sub- broader than long (fig. 117) 15 basal spots; fifth sternum lacking apical flanges or Vestiture of pygidium and scutellum yellowish gray or digital processes; $ genitalia with paired basal spines gray; metepisternum without distinct spots; ventral in internal sac 20 valve as long as or longer than broad 17 Pygidium with basal triangle or median stripe of setae 15. Body broad, elytra nearly quadrate (fig. 116); lateral distinct (fig. 115); apical spots lacking; fifth sternum parts of pronotal disk densely, conspicuously pubes- with or without apical flanges; 5 genitalia with or with- cent; pygidium densely pubescent except for median out basal spines in internal sac 21 denuded area (fig. 120); $ fifth sternum with promi- 20. Lateral margins of pronotum evenly arcuate (fig. 75); ely- nent processes flanking anal notch terani Kingsolver tral pattern ranging from nearly immaculcite with at Body narrow, elytra IVA times as long as wide (fig. 93); most faint brown spots on third interval to pattern in lateral parts of pronotal disk not conspicuously pubes- figure 75; base of median sclerite of internal sac cent except for lateral spots in some species; broad (figs. 81-85) placidus (Horn) processes of + fifth sternum variously developed 16 Lateral margins of pronotum slightly sinuate (fig. 106); 16. Pronotum with lateral patch of bright yellow setae near elytral pattern strongly mottled, usually with intensely anterior border; g genitalia with Y-shaped sclerite dark brown lateral maculae; median sclerite of internal skewed (fig. 94); $ fifth sternum lacking prominent sac slender, wedge-shaped (fig. 107) solitarius (Sharp) carínate processes; anal notch shallow 21. Pygidium black with small, narrow, white basal triangle porphyreus, new species (fig. 18); body small (2.1-2.3 mm); integumental color evenly dark red; dorsal vestiture inconspicuous (Yu- catan Peninsula) 2 chetumalae, new species Pygidium otherwise; integumental color and dorsal pubescence various 22 Major Group

22. Internal sac lacking basal spines (fig. 112); ventral valve The species included in the major group differ fronn all subtruncate or slightly emarginate apically (figs. 31 other Merobruchus by the following combination of charac- and 71) 23 ters: Postocular lobe prominent, elongated; elytra without Internal sac with basal spines (figs. 31 and 71); ventral gibbosities at bases of striae 3 and 4 but with prominent valve truncate or arcuate (figs. 31 and 71) 25 denticles at bases of striae 3-6; internal sac of 5 genitalia 23. Pygidium with broad, subtriangular pad of white setae without large sclerites. Additional characters are pronotum (fig. 114) strongly contrasted and distinct from remainder of pygidial vestiture, usually with short detached moderately gibbous with median sulcus and subbasal gib- line of pubescence at apex of triangle (Sonora, Mex- bosities evident; pectén of metafemur lamellar with five ico, to Colombia) sonorensis Kingsolver evenly spaced denticles; metatibia with muero about one- Pygidium with inconspicuous basal triangle of setae not half as long as apical width of tibia, lateral and coronal den- strongly contrasted with remaining pygidial vestiture ticles sharply developed; thoracic pleura lacking setal (figs. 17 and 62) 24 patches; elytral surface strongly concave mesally, apices of 24. Lateral thornlike sclerites in apical one-half of internal seventh and ninth intervals convex; pygidium medially sul- sac with axis of apex extending at 90° to axis of base cate before apex, both sexes with slight depression before (fig. 59) (Florida, Cuba, Haiti, Bahamas) apex; pattern dimorphic, 2 more strongly contrasting. One lysilomae, new species species. Thornlike sclerites elongated with axis of apex extending at 30° to axis of base (fig. 16) (Yucatan Peninsula) 5 chetumalae, new species Merobruchus major (Fall) 25. Ventral valve truncate (fig. 31); basal spines of internal sac thornlike, with broad base and acuminate (Figs. 63-69) apex hastatus Kingsolver Ventral valve arcuate (fig. 71); basal spines of internal Bruchus major Fall, 1912:320; White, 1941:189. sac subtriangular, minute paquetae Kingsolver Mylabris major: Leng, 1920:304. Merobruchus major: Bradley, 1946:41; Blackwelder and Blackwelder, 1948:44; Johnson, 1967:264; Bottimer, 1968:1023; Johnson and Kingsolver, 1982:418. Bruchus flexicaulis Schaeffer, 1904:229 (manuscript name). Acanthoscelides flexicaulis: Anonymous, 1940:36; ibid., 1942:7, 24; ibid., 1943:6, 28; ibid.. 1944:6, 24; ibid., 1945:7, 24; Zacher, 1952:465. Merobruchus flexicaulis: Wheeler, et al., 1948:23, 37. Acanthoscelides flexicaule: Zacher, 1952:470 (apparent lapsus). Bruchus julianus: Schaeffer, 1904:229 (misidentification); Gushman, 1911:491 (misidentification).

Color. Integument deep red to piceous. Recumbent vestiture of ochreous, gray, and brown hairs in mottled pattern, some pale yellow hairs with brassy sheen, distributed as follows: Head densely clothed with pale yellow hairs and scattered gray hairs on vertex and gena, antenna with segments 1-7 gray, 8-10 dense brown, 11 with brown and gray mixed. Pronotum with mostly pale yellow and some intermixed gray hairs in median one-third of disk, lateral parts of disk and flanks gray. Elytra with gray, brown, pale yellow, and brassy hairs in pattern as in figure 63. Pygidium of 5 (fig. 68) evenly gray to pale yellow with dark integument underlying vestiture, pygidium of 2 with pale yellow, gray, and brown in pattern of figure 69; venter of body mostly gray with some intermixing on legs, metepisternum, and posterior margins of abdominal sterna.

Structure. Body elongate-ovate (fig. 63). Head broad, obovate (fig. 66), ocular sinus less than one-half eye length; frontal carina vaguely defined, expanded dorsally into flattened, triangular boss with fine longitudinal sulcus; Irons, Body length 5.2-8.8 mm, width 2.8-4.7 mm; pronotal clypeus, and lateral face of mandible finely, densely punc- length 1.5-2.2 mm, width 1.9-2.9 mm. tate except boss minutely striate; postocular lobe broad, crescentic, vaguely limited posteriorly by shallow sulcus Type locality. Bruchus major. Texas, Brownsville, Esperanza (fig. 66); antenna gradually clávate from fourth segment, Ranch. Bruchus flexicaulis: Texas, Brownsville, Esperanza club segments eccentric and subserrate. Pronotum cam- Ranch. paniform in dorsal aspect (fig. 63), disk slightly elevated in median one-third with anterior part of elevation flat to Type depository. Bruchus major. Type 25056, in Museum of slightly depressed, posterior part sulcate, lateral parts of Comparative Zoology, Cambridge, Massachusetts. Bruchus disk convex, indistinct umbo present each side near basal flexicaulis: Lectotype here designated, Esperanza Ranch, border; lateral margin of disk demarcated by low, sinuate VII-1903, Siderocarpus flexicaule. Type 42446, National Mu- carina extending from posterolateral corner of disk to mid- seum of Natural History, Washington, DC. way between anterior and posterior borders; cervical sulcus narrow, deep; prosternum sharply triangular between coxae, Distribution. U.S.A.: Texas: Counties of Bexar, Cameron, then vertically emarginate to acute apex beyond contiguous Hidalgo, Nueces, Starr, Victoria, Webb, Wharton, Willacy. coxal apices. Scutellum quadrangular, deeply emarginate MEXICO: Tamaulipas, Veracruz. apically, bidentate. Elytra together about as long as wide, middle of lateral margins nearly straight; median one-third Biology. The only host plant recorded for this species is depressed between low ridges running along fifth intervals Pithecellobium flexicaule (Siderocarpus in older literature), two-thirds length from base; striae regular in course, slightly and M. major is the only known obligate bruchid predator of deflected laterad in basal one-third, striae 1 and 2 arising this tree. Stator beali Johnson has also been reared from from deep basal pits, striae 3, 4, 5, and 6 arising from be- this host, but it attacks other species of Pithecellobium as neath flattened, triangular, scalelike subbasal denticles, all well. The common name of the tree is Texas ebony. It is striae free apically except 4 and 5 conjoined; striai punc- used locally for fenceposts, cabinet wood, and fuel; the tures closely placed and slightly depressed in sharp, narrow seeds are sometimes used to make an ersatz coffee. grooves; mesosternum with intercoxal process flat, subtruncate; postmesocoxal sulcus rounded, following contour of l\Aerobruchus major attacks seeds contained in a woody, coxal cavity. Abdomen with first segment 1.8 x as long as indéhiscent pod and exits through holes drilled by the larva remaining segments in 5, 1.2 x in 2, fifth segment deeply through the pod wall. Two other species, Megasennius muri- emarginate apically for reception of pygidial apex in 5, catus (Sharp) from Central America and Pygiopachymerus emargination with distinct reflexion with posterior face con- lineóla (Chevrolat) from Central and South America, which cave and slightly lobed laterally, emargination shallow in $ have in common an elongated postocular lobe and elevated but with reflexion similar to that of $. Pygidium of 5 obovate basal elytral spines, also breed in woody pods (Whitehead (fig. 68) with apex truncate, apical one-half of disk and Kingsolver, 1975a). Whether these shared morphologi- shallowly, broadly sulcate, sulcus flanked by low umbones; cal features have any significance or not is unknown. pygidium of 2 semicircular but with apex slightly elongated and truncate, disk with paired slight convexities one-third Discussion. This species is the largest one treated in this from apex. Metacoxa with moderately dense, setigerous publication. It is sometimes confused with M. julianus, but it punctures except in polished area immediately laterad of is larger (length 5.2-8.8 mm vs. 3.2-5.5 mm ior julianus), trochanteral insertion; metafemur as in figure 67, pectén has prominent, flattened denticles at the bases of striae 3, with all denticles having common, bladelike base, ventral 4, 5, and 6 (denticles on striae 3 and 4 only), and lacks any face of femur slightly concave; metatibia slightly bent at ex- large sclerites in the male genitalia (large, U-shaped treme base (fig. 67), muero slender, acute, lateral denticle sclerite present). short, blunt, coronal denticles small, ventral carina flangelike, terminating in muero, lateral carina terminating in lat- Schaeffer (1904) discussed under the name julianus a "gi- eral denticle, ventrolateral carina terminating in sinus be- gantic Bruchus ... on Acacia flexicaulis, in the large seed tween muero and lateral denticle, dorsomesal carina pods of which it undoubtedly breeds." He noted the differ- prominent, terminating short of terminal margin. Male geni- ences in size between smaller, "starved" specimens de- talia: Median lobe (fig. 64) 3 x as long as wide, ventral scribed by Horn and the larger specimens from A. valve cordate, apex rounded; internal sac densely lined with flexicaulis, but he did not formally describe the larger speci- fine spicules, devoid of large sclerites, spicules condensed mens; instead, he noted that he distributed specimens into two lobate masses near base of sac, middle of sac with under the manuscript name flexicaulis, and some museums apparent wide rows of denticles, apex with cylindrical struc- may yet have specimens labeled flexicaulis. Since the name ture. Lateral lobes (fig. 65) bowed, rounded apically, cleft flexicaulis was first listed as a synonym of Bruchus julianus, two-thirds their length. it has been treated as an unavailable name by Johnson Columbinus Group

(1967) and Bottimer (1968) referring to Article lid of the The single species in this group is distinctive in several re- International Code of Zoological Nomenclature (1961). Ac- spects. Elytral intervals are of equal width basally and bear cording to the 1985 Code (Article lie), however, a name individual denticles (fig. 19), whereas in all other groups, first cited in synonymy may be made available if it was except the major group, the basal denticles of striae 3 and used as a name of a taxon prior to 1961. In 1940, the U.S. 4 are elevated on a gibbosity; striae 5 and 6 are joined ba- Department of Agriculture List of Intercepted Plant Pests (p. sally by a carínate loop, a unique character (fig. 19); the 36) listed Acanthoscelides flexicaulis, and in subsequent third interval, pygidium, and pleura lack distinct patches of years (1942, 1943, 1944, and 1945), the same annual publi- golden setae; postocular lobe narrow. The deep sternal cation listed Acanthoscelides flexicaulis (Schaeffer). In 1948, notch with lateral flanges in the female is consistent with Wheeler et al. used the combination Merobruchus other groups in Merobruchus. Male genitalic characters in- flexicaulis (Schaeffer), and in 1952, Zacher used Acantho- dicate a similarity more to the julianus group than to any scelides flexicaulis and A. flexicaule in a summary of bru- other. chid host plants. These listings constitute usage within the meaning of Article lie. Merobruchus columbinus (Sharp)

Although the name flexicaulis (Schaeffer, 1904) predates (Figs. 19-24) major (Fall, 1912), I prefer to use the latter name for this species in the interests of stability since major has been Bruchus columbinus Sharp, 1885:447. used in several publications, and I intend to apply to the Pseudopachymerus columbinus: Pic, 1913:10. International Congress of Zoological Nomenclature to Caryedes columbina: Blackwelder, 1946:758. suppress the name flexicaulis in favor of major. Merobruchus columbinus: Kingsolver, 1975:60; Janzen, 1975:178, 1977:164, 1980:938, 948, 1982:1274; Johnson One female specimen collected by Janzen at Puente Nacio- and Kingsolver, 1982:418; Janzen, 1983:738. nal, Veracruz, 21-VI-1962, is strongly melanistic and may represent a new species. Since Pithecellobium flexicaule Color. Integument dark red to piceous; antenna reddish apparently does not extend into Veracruz, the specimen brown except segments 8-10 piceous; eye usually black; probably emerged from a different host plant. metacoxal face piceous. Pubescence of gray, brown, and bronzy fine hairs in faint but variable pattern on pronotum and elytra ranging from most distinct pattern shown in figure 19 to uniform distribution of gray with bronzy highlights, sometimes with brown spots on third elytral interval; Î pygidium usually with small, paired median spots.

Structure. Body ovate, moderately deep. Head obovate (fig. 22), postocular lobe narrow; vertex finely, densely punctate, vestiture directed cephalad, frons more coarsely punctate, punctures tending to coalesce longitudinally, vestiture directed mesad, frontal carina prominent, border between vertex and frons marked by shallow, transverse depression; clypeus punctate as on head; ocular sinus one-half length of eye; antenna slender, reaching past posterolateral corner of pronotum, serrate from 5th segment, club strongly eccentric, 11th elliptical. Pronotum (fig. 22) elongate-trapezoidal, lateral margins gently concave, apical margin truncate, base bisinuate, disk evenly convex except shallow sulcus on basal lobe and shallow depressions at posterolateral corners; surface with scattered microfoveolae, each with central seta, interspaces punctulate; lateral carina represented by short, curved ridge above coxal cavity; cervical sulcus short, fine; prosternai apex separating procoxae and extending above and slightly beyond them; mesosternum truncate apically, postmesocoxal suici slightly widened behind coxae but not angulate. Scutellum small, quadrate, deeply emarginate. Elytra (fig. 19) as long as wide, depressed between seventh intervals; striae regular in course, 3, 4, and 5 with small basal denticles, basal margin Janzen (1977) discussed the life history of M. columbinas lacking gibbosity at striae 3 and 4; striae 6 and 7 on R saman in Guanacaste Province, Costa Rica. He stated connected basally with U-shaped carina (fig. 19); striae that larvae of this bruchid kill at least 43 percent of the fine, narrow; intervals of even width; minutely strigose. Met- seed crop of each tree. Females glue eggs on the pods acoxal face finely, densely punctate; hindleg as in figure 23, when the seeds are well formed in the pod yet green and pectén without gap; metatibia (fig. 23) slightly arcuate, soft. Within 2 weeks, larvae hatch and bore directly through channel between ventrolateral and ventral carinae polished; the pod wall into the seed. Before pupating, mature larvae muero slender; lateral denticle small, blunt; coronal denti- cut through the seed coat and partly through the pod wall cles three or four. Abdomen of 6 with fifth sternum broadly leaving a "trap door" through which the adult can easily emarginate to one-half length of sternum, emargination emerge usually immediately before the pods drop to the slightly flanged, in 2 more narrowly emarginate one-fourth ground. Merobruchus columbinas apparently has one gen- length of sternum, strongly flanged and with setal tufts; 5 eration per year and is not attacked by any parasites and 2 pygidia similar, truncate apically. Male genitalia: Me- (Janzen, 1977, p. 165). Another bruchid, Stator limbatus dian lobe (fig. 20) about 3 x as long as width across apex, (Horn), oviposits on seeds of P. saman through the exit ventral valve arcuate; internal sac with large, wishbone- holes of M. columbinus or through moth exit holes, and it shaped sclerite near base and two thornlike sclerites at completes its development in the seeds fallen to the middle; base of sac lined with minute denticles, apex with ground. mixture of fine, slender spines, minute denticles, and minute, pectinate sensory processes; apical closure valve Discussion. Pithecellobium saman is used locally as a ringlike, on dorsal side of apex. Lateral lobes as in figure shade and garden tree, for furniture and paneling, carvings, 21. and bowls. The seeds are edible.

Body length 4.0-5.4 mm, width 2.3-2.9 mm; pronotal length 1.3-1.5 mm, width 1.7-2.1 mm.

Type locality. GUATEMALA: Rio Maria Linda, 500'.

Holotype depository. British Museum (Natural History), London.

Distribution. GUATEMALA: Escuintla. HONDURAS: Cortés, Valle. EL SALVADOR: San Salvador, La Libertad, La Paz. NICARAGUA: Carazo, Chinandega, Granada. COSTA RICA: Guanacaste, Limón, Puntarenas, San José. PANAMA: Ca- nal Zone, Veragua. COLOMBIA: Valle del Cauca. VENEZU- ELA: Carabobo, Falcon, Lara, Los Andes, Miranda. TRINI- DAD. The countries but not specific provinces or localities were listed by Janzen (1977).

Specific records. HONDURAS: Cortés; Nacaome. EL SAL- VADOR: Lake llopongo; La Liberatad; San Vicente. NICA- RAGUA: Diria; Chinandega; San Cristobal. COSTA RICA: 4 mi N Las Cañas; Taboga, 6 mi SW Cañas; Bagaces; 10 mi NW Esparta; Turrubares; San José; Batán; Palo Verde. PANAMA: Gatun Locks; 35 mi NE Santiago. COLOMBIA: Rio Frio; Merida. VENEZUELA: Cagua; Rancho Grande; Choroni; Puerto Cabello; Caracas; Felipe; Zulia; Carafeobo. TRINIDAD (no locality).

Biology. The only host association known for this species is with Pithecellobium saman. The majority of collections were in March, but the range was from January to July and Sep- tember, with one collection in Venezuela in November.

10 Julianus Group

Members of the julianus group differ from ail other Merobru- antenna short, barely reaching posterior corner of protho- chus by the foliowing combination of characters: Postocular rax, clávate, subserrate from fifth segment. Prothorax sub- lobe short (fig. 44); base of each elytron with dentate gib- campaniform in dorsal aspect (fig. 41), lateral margins bosity at bases of striae 3 and 4; ventral valve arcuate (fig. nearly straight, disk evenly convex without asperities, small 42); lateral lobes elongated (fig. 43). Additional characteris- depression near each posterolateral corner, shallow longitu- tics are pronotum barely gibbous, median sulcus evident dinal sulcus in prescutellar lobe; disk densely, minutely, dis- only at base, subbasal gibbosities obsolete; elytral disk cretely foveolate, each foveola with recumbent seta arising convex or depressed, third interval with elongate yellow near anterior border; lateral carina a distinct ridge extend- spot; metepisternum with vaguely defined yellow spots; py- ing from posterolateral corner nearly to articulation of coxa; gidium not dimorphic. Three species. cervical sulcus short, deep; prosternum Y-shaped, intercoxal piece acutely triangular, not separating coxal apices. Merobruchus julianus (Horn) Scutellum angulately emarginate apically, densely setose. Elytra together slightly longer than wide, appearing quad- (Figs. 41-46) rate (fig. 41); disk flat mesally, slightly depressed around scutellum; striae regular in course, intervals of uniform Bruchus julianus Horn, 1894:410; Fall, 1910:186, 1912:320; width, integument strigose; striai punctures deep, separated Wenzel, 1912:140. by their diameter, striai suici shallow or obsolete, stria 1 Mylabris julianus: Leng, 1920:304; Bôving, 1927:141. arising at apex of scutellum, striae 3 and 4 arising basally Merobruchus julianus: Bridwell, 1946:54; Blackwelder and from distinct denticles on prominent gibbosity, striae 2, 5, Blackwelder, 1948:44; Johnson, 1967:264, 1968:1269 and 6 each arising from small, deep pit, all striae except (lectotype designation); Bottimer, 1968:1024, 1969:1187; conjoined 5 and 6 free apically. Mesosternum with inter- Johnson, 1969a:55, 1969b:676; Forister and Johnson, coxal process rounded apically, densely punctulate, postmes- 1970:84 et seq.; Kingsolver, 1970:374 (male genitalia); ocoxal sulcus angulate. Metacoxa densely, finely punctate Johnson, 1979:122; Johnson and Kingsolver, 1982:418. in posterior two-thirds, polished and impunctate in anterior Acanthoscelides julianus: Blackwelder, 1946:759. one-third; metafemur (fig. 45) moderately incrassate, Bruchus ochreolineatus Fall, 1910:186; Cushman, densely punctate on anterior face, subapical pectén with 1911:492; Johnson, 1968:1269; Johnson and Kingsolver, one long and three or four shorter denticles, all denticles 1982:418. about equally spaced; metatibia (fig. 45) strongly arcuate Bruchus ochrolineatus: Pic, 1913:38. basally, slightly arcuate in apical four-fifths; muero short, slender; lateral denticle prominent, coronal denticles usually Color. Integument dark red throughout except antennal club five; ventral carina flangelike in apical two-thirds, ventrolat- piceous. Pubescence of recumbent gray, coppery, and pale eral carina extending nearly to base of muero, lateral carina yellow hairs distributed as follows: Head, venter of body, ending in lateral denticle, dorsomesal carina complete. Ab- and legs sparsely gray except for vaguely condensed spots domen with first sternum as long as remaining sterna com- on metepisternum and similar spot on each abdominal ster- bined; fifth sternum strongly emarginate to receive apex of num near its lateral border, spots sometimes slightly yellow- pygidium in 5, shallowly emarginate and without prominent ish. Pronotum with broad, irregular median stripe of cop- ridges flanking emargination in 2; pygidium (fig. 46) in both pery hairs bisected by narrow median stripe of pale yellow sexes evenly convex. Male genitalia: Median lobe (fig. 42) 4 hairs, lateral margins of disk and flanks densely covered X as long as wide, ventral valve with apical margin evenly with gray hairs (fig. 41). Scutellum gray. Elytra with gener- arcuate; internal sac with wishbone-shaped median sclerite ally mottled pattern of coppery, gray, and pale yellow hairs and two long, slender flanking sclerites, interior of sac lined (fig. 41) with yellowish spots at extreme bases of third, fifth, with fine spicules. Lateral lobes as in figure 43. and seventh intervals, elongate pale yellow spot at middle of third interval, smaller elongate spot at basal one-third of Body length 3.2-5.5 mm, width 1.9-3.1 mm; pronotal fifth interval, and at middle of fifth and seventh intervals. length 1.1-1.5 mm, width 1.4-2.1 mm. Pygidium densely covered with gray and pale yellow hairs, latter in vague patches (fig. 46), integument showing Type locality. Texas (no locality). Lectotype 3905 in through pubescence in subbasal marginal spots and irregu- Museum of Comparative Zoology, Cambridge, Massachu- larly in middle of pygidial disk. setts. Designated by Johnson (1968).

Structure. Head turbiniform (fig. 44), eyes moderately pro- Distribution. U.S.A.: Texas: Counties of Atascosa, Bee, Bex- tuberant, frons with prominent median carina expanded ar, Brewster, Cameron, Dimmit, Frio, Goliad, Hidalgo, How- dorsally into small, flattened, impunctate boss; vertex, frons, ard, Jeff Davis, Kimble, La Salle, Maverick, McMullen, and clypeus densely punctate and setose; postocular lobe Nueces, Presidio, Shackelford, Starr, Terrell, Uvalde, Val narrowly expanded dorsally, fringed with cinereous hairs; Verde, Webb, Zavala. Louisiana: Caddo Parish. Arizona:

11 Coconino, Piñal, Santa Cruz, Yavapai. MEXICO: Baja Cali- Urosigalphus bruchi Crawford (Forister and Johnson, 1970), fornia Norte and Sur, Chihuahua, Coahuila, Nuevo Leon, and three unidentified species of Eurytoma (Johnson, Tamaulipas. 1967:269). Forister and Johnson (1970) have more exten- sively discussed the biology of this bruchid. Biology. Adults have been collected from April through Au- gust; specimens have been reared from host material col- Discussion. Some specimens oí ¡ulianus overlap in size lected throughout the year. Some specimens probably rep- those of ma¡or and are sometimes confused with it (cf. dis- resent flower visitation records: "cotton," Phacelia cussion of ma¡or). integrifolia Torr. (Hydrophyllaceae), and Sphaeralcea angustifolia (Cav.) G. Don. (Malvaceae). The species of Acacia listed as hosts oí ¡ulianus are excellent honey plants. Acacia berlandieri, wrightii, and roemer- The host plants for M. ¡ulianus for which authentic rearing iana are used as ornamental shrubs, and berlandieri and records are known include at least Acacia berlandieri greggi are potential commercial sources of plant gums. Benth., A. coulteri Benth., A. greggi Gray, A. roemeriana Scheele, and A. wrightii Benth. Specimen labels were taken Merobruchus triacanthus, new species at face value. Some records were obtained from herbarium specimens. Records in the literature of the following host (Figs. 121-125) plants need confirmation: Acacia juncifolia Benth., A. rigi- dula Benth., Desmanthus illinoensis (Michaux) Kuntze, Mi- Merobruchus sjp. 3: Johnson, 1979:123. mosa borealis Gray, and Pithecellobium sp. Acacia juncifolia is an introduced Australian species. Records from A. rigi- Color. Integument deep red to piceous. Head usually red, dula and Pitliecellobium are based on plant quarantine in- sometimes with vertex piceous, antenna with segments 1-5 terceptions from Mexico. No Merobruchus species are oth- and 11 reddish, intermediate segments piceous; pronotum erwise known to infest seeds of either Mimosa or with most of disk piceous, flank deep red, prosternum pic- Desmanthus, and these records are probably the result of eous; most of elytra piceous with small reddish areas later- misidentifications of plants. Seeds of Mimosa borealis are ally, mesosternum and metasternum deep red to piceous, too small to support even the smallest individuals of M. ¡u- metepisternum reddish, abdomen reddish with piceous lianus. Seeds of Desmanthus are likewise too small. blotches; pygidium reddish with piceous median spots (fig. 125); legs mostly deep red with ventral margins piceous. L.J. Bottimer's unpublished field notes for 1924-25 for/¿y- Pubescence of recumbent gray, coppery, brown, and pale lianus at Uvalde, Texas, stated "In Acacia berlandieri, larva yellow hairs distributed as follows: Head usually yellowish eats an irregularly oval hole in one side of seed then fas- mixed with gray. Pronotum on darker area of pronotal disk tens edge of hole to valve. Adult cuts a hole through pod to (fig. 121) with coppery and yellowish hairs, rarely with frag- escape; in Acacia greggi, however, the larva cuts a clean mented median line of yellowish hairs, flanks of pronotum hole in seed but does not fasten seed to pod." He also largely gray, occasionally intermixed with pale yellow. Elytra found ¡ulianus hibernating in capsules of buckeye at Ft. predominantly gray in middle of disk intermiixed with brown Davis, Texas. Johnson (1967) stated that A. berlandieri pods and coppery (fig. 121), middle of third interval with elon- dehisce soon after they ripen, and seeds fall to the sub- gated pale yellow spot, lateral areas of elytra with vague strate except for those infested by ¡ulianus, which remain bar pattern, bases of elytra occasionally with pale yellow glued to the inner wall of the pod. A larva may feed in more patches. Pygidium (fig. 125) mostly pale yellow with inter- than one seed of this host by cutting a passage between mixed grayish patches, base with three yellowish patches in seeds and gluing the seeds together forming a tube-like both sexes, piceous integumental spots showing through structure, but in A. greggi, the larvae develop one to a vestiture. Venter of body with intermixed gray and yellowish seed. This may be explained by the fact that seeds of A. hairs in no particular pattern except mesepimeron yellowish greggi are larger than those of A. berlandieri and have suffi- and metepisternum with three yellowish patches, each ab- cient nutrients in one seed for a larva to develop fully. For- dominal segment with small, indistinct yellowish patch near ister and Johnson (1970) found that eggs oi ¡ulianus are laid lateral margin; legs mostly gray, sometimes mixed with yel- on immature pods and that they are fastened to the pod low mottled pattern evident on metafemur and metatibia. surface with several strands of mucilage to permit the egg to remain attached during growth expansion of the pods, Structure. Head turbiniform, not sexually dimorphic, post- but egg shells are sloughed off after eclosión and no evi- ocular lobe narrow, setose; vertex, frons, and basal two- dence remains on mature pods. Adults harden in the pupal thirds of clypeus strongly but finely punctate, frons with chamber, then cut a symmetrical hole in the pod wall to es- prominent carina ending dorsally in small impunctate boss; cape. The cycle from egg to adult is about 60 days. This antenna short, barely reaching posterior corner of prono- bruchid is parasitized by Eupelmus cushmani (Crawford), tum, subserrate-clavate from fifth segment. Pronotum cam-

12 paniform (fig. 121), disk convex without asperities except Holotype, allotype, and paratypes deposited in National Mu- short, shallow sulcus on prescutellar lobe and slight de- seum of Natural History, Washington, DC. Paratypes also pression on each posterolateral corner of disk; surface with deposited in CD. Johnson collection. Flagstaff, Arizona; densely placed, fine, circular foveae, each with a seta aris- Canadian National Collection, Ottawa; British Museum of ing beneath its anterior rim, most foveae discrete; lateral Natural History, London; and Museum G. Frey, Tutzing, Mu- margin of disk marked with blunt carina extending from nich, West Germany. posterolateral corner one-half distance to coxal cavity; cervical sulcus short; prosternum with apex separating coxae Distribution. MEXICO: Michoacán, Colima. at their apices. Scutellum quadrate, emarginate, and bidentate apically. Elytra together (fig. 121) slightly longer than Biology. Five host plants are known for this species: Acacia wide, disk flat or slightly depressed mesally; striae regular acatlensis, coulteri, and aff. riparioides; Leucaena guatema- in course, intervals of uniform width, surface strigose, striai lensis; and Lysiloma divaricata. Lysiloma divaricata is an ex- punctures small, deep, striai suici shallow; stria arising at cellent source of tanbark in Mexico and Central America. end of scutellum, 2, 5, and 7 arising from deep basal pits beneath anterior rim of elytron, 3 and 4 each arising from Discussion. Merobruchus triacanthus is most similar to M. small denticle on prominent gibbosity, all striae ending free ¡ulianus in color pattern, body shape, details of the male apically or occasionally with 4 and 5 conjoined. Mesoster- genitalia, and lack of a definite ridge on either side of the num with intercoxal process rounded apically, postmeso- genital notch of the terminal sternum. Specimens of triacan- coxal sulcus angulate. Metacoxa densely punctate; metafe- thus average smaller than those of ¡ulianus, the muero is mur as in figure 124, with lateral face finely, densely shorter (fig. 45, cf. with 124), the elongated ochreous spot punctate, pectén with one long, acute, slightly curved denti- in the middle of the third interval is not as prominent, the cle, and three shorter denticles, long denticle separated gap between the first and second denticles of the pectén is from others by a gap (fig. 124); metatibia strongly arcuate at wider, and the middle spine of the internal sac is more slen- base, muero short, slender; lateral denticle blunt, coronal der (fig. 42, cf. with 122). denticles minute, ventral carina flangelike in apical three- fourths, ventrolateral carina extending nearly to notch be- The name triacanthus refers to the three nearly equal tween muero and lateral denticle, lateral carina ending in spines in the internal sac of the male genitalia. lateral denticle, dorsomesal carina prominent, complete. Abdomen with first sternum longer than remaining sterna Merobruchus politus, new species combined in 5, subequal to remaining segments in 2; caudal margin of fifth segment deeply, broadly emarginate for (Figs. 86-92) reception of reflexed pygidial apex in 5, slightly emarginate but without lateral ridges in 2; pygidium densely, finely punc- Color. Integument reddish brown to pieeous; head with tate. Male genitalia: Median lobe broad (fig. 122), 4 x as frons and elypeus pieeous except for narrow reddish band long as wide, ventral valve evenly arcuate; internal sac with inside each eye; antenna with segments 1-5 reddish, 6-11 three elongated spines of nearly equal size and shape, en- pieeous; legs mostly reddish, metacoxa and polished area tire sac lined with minute spicules. Lateral lobes (fig. 123) on first and second abdominal sterna pieeous. Pubescence nearly straight, apices rounded, cleft about two-thirds their of gray, yellow, and dark brown setae; flanks of prothorax length. densely clothed with yellowish gray to gray setae contrasting sharply with yellow discal stripe; venter of body densely Body length 2.3-3.5 mm, width 1.5-2.3 mm; pronotal clothed with gray; elytra mottled gray, yellowish gray, and length 0.9-1.1 mm, width 1.1-1.6 mm. brown in vague pattern, third interval with elongate yellowish gray (in some specimens golden yellow) median stripe Holotype J. MEXICO: Michoacán, 16 mi E Jiquilpan, 5300', limited by dark brown spots, a short, similarly colored spot ex seeds Acacia aff. riparioides, CD. Johnson 256-68. Al- near apex; fifth interval and small apicolateral and mediolat- lotype 2 and 28 paratypes, same data. Other paratypes: eral spots yellowish gray, scattered spots gray; 5 pygidium MEXICO: Colima, 7 mi S Colima, ca. 1100', 7-III-1973, ex (fig. 91) evenly clothed with yellow except for small median seeds Acacia acatlensis, Johnson 380-73 (listed as Lysi- spot at basal one-third and two lateral spots; 2 pygidium loma desmostachys by Johnson, 1979:123) (9); 3 mi S Co- (fig. 92) with median line and lateral stripes yellowish. lima, 7-III-1973, ex seeds Leucaena guatemalensis, John- son 373-73. Oaxaca, Tehuantepec, 1000 m, 12-X-1966, G. Structure. Body somewhat elongated, nearly 2 x as long Frey (3). Michoacán, 20 mi S Nueva Italia, 6-III-1979, in as wide (fig. 86). Head turbiniform, eyes protuberant, vertex seeds Acacia coulteri, Johnson 633-79 (10); Huetamo, densely microfoveolate, each foveola with recumbent seta 27-11-1934, Hinten 5701, Lysiloma divaricata (1). arising from middle, frontal carina prominent, frons microru-

13 gose with foveolae crowded and convergent; clypeus irregu- Durango, 7800', 18-VII-1964, Powell (2); 24 mi E El Salto, larly microfoveolate except apex polished; postocular lobe 7-VI-1967, 1575-BR, Moldenke, on Acacia farnesiana (2); narrow; antenna capitate, eccentric from fifth segment, club Palos Colorados, 8000', 5-VIII-1947, Michener and Schra- segments nearly 2 x as wide as long, terminal segment mel (2); 9 mi W Durango, 9-V-1964, 1607-BR, Moldenke, ovate. Pronotum campaniform (fig. 86), lateral margins sin- on Ouercus undulata (1); 11 mi W Durango, 7000', uate, disk convex with subbasal, rounded gibbosity either 23-VI-1964, McAlpine (1). Chihuahua, 15 mi E Cuauhte- side at basai one-third, disk appearing slightly constricted moc, 6000', 11-VII-1964, Chemsak et al., at blacklight (1). apicad of gibbosities, middle of disk microfoveolate, depres- Distrito Federal, 30 mi SE Mexico City, 1-VII-1954, Kissin- sions irregularly spaced; lateral carina obtuse, becoming ger (1); Valley of Mexico, 18-IX-1896, ex herbarium sheet obsolete at one-third distance from procoxal fossa; cervical Echinopepon milleflorus, C.C. Pringle (1). Mexico, km 17 sulcus short, deep, nearly hidden in vestiture; prosternum Toluca Road, 11-VII-1934, Plummer (1). TIaxcala, 8 mi short, not separating procoxal apices; mesosternum sub- WNW Apizaco, 8200', 18-VI-1961, on flowers Heterotheca triangular, truncate apically, postmesocoxal sulcus narrow chrysopsides (1). following contour of coxal cavity. Scutellum transverse, bidentate, emarginate apically. Elytra together (fig. 86) longer Holotype, allotype, and paratypes deposited in National Mu- than wide, disk depressed in middle one-third; striae regu- seum of Natural History, Washington, DC. Paratypes also lar in course, stria 1 arising in scutellar depression, stria 2 deposited in American Museum of Natural History, New from pit beneath oblique ridge, striae 3 and 4 from closely York; Canadian National Collection, Ottawa; California In- set flat denticles atop slight gibbosity, striae 5 and 6 from sect Survey, University of California, Berkeley; Snow Mu- minute basal denticles; striae free apically except striae 4 seum, Lawrence, Kansas; Naturhistoriska Riksmuseet, and 5 sometimes conjoined; intervals flat, third and fifth Stockholm, Sweden; and CD. Johnson collection, Flagstaff, wider than others, surface finely imbricate. Metacoxa Arizona. densely, finely punctate except for glabrous band on anterior margin, pectén of metafemur (fig. 90) with one long, Distribution. MEXICO: Durango, Chihuahua, TIaxcala, Dis- slender subapical denticle separated from succeeding two trito Federal, Mexico. or three denticles by slight gap; metatibia (fig. 90) strongly bent at base, apical two-thirds nearly straight, gradually ex- Biology. Of all the plant names given in the paratype lists, panded base to apex, muero short, scarcely longer than only Pithecellobium leptophyllum is a rearing record and lateral denticle; ventral, lateral, and dorsomedial carinae was found on a herbarium sheet in the National Museum of complete, lateroventral carina abbreviated. Abdomen with Natural History; the others are probably pollen feeding first sternum slightly longer than remaining sterna together records. Apparently only one beetle develops in a seed. in 5, subequal in 2; fifth sternum broadly emarginate in 5, Pods are rather thin walled and adults emerge through cir- nearly evenly rounded in 2 but with submarginal carina cular exits cut in the pod wall. slightly lobed either side of pygidial apex, polished spot on lateral area of first and second abdominal sterna (fig. 89). Discussion. This species is distinguished from all other Pygidium of 5 convex, reflexed into emargination of termi- members of the genus by the polished areas on abdominal nal sternum, of 2 nearly flat, disk in both sexes densely sterna 1 and 2 and by details of the internal sac armature microfoveolate, surface nearly obscured by vestiture in of the male genitalia. male. Male genitalia: Median lobe about three times as long as width across apex (fig. 87), ventral valve arcuate, The specific name is derived from polio (Latin) = polished dorsal valve truncate, membranous; internal sac with one and refers to the polished area on the abdominal sterna. large forked sclerite at basal one-third and two slender, tapered spicules at middle, basal one-fourth of sac lined with minute, blunt denticles, apical one-third lined with minute, acute denticles, apical closure valve circular. Lateral lobes (fig. 88) slightly bowed, perceptibly expanded apically, cleft three-fourths their length.

Body length 4.0-4.9 mm, width 2.6-2.9 mm; pronotal length 1.1-1.5 mm, width 1.7-1.9 mm.

Holotype 5. MEXICO: Durango, Durango, IV-XI-1896, E. Palmer 338, seeds of Pithecellobium leptophyllum. Allotype 2 and six paratypes, same data. Other paratypes: MEXICO: (No State given), in seeds Bauhinia (3). Durango, 21 mi W

14 Vacillator Group

The vacillator group is distinguished by the following combi- Distribution. U.S.A.: Texas. MEXICO: Chihuahua, Tamau- nation of characteristics: Integument usually dark red; pu- lipas, Guerrero, Durango, Morolos, Distrito Federal, Puebla, bescence yellow, especially on the pygidium; pygidium di- Jalisco, Oaxaca. morphic in some species; three prominent spots of yellow hairs on metepisternum; postocular lobe short; elytra longer Specific records. MEXICO: Durango, 14 mi W La Ciudad, than wide; basal elytral gibbosities prominent on striae 3 15-VI-64, Howden; 24 mi W Ciudad, 20-VI-1964, Howden. and 4; carinae flanking anal notch not prominent (except in Chihuahua, La Bufa Mine, 81 mi S Creel, 26-VIII-1950, R. vacillator)] lateral margins of pronotum straight or slightly Smith; Bacugachic, 27-VIII-1950, R. Smith. Morolos, Cuer- bowed; postmesocoxal sulcus angulate behind coxa; ventral navaca, 22-VI-1937. Tamaulipas, 25 mi N Monte valve broad, truncate in most species; internal sac with Nacimiento del Rio Frio, O'Brien. Distrito Federal, La Venta, large, wishbone-shaped sclerite; lateral lobes bowed except VI. Guerrero, Tejupiico, Temascaltepec, 4000', 1932, Hinton in porphyreus and cristoensis; all known host associations and Usinger; Real de Arriba, Temascaltepec, 6-7000', with Lysiloma spp. Six species. 1932. Puebla, Puebla, V Jalisco, 3 mi S Tecolatlan, 1-1-1973, 4000', ex seeds Lysiloma divaricata, CD. John- This group appears to be most similar to the terani group. son 3-73. Oaxaca, 9 mi E El Cameron, 27-VI-1971, O'Brien. Merobruchus vacillator (Sharp) Bottimer (1968) reported vacillator from Big Bend, Texas, (Figs. 126-130) and I have confirmed that identification.

Bruchus vacillator Sharp, 1885:457; Pic, 1913:54. Biology. Nothing is known of the biology of this species Acanthoscelides vacillator: Blackwelder, 1946:761. except the host plant record. Merobruchus vacillator: Bottimer, 1968:1024; Johnson and Kingsolver, 1982:418. Discussion. Characters to separate M. vacillator Uom others in the group are the lack of distinctive setal patches on the Color. Integument reddish brown to deep purplish red with pronotum and lateral pronotal sclerites, the prominent ely- pale reddish variegation; antenna with basal segments red- tral gibbosities, two prominent processes on the 2 fifth ster- dish yellow, apical four segments contrastingly dark red, num, and conformation of the internal sac sclerites (cf. figs. terminal segment sometimes reddish yellow; forelegs and 26, 94, and 127). midlegs reddish brown. Pubescence of mostly yellowish gray setae in vague pattern on pronotum and elytra (fig. The only known host plant, Lysiloma divaricata, is used lo- 126) except for pair of usually prominent spots on disk of cally in Mexico and Central America for tanbark. pronotum; scutellar and metepisternal spots yellow; pygidial setae yellow in pattern illustrated (fig. 130). Merobruchus porphyreus, new species

Structure. Essentially as for M. porphyreus except basal (Figs. 93-97) striai gibbosities more prominent, 2 fifth sternum shallowly emarginate, emargination limited by prominent lateral proc- Color. Integument reddish brown to deep purplish red with esses, and in details of 5 genitalia. Male genitalia: Median paler red variegation; antenna yellowish red to piceous; lobe short (fig. 127), slightly more than 2 x as long as forelegs and midlegs yellowish, occasionally with piceous width across apex; ventral valve transverse, gently arcuate, suffusion, hindlegs reddish brown to piceous, all tarsi usu- without median process; internal sac with wishbone sclerite ally yellowish. Pubescence of gray, black, and yellow hairs; at base, middle of sac with paired, broad-based, hollow head with fine yellow setae, pronotal disk with sparse gray spines, apical one-half of sac sparsely lined with slender on intervals between foveolae, each foveola with centrally spicules, apex with arched, slightly sclerotized structure, placed fine black seta, anterior flank of pronotum with and terminal closure ring. Lateral lobes slender (fig. 128), patch of yellow hairs, middle of basal lobe with few yellow less bowed than in cristoensis and porphyreus. hairs; scutellum yellow, yellow patches on mesepimeron, middle and caudal margin of metepisternum, and lateral Body length 2.7-3.3 mm, width 1.6-1.9 mm; pronotal margin of abdominal sterna 2-5; venter of body otherwise length 0.9-1.1 mm, width 1.2-1.4 mm. gray; legs gray except tarsal pads yellow; pygidium with variegated pattern of yellow and black setae, usually with Type locality. MEXICO: Guanajuato. vague, paired dark spots near apex (fig. 97).

Holotype depository. British Museum (Natural History), Structure. Body subfusiform, subdepressed dorsally (fig. London. 93). Head turbiniform, frons and clypeus densely, finely fo-

15 veolate, each foveola with centrally placed seta, frontal ca- Holotype, allotype, and paratypes deposited in National Mu- rina obtuse, postocular fringe narrow; antenna gradually seum of Natural History, Washington, DC. F^aratypes also clávate from fourth segment, terminal segment elliptical. deposited in Canadian National Collection, Ottawa; Califor- Pronotum campaniform (fig. 93), lateral margins straight, nia Academy of Sciences, San Francisco; J.E. Wappes col- basal lobe prominent, shallowly sulcate; disk convex, with- lection, Chadds Ford, Pennsylvania; and CD. Johnson col- out major asperities, slightly depressed near posterolatêral lection. Flagstaff, Arizona. angles, surface of disk densely, irregularly, finely foveolate, intervals finely rugose; lateral margins of pronotal disk Distribution. MEXICO: Jalisco, Sinaloa, Durango, Veracruz, marked by poorly defined, obtuse ridge, cervical sulcus in- Chiapas. distinct, mtercoxal process barely separating procoxal apices. Scutellum quadrate, deeply incised apically. Postmeso- Biology. Only two host plants are recorded here for porphy- coxal sulcus strongly angulate. Elytra together (fig. 93) reus, Lysiloma acapulcensis and L. kellermani. Nothing else slightly longer than wide, subdepressed, stria 1 arising near is known of the life history of this species. scutellar apex, 3 and 4 from low gibbosity surmounted by two blunt denticles, 5 and 6 from basal pits, all striae free Discussion. The characters to separate M. porphyreus from apically except 6 and 7 conjoined; striai lines shallow, regu- others in the vacillator group are the bright yellow setal larly punctate; intervals flat, finely imbricate. Forelegs and patch on the pronotal flank, on the mesepimeron, and in midlegs not modified; metacoxal face densely punctate; the middle and on the caudal margin of the metepisternum, pectén of metafemur with one long denticle separated by the short apicomedian line of setae on the disk of the pro- gap from one or two shorter denticles (fig. 96); metatibia notum, lack of apical processes at the apex of the 2 fifth (fig. 96) with ventral, lateral, and dorsomesal carinae com- sternum, and wishbone sclerite skewed and paired median plete, lateroventral carina obsolete apically; muero short, spines absent in the 5 genitalia. acute, lateral denticle barely evident, coronal denticles two or three. First abdominal sternum as long as remaining The specific name is derived from porphyra (Gr. = purple) sterna together; fifth sternum broadly emarginate in 5, more alluding to the reddish-purple integument. narrowly emarginate in 2, and lacking lateral carínate processes flanking emargination. Pygidium oblique in 2, more Lysiloma acapulcensis is a source of plant gums. The bark convex and slightly reflexed apically in 5, discal surface is astringent and is used in local medicine in Mexico. densely, finely imbricate in both sexes. Male genitalia: Me- dian lobe (fig. 94) about 3 x as long as apical width, ven- Merobruchus cristoensis, new species tral valve short, broad, apical border somewhat angulate, tip rounded, dorsal valve membranous with brush of hairs (Figs. 25-29) apically; internal sac lined with fine denticles in basal one- half, fine setae in apical one-half, middle of sac with large, Color. Integument reddish purple, pygidium, antennae, fore- wishbone-like, skewed sclerite with attenuated flattened legs, and midlegs reddish yellow. Vestiture mostly yellowish arms, acute lateral process at fork. Lateral lobes bowed (fig. gray, vaguely condensed into patches (fig. 25); pronotal 95), attenuate apically. disk with thin median line of setae; pygidium (fig. 29) with yellow setae nearly obscuring disk, inconspicuous median Body length 2.7-3.5 mm, width 1.4-2.1 mm; pronotal stripe of condensed hairs constricted at middle. length 1.5-2.2 mm, width 1.1-1.3 mm. Structure. Similar to porphyreus except for distinctive 5 Holotype ¿. MEXICO: Jalisco, 3 mi E Tuxcueca, ca 5700', genitalia. Pectén of metafemur with distinct gap between 5-III-1979, ex seeds Lysiloma acapulcensis, Johnson long proximal denticle and two distal denticles. Male genita- 606-79. Allotype 2 and four paratypes, same data. Other lia: Median lobe (fig. 26) 2y2 times as long as width across paratypes: MEXICO: Sinaloa, 15 mi W El Palmito, 8, 17, 19, apex, ventral valve as illustrated, similar to that of porphy- 28-V-1964, Howden (10); 18 mi W El Palmito, 11-VII-1964 reus; armature of internal sac with flat, wishbone-shaped (1), 1-VIII-1964, Howden (1); 8 mi W El Palmito, sclerite near base of sac, sclerite not skewed, and pair of 11-VII-1964, Howden (1); (Mexico, no locality). El Paso bor- slender spines at middle of sac, apex of sac lined with ex- der station, 14-V-1976 (2). Tamaulipas, 1 mi E Gomez Fa- tremely fine spicules. Lateral lobes as illustrated (fig. 27), rias, 26-X-1979, Wappes (2). Durango, San Antonio, 5000', nearly identical to those of porphyreus. 10-VI-1937, Van Dyke (3); 24 mi W La Ciudad, 20-VI-1964, 7-VII-1964, Howden (2); 11 mi SW El Salto, Body length 3.4 mm, width 1.9 mm; pronotal length 1.0 9-VI-1964, Howden (2). Veracruz, Jalapa (1). Chiapas, Ar- mm, width 1.4 mm. riba Encanada, 8-1-1950, ex Lysiloma kellermani, Miranda 5888 (2).

16 Holotype J. EL SALVADOR: Monte Cristo, 23 km N Meta- of depression; lateral ridge sinuate, extending one-half way pan, 2800 m, 8-10-V-1971, H.F. Howden. from posterolateral corner toward procoxal cavity; cervical sulcus short, prominent; prosternum not separating pro- The unique type is deposited in Canadian National Collec- coxal apices; mesosternal apex truncate, postmesocoxal tion, Ottawa, Ontario. sulcus strongly angulate. Scutellum subquadrate, deeply emarginate apically. Elytra together slightly longer than Distribution. El Salvador. wide (fig. 53), evenly convex except slightly depressed behind scutellum; striae normal in course, striae 3 and 4 each Biology. Nothing is known of the host plants or habits of arising from small, blunt denticle on low basal gibbosity; all this species. striae free apically, 4 and 5 abbreviated; intervals flat, finely imbricate. Metacoxa finely, discretely punctulate; pectén of Discussion. The principal characters that distinguish M. metafemur (fig. 56) with one large denticle separated by cristoensis are the fine median line of setae on the pronotal gap from two or three progressively smaller subapical denti- disk, absence of a yellow setal patch on the pronotal flank, cles; metatibia (fig. 56) strongly arcuate; ventral, lateral, and grayish yellow setal patches on the mesepisternum and met- dorsomesal carinae complete, lateroventral carina obsolete episternum, the flat, not skewed, wishbone sclerite, and the in apical one-sixth; muero short, acute, lateral denticle pair of acute median spines in the male genitalia. short, coronal denticles three or four. First abdominal sternum subequal in length to remaining sterna combined, fifth The specific name is derived from the type-locality. sternum emarginate. Pygidium nearly flat in basal one-half, strongly convex in apical one-half, apex truncate in 5, Merobruchus lineaticollis (Sharp) rounded in $. Male genitalia: Median lobe apically broad (fig. 54), ventral valve transverse, slightly arcuate apically, (Figs. 53-57) dorsal valve membranous; internal sac with elongate wishbone sclerite, interior of sac lined with minute setae visible Bruchus lineaticollis Sharp, 1885:477; Pic, 1913:31. only under high magnification, apex of sac with patch of Acanthoscelides lineaticollis: Blackwelder, 1946:760. ciliate scales and circular closure valve. Lateral lobes as Merobruchus lineaticollis: Johnson and Kingsolver, illustrated (fig. 55). 1982:418. Body length 2.7-3.1 mm, width 1.5-1.7 mm; pronotal Color. Integument reddish brown to piceous; basal fifth and length 0.7-0.8 mm, width 1.0-1.3 mm. terminal segments of antenna, forelegs, and midlegs yellowish brown, pygidium mostly dark brown. Vestiture of yellow- Type locality. GUATEMALA: Panjachel, 5000'. ish gray and dark brown setae, those on pronotum mostly dark brown with contrasting, intensely yellowish white me- Holotype depository. British Museum (Natural History), dian stripe (fig. 53); elytra with second interval yellowish London. nearly to apex, third interval yellowish but interrupted by two dark brown spots, apical one-half of elytra with vaguely Distribution. MEXICO: Chiapas. GUATEMALA: Salama. defined, oblique, grayish yellow band, median transverse band vaguely defined; pygidium dark brown in basal three- Specific records. MEXICO: Chiapas, 22 mi N fourths with median stripe and lateral margins yellowish Ocozocoautia, 2-VII-1969, Bright and Campbell; 5 mi E El (fig. 57); venter of body densely clothed with yellowish gray Bosque, 6-VII-1969, Campbell and Bright; 8 mi S Simojo- setae. vel, 10-VI-1969, Campbell.

Structure. Body subfusiform, subdepressed dorsally. Head Biology. No host data are available for this species. turbiniform, eye with postocular fringe narrow; frons with narrow median carina, vertex finely, densely cribrate, frons Discussion. Only four specimens of this species have been more coarsely punctate, clypeus coarsely punctate to seen. It is placed in the vacillator group by virtue of the strigate, antenna with segments 1-4 slender, 5-10 eccen- shape of the ventral valve and internal sac sclerites and by tric, 11 elliptical. Pronotum campaniform (fig. 53), lateral the red integument and yellow pubescence, but it is easily margins slightly sinuate, disk evenly convex except slightly distinguished from other members of the group by the dis- depressed near posterolateral corners, and perceptibly tinct median stripe of yellow setae on the pronotal disk and channeled on basal lobe; surface of disk densely microfo- pygidium and again by characters in the male genitalia. It is veolate, with foveolae generally separated by less than one- illustrated in color in Sharp's treatment in the Biología half diameter, on flanks separated by more than 1 diameter, Centrali-Americana (vol. 5, plate 26, fig. 12). each foveola with fine, dark brown seta arising from center

17 Merobruchus knulli (White) and dorsomesal carinae complete from base to apex, muero short, broad, coronal denticles three or four. Abdomen (Figs. 47-52) of Í with fifth sternum broadly emarginate with fine carina on margin, slightly flared either side of pygidial apex, 2 with Bruchus knulli White, 1941:189. emargination shallow but flared carina more prominent; 5 Mylabris knulli: Blackwelder and Blackwelder, 1948:44. pygidium (fig. 51) subtriangular, 2 subovate (fig. 52), apical Merobruchus knulli: Johnson, 1968:1269; Bottimer, one-half somewhat more attenuated in 5 than in 2; sculpture 1968:1024; Johnson, 1979:123, 1981b:251; Johnson and mostly hidden by vestiture in 5, minutely pustulate in 2, Kingsolver, 1982:418. each pustule a modified setal insertion. Male genitalia: Me- dian lobe (fig. 48) broad, about 3 x as long as width Color. Integument reddish brown, eyes black, apical one- across ventral valve, base broad, emarginate, ventral valve third of elytra, especially in 5, usually clouded with dark truncate, dorsal valve membranous, somewhat angulate, brown or piceous (fig. 47); 5 pygidium with small, irregularly internal sac with large, skewed wishbone sclerite near base rounded median spot divided by narrow line of setae and of sac, with pair of brushlike setal patches near middle, vague diagonal bands (fig. 51), 2 pygidium with large, ringlike closure valve at apex. Lateral lobes elongate (fig. brown, bare cordate patch (fig. 52); terminal four antennal 49), setose. segments often dark brown. Vestiture of pale yellow, gray, dark brown, and black slender setae; head and pronotum Body length 3.2-4.1 mm, width 1.9-2.6 mm; pronotal with mostly yellow setae, few gray intermixed setae, foveo- length 0.9-1.4 mm, width 1.4-1.9 mm. lae set with black setae; elytra in both sexes with mostly yellow setae in basal one-half, 2 with mixed yellow and gray Type locality. Huachuca Mts., Arizona, 20-VII-1936, J.N. apically and laterally, 5 with darker apical areas of mostly Knull. black with some gray intermixed; venter of body gray except for three yellow patches on mesepisternum and metepister- Holotype depository. Ohio State University, Columbus. num, and scattered yellow setae along dorsolateral margins of abdomen; legs gray. Distribution. U.S.A.: Arizona. MEXICO: Jalisco, Michoacán, Morolos, Oaxaca, Sonora. HONDURAS: Comayagua. GUA- Structure. Body subfusiform, subdepressed above, moder- TEMALA: El Progreso. EL SALVADOR: San Vicente. ately deep. Head turbiniform; vertex, frons, and clypeus coarsely punctate to foveolate, frontal carina distinct but Specific records. U.S.A.: Arizona: Pima Co., Rincón Mts., with margins somewhat irregular, labrum sparsely punctate, Chimenea Cyn., in Lysiloma microphylla var. thornberi, CDJ setose; antenna eccentric from fifth segment, gradually 38-76. MEXICO: Sonora, Sierra de Alamo, in Lysiloma wat- clávate, terminal segment subelliptical. Pronotum (fig. 47) soni, Gentry 4787; 45 mi S Nogales. Jalisco, 12 mi NW Jo- campaniform, disk nearly evenly convex except shallowly cotepec, in Lysiloma acapulcensis, CDJ 342-73; nr concave near posterolateral angles and on either side of Tuxcueca, in Lysiloma acapulcensis; nr Atenquique, in Lysi- basal lobe, lobe shallowly sulcate, discal surface closely, loma acapulcensis, CDJ 357-73. Morelos, Cuernavaca. Mi- irregularly foveolate, each foveola with centrally placed choacán, Uruapan (no locality). Oaxaca, 15 mi S Sola de black seta, interspaces micropunctate, each puncture with Vega, Howden. GUATEMALA: El Progreso, 17 mi W Sana- yellow seta; flanks of pronotum arcuately carínate in middle rate, in Lysiloma acapulcensis, CDJ 1953-80. EL SALVA- one-third; cervical sulcus short, deep; prosternum barely DOR: San Vicente, in Lysiloma acapulcensis, Standley separating apices of procoxae. Scutellum subquadrate, bifid 21650. HONDURAS: 22 km SE Comayagua, in Lysiloma apically, set with yellow setae. Elytra together (fig. 47) about acapulcensis, CDJ 2102-80. as long as wide; subdepressed around scutellum, surface transversely imbricate on intervals, striae well marked by Biology. Three host plants are known for this species: Lysi- narrow suici, foveolate; stria 1 arising from posterior angle loma microphylla var. thornberi in Arizona, L. watsoni in So- of scutellum, stria 2 from deep basal pit, 3 and 4 from nora, and L. acapulcensis in Jalisco, Guatemala, El Salva- prominent bidentate gibbosity, 5, 6, and 7 from basal pits, dor, and Honduras. striae free apically except 4 and 5 abbreviated and conjoined; intervals subequal in width. Mesosternal apex Discussion. This species closely resembles M. rounded, margins subcarinate; postmesocoxal suIci meeting xanthopygus. Salient characters are the simply punctate on midline, slightly angulate caudad of coxae; metacoxal pronotal disk, 2 pygidium with a small median bare spot, 5 face densely punctate; pectén of metafemur (fig. 50) with pygidium immaculate or with a very small, irregular median one long slender denticle and two or three shorter succeed- spot, and 5 genitalia with a semicircular wishbone sclerite. ing denticles; metatibia strongly arcuate basally, nearly straight in apical three-fourths, lateral, ventral, lateroventral. Lysiloma acapulcensis is a source of gums and local medicines.

18 Merobruchus xanthopygus, new species foveolate, 2 pygidium with a large cordate spot, $ pygidium with a bisected median spot, and 5 genitalia with an angu- (Figs. 131-136) lar, usually skewed wishbone sclerite. These two species are extremely similar, and three series of rearings yielded Color. As for knulli except distal one-half of elytra darker both species. No specimens of xanthopygus have been col- than proximal one-half in most specimens (fig. 131); prono- lected north of Jalisco, whereas knulli is known from as far tum uniformly clothed with yellow setae; 5 pygidium (fig. north as Arizona. The temptation is strong to combine 135) almost uniformly yellow, occasionally with faint medio- these two species as a variable species, but the characters basal dart of pale yellow and small, median bare spot bi- appear to be clearly diagnostic. sected by dart; S similar to that of 5. The name xanthopygus, which is derived from xanthos Structure. As for knulli except for following differences: Pro- (Gr. = yellow) and pygus (Gr. = rump), refers to the yellow notum sparsely, simply punctate, not densely foveolate; 5 pygidium. genitalia with wishbone sclerite nearly semicircular, flat, not skewed (fig. 132). Lateral lobes elongate (fig. 133), bowed toward apices. Hindleg as in figure 134.

Body length 3.8-4.0 mm, width 2.1-2.5 mm; pronotal length 1.2-1.3 mm, width 1.7-1.9 mm.

Holotype 5. MEXICO: Jalisco, 3 mi E Tuxcueca, ca. 5700', 5-III-1979, ex seed Lysiloma acapulcensis, CD. Johnson 606-79. Allotype 2 and 26 paratypes, same data. Other paratypes: MEXICO: Jalisco, Chápala, 18-VIII-1949, Bottimer 125T (1); Chápala, 6 mi W Jalisco, 30-VI-1963, Doyen (1); 12 mi NW Jocotepec, ca. 5500', 31-X-1974, ex seeds Lysi- loma acapulcensis, Johnson 342-73 (4). Morelos, Cuerna- vaca, June (2). Guerrero, Azula, 6000', Smith (1); 28 mi N Chilpancingo, 19-VII-1974, D. Chandler. Mexico, Temascal- tepec. Real de Arriba, 6-7000', V-VII-1933, Hinten and Usinger (2); Temascaltepec, Tejupilco, 4000', 1932, Hinten and Usinger (1). Oaxaca, Tamazulapan, 7-8-VI-1967, Flint and Ortiz (1); Yojoa Lake, 1-IX-1972, Rolston (1). GUATE- MALA: 17 km W Sanarate, El Progreso, 15-111-1980, ex seeds Lysiloma acapulcensis, Johnson 1953-80 (2). HON- DURAS: 22 km SE Comayagua, 28-111-1980, ex seeds Lysi- loma acapulcensis, Johnson 2102-80 (7).

Holotype, allotype, and paratypes deposited in National Mu- seum of Natural History, Washington, DC. Paratypes also deposited in British Museum (Natural History), London; Cal- ifornia Academy of Sciences, San Francisco; California In- sect Survey, Berkeley; Louisiana State University, Baton Rouge; Canadian National Collection, Ottawa; and CD. Johnson collection, Flagstaff, Arizona.

Distribution: MEXICO: Jalisco, Morelos, Mexico, Oaxaca. GUATEMALA: El Progreso. HONDURAS: Comayagua.

Biology Only one host plant, Lysiloma acapulcensis, has been recorded for this species.

Discussion. The characters in the key and in the preceding description should suffice to distinguish this species from M. knulli. The salient characters are pronotal disk irregularly

19 Teraní Group

The terani group with one known species is most similar to of 5 deeply emarginate with lateral margins of notch slightly the vacillator group but with the following differences: Elytra elevated and carínate, of 2 with lateral margin more promi- quadrate, not longer than wide; postocular lobe short; anal nently elevated and angulately carinate; pygidium as in fig- notch of female flanked by prominent processes; lateral ure 120. Male genitalia: Median lobe (fig. 117) broad api- margins of pronotum concave; metepisternum with promi- cally, base emarginate; ventral valve broad, arcuate; nent patches of yellow hairs; lateral lobes slightly bowed; all internal sac with single wishbone-shaped sclerite, apex of known host plants of larva in the genus Acacia. sac with large patches of fine denticles. Lateral lobes as in figure 118. Merobruchus terani Kingsolver Body length 2.9-4.5 mm, width 1.9-2.5 mm; pronotal (Figs. 116-120) length 1.0-1.5 mm, width 1.3-1.7 mm.

Merobruchus sp. 4: Johnson, 1979:123. Type locality. COSTA RICA: Gste. Prov., Santa Rosa N.R Merobruchus terani Kingsolver, 1980:256; Janzen, 1980:936, 948; Johnson and Kingsolver, 1982:418. Type depository. In National Museum of Natural History, Washington, DC. For detailed description, see Kingsolver, 1980:256. Distribution. U.S.A.: Texas. MEXICO: Nuevo Leon, Guer- Color. Integument deep red to piceous, prosternum, mes- rero, Jalisco, Puebla, Oaxaca, Yucatan, Campeche. GUATE- osternum, and metasternum piceous; forelegs, midlegs, MALA: Zacapa. HONDURAS: Yoro, La Paz. COSTA RICA: and proximal five or six and terminal segments of antennae Guanacaste. reddish yellow, middle segments of antennal club sometimes dusky. Vestiture of yellowish orange, gray, and dark Specific records since 1980. U.S.A.: Texas: Cameron Co., brown recumbent hairs, those on head, pronotum, and py- Sabal Palm Grove Sane, Southmost (new U.S. record). gidium orange; elytra light orange, gray, and brown, with MEXICO: Nuevo Leon, nr. Linares, in Acacia berlandieri. broad, transverse orange band and basal spots orange, Jalisco, 56 mi S Puerto Vallarta, in Acacia sp., CDJ 446-73. apical one-third mottled with orange, brown, and gray hairs Oaxaca, 75 mi SE Oaxaca, in Acacia angustissima, CDJ intermixed; scutellum orange; body beneath mostly gray 225-68; 32 mi NW Tehuantepec, in Acacia picachensis, with orange spots on mesepimeron, metepisternum, and on CDJ 566-79. Yucatan, nr. Uxmal, in Acacia tenuifolia, CDJ lateral margins of fourth and fifth abdominal segments; py- 1677-80. Campeche, 15 km E Campeche, in Acacia tenui- gidium (fig. 120) with densely placed orange vestiture, dark folia, CDJ 1658-80. GUATEMALA: 23 km SW Rio Hondo, in areas often with brown hair but usually bare; legs ciner- Acacia picachensis, CDJ 1939-80. HONDURAS: Nr. La eous, apices of protibia, mesotibia, and tarsal pads Paz, in Acacia gaumeri, CDJ 878-79. yellowish. Biology. This species has been reared from the following Structure. Body ovate (fig. 116), broadest at middle of ely- host plants: Acacia angustissima, berlandieri, gaumeri, pica- tra. Head short, subtriangular; vertex finely, densely punc- chensis, and tenuifolia. It has been collected in January, tate, punctures on frons and clypeus somewhat coarser, March, and May to September. deeper, and approximate, sometimes merging, each puncture setose, frontal carina prominent but not sharp, slightly Discussion. Merobruchus terani is perhaps most similar to expanded dorsally; postocular lobe narrow, setose; antenna vacillator in the vacillator group, but the elytra of terani are reaching posterolateral corner of pronotum, strongly clávate nearly quadrate and the pronotal lateral margins are con- from 5th segment, 11th subtriangular. Pronotum campani- cave, whereas the elytra of those species in the vacillator form (fig. 116), convex, subdepressed basally, posterolateral group are longer than wide, and the pronotal margins are corners strongly depressed, depression delimited laterally straight or slightly bowed. The bright yellow vestiture, dark by oblique, obtuse ridges, basal lobe shallowly sulcate; disk red integument, broad truncate ventral valve, and flat wish- densely foveolate, each foveola round and setose, punc- bone sclerite in the internal sac are associating characteris- tures generally crowded, separated by less than diameter of tics between the two groups. puncture, sometimes coalescing; cervical sulcus short, deep; prosternum barely separating procoxal apices. Striae Members of the vacillator group are associated with species of elytra shallow, deepening toward apex, 2, 5, and 6 aris- of Lysiloma, whereas terani is known only from species of ing basally from deep pits, 3 and 4 from prominent, biden- Acacia. tate subbasal gibbosity; intervals minutely strigose; postmesocoxal sulcus strongly angulate. Mesocoxal face densely punctate, hindleg as in figure 119. Fifth abdominal sternum

20 Insolitus Group

The pattern of pubescence on the pygidium and elytra is The insolitus group is characterized by the following in similar to that of M. boucheri but is bright yellowish orange, combination: Small relative size (2.2-3.3 mm); elytra subde- whereas that of boucheri is grayish yellow. Comparison of pressed (except placidus and solitarius); postocular lobe the male genitalia (figs. 10 and 117) shows fundamental narrow; metepisternum lacking prominent patches of hairs; differences in the form of the median lobe and armature of striae 3 and 4 with basal denticles; postmesocoxal line the internal sac. angulate (except paquetae); pygidium with basal triangular pad of white or yellowish hairs sometimes extending as nar- Acacia berlandieri is used as an ornamental shrub, is a row line toward apex or broken into median segments; fe- good honey plant, and is a potential source of commercial male sometimes with well-developed lateral flanges or proc- plant gum. esses flanking anal notch; male genitalia with median triangular sclerite (wishbone) and pair of lateral, usually thornlike sclerites near apex of sac, some species with pair of spines near base of sac; lateral lobes broad, spatulate (except boucheri).

The 10 species in the insolitus group are separated into 4 subgroups: Insolitus, sonorensis, placidus, and boucheri.

Insolitus Subgroup

Characteristics of the insolitus subgroup are elytral pattern contrastingly marked, pronotum subconical with lateral margins straight, elytral disk subdepressed, pygidium with distinct pattern, processes flanking anal notch well developed. Three species.

Merobruchus insolitus (Sharp)

(Figs. 35-40)

Bruchus insolitus Sharp, 1885:476. Acanthoscelides insolitus: Blackwelder, 1946:759. Merobruchus sp.\ Gibson, 1972:148. Merobruchus insolitus: Johnson, 1979:122; Janzen, 1980:948; Johnson, 1981b:251; Johnson and Kingsolver, 1982:418.

Color. Integument dark red to piceous; head dark red, eyes black, antenna reddish yellow, sometimes with segments 8, 9, and 10 darker than remaining segments; pronotum red tending toward piceous; elytra red with piceous umbones (fig. 35), lateral borders usually dark in basal one-half, third interval with rounded dark spots at basal one-sixth and apical two-thirds, lateral intervals with variable individual dark spots, or spots merged; pygidium piceous with reddish median streak; body venter reddish except prosternum and mesosternum tending to be piceous. Vestiture of yellowish white, white, and brown slender hairs; head, pronotum, elytra, and venter of body yellowish white, brown elytral spots with brown hairs, pygidium (figs. 39 and 40) with yellowish white hairs sparsely distributed except for conspicuous, narrow, white, median "dart" usually extending about two-thirds from base, occasionally to apex, usually evenly attenuated, occasionally "pinched," or rarely interrupted.

21 Structure. Body subfusiform, subdepressed dorsally. Head Albizia occidentalism CDJ 141-75, and A. Iebbel<, CDJ with vertex densely punctulate, vestiture directed cephalad; 129-75. Sonera, Alamos, in Albizia sinaloensis', San Ma- frons more coarsely punctate, vestiture directed mesad, rino; Hermosillo; 9 mi N Guaymas, in Acacia occidentalis, frontal carina ridgelike, frons separated from vertex by CDJ 141-73; 10 mi N Guayamas, in Lysiloma divaricata, transverse sulcus; clypeus scabriculous; antenna slender, CDJ 220-78; Lake Mocuzari, in Lysiloma divaricata, CDJ extending beyond posterolateral corner of pronotum, serrate 131-76; Navojoa, in Pithecellobium sonorae, CDJ 151-75; from 4th segment, 11th elliptical. Pronotum (fig. 35) nr. Santa Maria, Escuela de Agricultura. Tamaulipas, Hwy. elongate-campaniform, lateral margins slightly sinuate, apex A-70, nr. Jet. 85; Ciudad Mante, ex ?Albizia; Gomez Farias. rounded; disk evenly convex except for slight depression on Nuevo Leon, Linares; between Linares and Iturbide, in Pi- basal lobe and at posterolateral corners; lateral carina thecellobium pallens; Tope Chico nr. Monterrey; Chipinque ridgelike, sinuate; cervical sulcus hidden in vestiture; pro- Mesa. Durango, San Antonio; El Salto; La Ciudad. Sinaloa, sternum barely separating procoxal apices; postmesocoxal Mazatlan, 30 mi S Guamuchil, in Lysiloma divaricata, CDJ sulcus strongly angulate. Scutellum wider than long, deeply 137-73. Jalisco, Chiquilistlan, in Lysiloma acapulcensis; emarginate, and bidentate. Elytra together (fig. 35) about as Tuxcueca, in Lysiloma divaricata, CDJ 606-79; Chamela wide as long, slightly depressed mesally; striae on disk nar- Biological Preserve, in Lysiloma divaricata. Michoacán, N of row and shallow, somewhat sinuate especially in basal one- Playa Azul, in Lysiloma divaricata, CDJ 1212-79; 3 mi E third, striae 3 and 4 arising from prominent, bidentate basal Tuxcueca, in Lysiloma acapulcensis, CDJ 606-79. Nayarit, gibbosity, striae free apically; intervals finely imbricate, third Pichen, nr. Acaponeta, in Pithecellobium aff. máncense, and fifth wider than others. Metacoxal face densely, evenly CDJ 482-73. Mexico, Tejupiico; Temascaltepee. Puebla, punctate; pectén of metafemur (fig. 38) with distinct gap Puebla (no date); Tehuaean; Atlixco. Oaxaca, Salina Cruz; between long denticle and four smaller denticles; metatibia 35 km W Zanatepee, in Pithecellobium undulatum, CDJ with all carina complete, lateroventral carina joining flanged 341-78; 22 km NW Puerto Escondido, in Lysiloma micro- ventral at base of muero, space between them canaliculate; phylla van thornberi, CDJ 1369-80; El Cameron; 93 km SE muero short, coronal denticles three. Fifth sternum of 5 ab- Oaxaca, in Lysiloma divaricata, CDJ 574-79. Chiapas, Ar- domen with broad, shallow emargination; 2 with narrow riaga; Arriba, Encanada, in Lysiloma kellermani; 20 mi S emargination, lateral flanges moderately prominent. Pygid- Comitan, in Lysiloma acapulcensis, CDJ 1041-79; Jet. ium in both sexes finely, densely punctate. Male genitalia: routes 190 and 195; Tuxtia Gutierrez. Yucatan, 7 km N Median lobe (fig. 36) about 3 x as long as wide; ventral Uxmal, in Acacia tenuifolia, CDJ 1677-80. Campeche, 15 valve semicircular with distinct apical nipple; internal sac km E Campeehe, in Acacia tenuifolia, CDJ 1658-80. GUA- with three sharply triangular sclerites at middle of sac. Lat- TEMALA: Barranquillo, in Pithecellobium pallens; 17 km N eral lobes as in figure 37. Sanarate, in Lysiloma acapulcensis, CDJ 1953-80; 20 mi SE Taxisco, in Lysiloma divaricata, CDJ 810-79; Baja Vera- Body length 2.9-3.3 mm, width 1.6-2.0 mm; pronotal paz. HONDURAS: Comayagua. COSTA RICA: Bagaces, in length 1.9-2.2 mm, width 1.2-1.5 mm. Lysiloma seemani; Santa Rosa N.R, in Albizia adinocephala; 8 km N Las Cañas, in Lysiloma sp., DHJ Vl~20-1972-lll; Type locality. GUATEMALA: Nr. Guatemala City. Finca La Pacifica, in Lysiloma seemani; Finca Taboga, in Lysiloma seemani. Holotype depository. In British Museum (Natural History), London. Biology. Host plants recorded for M. insolitus are in the genera Acacia, Albizia, Lysiloma, and Pithecellobium. See Distribution. U.S.A.: Arizona, Texas. MEXICO: Baja Califor- Appendix 3 for specific hosts. This species has the longest nia, Sonora, Tamaulipas, Nuevo Leon, Durango, Sinaloa, host list of any Merobruchus species. It was collected each Jalisco, Nayarit, Michoacán, Mexico, Puebla, Oaxaca, month except September. Merobruchus insolitus is parasit- Chiapas, Yucatan, Campeche. GUATEMALA: Jutiapa, Pro- ized by the braconid wasp Urosigalphus neobruchi Gibson gresse, Salama. HONDURAS: Comayagua. COSTA RICA: and by the chalcid wasp Chryseida bennetti Burks (Burks, Guanacaste. 1956).

Specific records. U.S.A.: Texas: Cameron Co., Brownsville, Host data labels in some collections listing Havardia brevifo- ex seeds Pithecellobium pallens, Harlingen, Bottimer 50E1; Ha are referable to Pithecellobium pallens. Hidalgo Co., Bentsen; Brewster Co., Big Bend N.R; Frio Co., Pearsall. Arizona: Pima Co., Chimenea Cyn., Rincón Discussion. The principal distinctive characters for this spe- Mts., in Lysiloma microphylla van thornberi. MEXICO: Baja cies are the dark red body, piceous elytral margins, white California (no locality), in Lysiloma candida; Baja California basal pygidial triangle elongated into an evenly tapered Sur, Sierra de la Giganta, in Lysiloma divaricata; nr. Los Di- dart, and, in the 5 genitalia, the apical nipple on the ventral visaderos; La Burrera; nr. Mulega; nr. San Jose del Cabo, in

22 valve, three subequal median sclerites, and the basal pair basal thornlike sclerites in hastatus, insolitus, and paquetae of small, thornlike sclerites. The apparently nearest relative are absent in the other four species.This is the only species of insolitus is paquetee. in the insolitus group in which the postmesocoxal sulcus is not angulate. Lysiloma acapulcensis is a local source of medicines and gums, and L divaricata is a source of tanbark. Merobruchus paquetae Kingsolver

Merobruchus hastatus Kingsolver (Figs. 70-74)

(Figs. 30-34) Merobruchus paquetae Kingsolver, 1980:252; Janzen, 1980:948; Johnson and Kingsolver, 1982:418. Merobruchus hastatus Kingsolver, 1980:250; Janzen, 1980:948; Johnson and Kingsolver, 1982:418. For detailed description, see Kingsolver, 1980:252.

For detailed description, see Kingsolver, 1980:250. Color. Integumental color dark red to piceous, forelegs, midlegs, and antennae reddish yellow, in some darker Color. Integument reddish to piceous; eyes black; forelegs, specimens with segments 8-10 piceous; eyes black; meta- metatibia, and usually apex of metafemur reddish yellow; femur mostly piceous with dorsal one-third dark red, prono- antenna with proximal 7 segments and 11 reddish yellow, tum with broad median stripe dusky; elytra with lateral mar- 8-10 piceous to black. Dorsal vestiture of grayish yellow gins usually clouded with piceous, sometimes with strong and dark brown hairs, ventral of silvery gray hairs; dark lateromedian, dark rounded spot; pygidium usually with brown hairs on piceous spots on pronotum, elytra, and py- paired, median, irregular dark areas showing through vesti- gidium in pattern in figure 30. ture, more prominent in darker specimens. Vestiture of yellowish gray, silvery gray, white, and dark brown slender Salient characters. Pronotal margins slightly sinuate (fig. hairs in pattern shown in figures 70 and 74, dark brown 30), vestiture nearly concealing punctation; elytral pattern hairs restricted to piceous areas; scutellum white; pygidium more contrasting in 5 than in 2; pygidium with dense white (fig. 74) with basal triangle and median lozenge-shaped basal triangle (fig. 34), usually with small, median, elongate mark yellowish gray to white, rarely with basal triangle con- white spot separated from basal triangle by black integu- nected to lozenge, paired dark, median patches encroach- ment; anal notch of 2 short, deep, lateral tufts prominent, ing on white patches; venter of body with evenly distributed that of S broad, nearly dividing fifth sternum; postmesocoxal silvery gray hairs. sulcus arcuate, not angulate; antenna with segments 8-10 piceous; hindleg as in figure 33. Male genitalia as in figures Salient characters. Pronotum with dense vestiture but with 31 and 32. discal foveae showing through vestiture, middle of disk darker than lateral areas, pronotum evenly arcuate in lateral Body length 3.25-3.75 mm, width 2.0-2.7 mm; pronotal aspect; antennae entirely yellow; elytra with moderately length 1.9-2.0 mm, width 1.4-1.5 mm. strong pattern of brown maculae; pygidium with basal triangle yellowish constricted by lateral, depressed dark spots; Type locality. COSTA RICA: Gste. Prov., Santa Rosa N.R postmesocoxal sulcus angulate; anal notch of 2 fifth sternum shallowly sulcate, slightly reflexed; 5 ventral emargina- Holotype depository. In National Museum of Natural His- tion nearly dividing sternum; postmesocoxal sulcus angu- tory, Washington, DC. late. Male genitalia as in figures 71 and 72.

Distribution. COSTA RICA: Gste., Santa Rosa N.R No addi- Body length 2.0-3.1 mm, width 1.1-1.9 mm; pronotal tional records are known. length 0.6-1.0 mm, width 0.9-1.4 mm.

Biology. This species has been reared from seeds of Lysi- Type locality. COSTA RICA: Gste. Prov., 1 mi W Tilaran. loma desmostachys, erroneously identified as Piptadenia flava in the original description (Kingsolver, 1980) and in Holotype depository. In National Museum of Natural His- Janzen (1980). No other hosts are known. tory, Washington, DC.

Discussion. Despite the resemblance of the ventral valve of Distribution. COSTA RICA: Guanacaste. NICARAGUA: Gra- hastatus to that of santarosae, sonorensis, lysilomae, and nada. PANAMA: Canal Zone. COLOMBIA: Magdalena, chetumalae, hastatus appears to be somewhat isolated as Valle del Cauca. VENEZUELA: Mérida. SURINAM. BRAZIL: evidenced by the bold elytral and pygidial patterns. The Rio de Janeiro, Mato Grosso.

23 Specific records since 1980. COLOMBIA: Antioquia, Sope- Salient characters. Color pattern muted; antenna with seg- tran, in Pseudosamanea sp. (new host record). SURINAM: ments 8-10 piceous, scutellum prominently white; pygidium (no locality) USDA Plant Quarantine interception, in ?Mi- with prominent, densely white basal triangle narrowly con- mosa sp. nected to median diamond-shaped white spot; postmesocoxal sulcus sharply angulate; ventral notch of 2 fifth ster- Biology. This species has been reared from the following num broad, lateral carínate flanges moderately prominent, host seeds: Albizia adinocephala, caribaea, lebbek (Rio de emargination in 5 broad, about one-half length of sternum, Janeiro), longipedata, Albizia sp. (Mato Grosso, Brazil), Lysi- slightly flanged; in lateral aspect, basal two-thirds of prono- loma sp., Pseudosamanea guachepele (Colombia), and tum plane, apical one-third arcuate. Male genitalia as in questionably Acacia sp. Collections have been made in figures 112 and 113. February, March, and July. Body length 2.9-3.5 mm, width 1.6-2.0 mm; pronotal Discussion. Characters sufficient to separate paquetae are length 0.9-1.2 mm, width 1.2-1.4 mm. the drab, yellowish gray appearance, entirely yellow antennae, yellowish gray pygidial stripe divided or nearly divided Type locality. COSTA RICA: Gste. Prov., Santa Rosa N.R by lateral integumental spots, the very shallow 2 anal notch, and 5 genitalia with the ventral valve arcuate, median Holotype depository. In National Museum of Natural History, sclerite wishbone shaped, and a pair of small, basal, thorn- Washington, DC. like denticles. The pronotal disk is often darker than in figure 70. Compare this species with the closely related Distribution. MEXICO: Sonora, Sinaloa, Jalisco, Nayarit, insolitus. Campeche. HONDURAS: Comayagua. COSTA RICA: Gua- nacaste. COLOMBIA. The extensive host list and broad geographical range are noteworthy. Specific records since 1980. MEXICO: Sonora, nr. San Carlos Bay, in Albizia, CDJ 874-79; 10 mi N Guaymas, in Sonorensis Subgroup Lysiloma divaricata, CDJ 1142-79 (new host record). Ja- lisco, Chamela Biological Preserve, in Albizia occidentalis Characteristics of this subgroup are internal sac lacking and A. tomentosa. Campeche, nr. Francisco Escarcega, in basal paired sclerites; ventral valve truncate; wishbone Albizia, CDJ 1619-80. HONDURAS: 11 mi SW Comayagua, sclerite broad-based, acute apically; dorsal pattern obscure in Albizia, CDJ 874-79. (except santarosae usually with black lateral maculae). Four species. Biology. The host plants for sonorensis include Albizia adinocephala, caribaea, lebbek, occidentalis, ortegae, Merobruchus sonorensis Kingsolver sinaloensis, tomentosa; Lysiloma divaricata, seemani; and Pithecellobium sonorae. Collections were made in March, (Figs. 111-115) April, July, and December.

Merobructius sonorensis Kingsolver, 1980:254; Janzen, Discussion. Despite the close resemblance of the male 1980:948; Johnson and Kingsolver, 1982:418. genitalia to those of santarosae (figs. 101 and 112), sonorensis is distinctive in the dark red integument with an indistinct elytral and pronotal pattern, prominent, elongate, white For detailed description, see Kingsolver, 1980:254. pygidial triangle, and broad anal notch of the 2 fifth sternum. See also the discussions of chetumalae, lysilomae, Color. Integument mostly dark red with some piceous ob- and santarosae. The wood of Albizia occidentalis is used in fuscation on head, pronotum, and pygidium, eyes black, carpentry in northern Mexico, and A. lebbeck, introduced antenna yellowish red with segments 8-10 dark, forelegs from Asia, is a useful shade and ornamental tree. and midlegs yellowish red. Vestiture of grayish yellow, dark brown and white hairs; head, pronotum, and venter of body Merobruchus santarosae Kingsolver grayish yellow; pronotum with vague, median, grayish yellow median stripe in most specimens; scutellum white; elytra (fig. Ill) grayish yellow except dark brown on dark integ- (Figs. 98-105) umental spots; pygidium (fig. 115) with dark brown hairs on piceous spots, basal triangle and median spot white, other- Merobruchus sjp. 1: Johnson, 1979:123. wise grayish yellow. Merobruchus santarosae Kingsolver, 1980:246; Janzen, 1980:948; Johnson and Kingsolver, 1982:418.

24 For detailed description, see Kingsolver, 1980:246. distinct from the sparse, dark brown pubescence of the other two species. See the discussion of chetumalae. Color. Integument deep red above with piceous maculae on elytra; venter of body usually somewhat darker especially Acacia coulteri is a local source of fuel in Mexico. on thoracic sterna; eyes piceous, antenna reddish yellow with segments 8-10 piceous; forelegs and midlegs reddish Merobruchus lysilomae, new species yellow, hindlegs usually dark red. Vestiture of yellowish gray and dark brown, slender, recumbent hairs, brown hairs only (Figs. 58-62) on piceous integumental maculae on elytra and occasionally on pronotum. Hindleg as in figure 103. Color. Integument dark reddish brown except antennae, forelegs and midlegs, and metatarsi yellowish; elytra mot- Salient characters. Pronotum (fig. 98) immaculate, densely tled with dark brown spots and vaguely defined transverse covered with pubescence concealing punctation, in lateral bars; pronotum usually with narrow yellowish red median aspect dorsal profile evenly arcuate; dark brown or piceous stripe. Vestiture of short yellowish gray hairs evenly distrib- lateral elytral maculae (figs. 98-100) prominent in most uted over body except in some dark elytral spots and dark specimens; pygidium (figs. 104 and 105) with basal triangle areas on pygidium; postocular spot and pronotal flanks with of setae of same color as those on remainder of pygidium, dense yellow setae; pygidium with vaguely defined to dis- not strongly differentiated, indistinct median dart usually tinct basal triangle of yellowish white setae delimited by constricted by dark integumental spots, apex of 2 fifth ster- dark integument. num with moderately long tufts, ventral notch deep; postmesocoxal sulcus strongly angulate. Male genitalia as in Structure. Body elongate-ovate, widest behind humeri, figures 101 and 102. somewhat depressed dorsally. Head turbiniform, frons slightly convex, some specimens with narrow, median gla- Body length 3.1-4.2 mm, width 1.9-2.1 mm; pronotal brous line; vertex finely, transversely punctate, frons and length 0.9-1.3 mm, width 1.3-1.4 mm. clypeus more coarsely punctate; postocular lobe narrow; antenna clávate from fifth segment, terminal segment ovate. Type locality. COSTA RICA: Gste. Prov., Santa Rosa N.P Pronotum campaniform in dorsal aspect (fig. 58), lateral margins nearly straight, apical margin evenly rounded, Holotype depository. In National Museum of Natural His- basal margin sinuate; disk slightly convex, discal surface tory, Washington, DC. with distinct and discrete microfoveolae, each bearing recumbent hair in its center, basal lobe with shallow median Distribution. COSTA RICA: Guanacaste. MEXICO: Sonora, sulcus; lateral carina obsolete, present only as intermittent Sinaloa, San Luis Potosí, Tamaulipas. HONDURAS: ridge; cervical sulcus short, deep; prosternum barely sepa- Comayagua. rating procoxae; postmesocoxal sulcus vaguely angulate. Scutellum short, broad, deeply emarginate. Elytra together Specific records since 1980. MEXICO: Sonora, nr. Navojoa, slightly longer than broad (fig. 58), widest at apical one- in Acacia coulteri, CDJ 179-73 (new host record); Lake third, subdepressed, especially around scutellum; striae Mocuzari, in Acacia coulteri, CDJ 191-77. Sinaloa, nr. Cu- narrow, shallow, regular in course, 3 and 4 arising basally liacan; Valle de Carrizo, Carranza River. HONDURAS: Com- from bidentate basal gibbosity; interspaces flat, finely imbri- ayagua, in Albizia sp., CDJ 874-79. cate. Metacoxal face finely, evenly punctate, metafemoral pectén (fig. 61) with four denticles on triangular lamina, Biology. Host plants recorded for this species are Acacia basal denticle 2 x as long as others; metatibia abruptly coulteri, Albizia sp., and Lysiloma desmostachys. It has bent at base, nearly straight and gradually broadened to- been collected from December to March and in May, July, ward apex; muero short, slightly longer than lateral denticle, and August. In the original description (Kingsolver, 1980), coronal denticles three, lateroventral carina obsolete api- the common name "palo amarillo" was listed, but since this cally, not joined to ventral carina at muero base. Terminal name can be applied to several plants in Mexico, no defi- sternum of 5 broadly emarginate to one-half its length, nite species can be indicated. broadly emarginate also in 2 but with lateral borders of emargination slightly prominent and carínate; 5 pygidium Discussion. Although details of the male genitalia of santa- oblique in basal two-thirds, strongly convex and reflexed rosae indicate a close similarity to sonorensis, lysilomae, apically, extreme apex truncate, S pygidium similar but apex and chetumalae, its external characters differ. The dense not reflexed, extreme apex rounded, disk in both sexes yellowish gray pubescence of saritarosae with dark brown finely strigose, strigae unevenly distributed. Male genitalia: or black lateral maculae in 90 percent of the specimens is Median lobe more than 4 x as long as wide (fig. 59), ventral valve broad, quadrate, slightly emarginate apically; in-

25 ternal sac with rows of fine, sharp denticles basally, middle Discussion. Characters sufficient to separate this species of sac with triangular median sclerite and two identical, lat- from others in the sonorensis group are the purplish red eral, thornlike sclerites, apex lined with fine denticles. Lat- integument, sparse body vestiture, poorly defined basal tri- eral lobes as illustrated (fig. 60). angle of the pygidium, ratio of pronotal width to length 1.3:1, paired median sclerites of the internal sac each with Body length 2.4-3.1 mm, width 1.4-1.7 mm; pronotal a broad base and with the lateral spine set at 90 degrees, length 0.8-0.9 mm, width 1.1-1.3 mm. wishbone sclerite elongate-triangular, and carínate processes flanking the $ fifth sternal notch prominent. Holotype 5. U.S.A.: Florida: Royal Palm Park, 9-IX-1931, L. Bottimer 63g. Allotype $ and nine paratypes, same data. The species is named for one of its host plant genera. Other paratypes: Same data as holotype except with dates IX-1931. 20-XI-1931, 1-XII-1931, Bottimer 63h (89); same Merobruchus chetumalae, new species data as holotype except IX-1931, 63i (3); Royal Palm Park, 13-111-1924, Blatchley (2); Key Largo, 5-XI-1964, S. Kemp, (Figs. 15-18) light trap (11); Upper Key Largo, 15-IX-1960, in Lysiloma prob, latisiliqua, Bottimer 108v (45); Upper Key Largo, Color. Integument dark purplish brown, forelegs and mid- 12-V-1972. 18-111-1972, Wappes (6); Key Largo, Tavernier, legs yellowish, metafemur sometimes reddish brown; an- 25-111-1975, Steiner (2); Dade Co., V-1953, Paulson (1); Ft. tenna yellowish with segments 8-10 reddish brown, seg- Lauderdale, 6-X-1963, McLean, in seeós Acacia simplicifo- ment 11 usually yellowish; 5 pygidium reddish brown, lia (10); Everglades N.R, Flamingo, 1-5-XII-1961, Munroe sometimes with lateral yellowish areas, 2 pygidium dark (1); Everglades N.R, Paradise Key, 31-111-1962, Woodruff brown. Vestiture of short, silvery gray, and brown hairs ar- (41); Coconut Grove, 9-IX-1944, in Albizia lebbek seed (1), ranged in pattern similar to that of lysilomae; pronotum of- 4-IX-1945, in Albizia polyphylla seed (1), 25-VII-1933, in ten with slender median line of gray hairs; flanks of prono- Acacia richii seed (1). CUBA: Cienfuegos, 9-VI-1933, in tum and of venter of body with vestiture dense; 5 pygidium Lysiloma latisiliqua seed (2); Habana, 23-11-1943, E.E.A. (fig. 17) evenly clothed with gray hairs except for short, Ent. 11158, in seeds "sabicu," Lysiloma sabicu (7); Cuba broad basal triangle extending to one-third from pygidial (no locality), 17-XII-1928, Gouldman 76381, in Lysiloma base; 2 pygidium (fig. 18) with narrow basal triangle of sil- seeds (336). HAITI: Port au Prince, 1925, Hoffman (1). BA- very gray hairs, in some specimens reaching nearly to HAMAS: Nassau, 31-1-1962, in fly trap (1); Mayaguana Is., apex, sometimes interrupted at basal one-third, remainder 3-IX-1963, Murvosh, light trap (1). of pygidium covered with dark brown hairs.

Holotype and paratypes deposited in Canadian National Structure. Similar to lysilomae, with following differences: Collection, Ottawa, Ontario; allotype and paratypes in Na- Ratio of pronotal width to length 1.24:1 in chetumalae (fig. tional Museum of Natural History, Washington, DC. 15), (1.31:1 in lysilomae, fig. 58); paired median sclerites of Paratypes deposited in Blatchley collection, Purdue Univer- internal sac more slender with spine projecting at about sity, W. Lafayette, Indiana; CD. Johnson collection. Flag- 30° in chetumalae (fig. 16) (sclerite broad based and with staff, Arizona; J.E. Wappes collection, Chadds Ford, Penn- spine at 90° in lysilomae, fig. 59); overall measurements sylvania; Division of Plant Industry, Florida Department of average smaller in chetumalae; carínate processes flanking Agriculture, Gainesville; and British Museum (Natural His- fifth sternal emargination in 2 not as prominent in chetuma- tory), London. lae. Male genitalia: Median lobe (fig. 16) 4.5 x as long as width across ventral valve; similar to lysilomae but with ven- Distribution. U.S.A.: Florida. WEST INDIES: Cuba, Haiti, tral valve longer, paired sclerites of internal sac elongated, Bahamas. and with lateral spine set at about 30°, not 90° as in lysilomae; wishbone sclerite much smaller than in lysilomae. Biology. This species has been reared from seeds of several host plants, including Lysiloma sabicu and latisiliqua, Body length 2.1-2.3 mm, width 1.2-1.3 mm; pronotal Albizia lebbek and polyphylla, and Acacia richii and simplici- length 0.6-0.7 mm, width 0.8-0.9 mm. folia. It was collected during June and July, September through January, and March. Holotype ¿. MEXICO: Quintana Roo, 46 mi W Chetumal, 28-XII-1978, reared from seeds Lysiloma sp., CD. Johnson Lysiloma latisiliqua is planted as an ornamental tree around 412-78. Allotype 5 and six paratypes, same data. Other par- homes and along highways, and L. sabicu is used in ship atypes: MEXICO: Campeche, 160 km E Francisco Escar- building in the West Indies. cega, 27-XII-1978, reared from seeds Lysiloma sp., CD. Johnson 411-78 (4). Yucatan, Chitzén Itzá, 10/11-VI-1983;

26 1 km S Xcalacoop, 11-VI-1983, Wappes (3); 13 km E Me- Structure. Body ovate, convex dorsally (fig. 75). Head turbi- rida, 10-111-80, Lysiloma latisiliqua, CDJ 1853-80 (2). niform, eyes with ocular sinus about five-eighths length of eye; postocular fringe narrow, densely setose; vertex Holotype, allotype, and paratypes deposited in National Mu- densely, finely punctulate, punctures becoming coarser and seum of Natural History, Washington, DC. Paratypes in CD. striated on frons; antenna serrate from fifth segment, termi- Johnson collection, Flagstaff, Arizona, and J.E. Wappes nal segment subelliptical. Pronotum campaniform, lateral collection, Chadds Ford, Pennsylvania. margins straight to slightly arcuate, disk evenly convex with slight depressions near posterior corners and on basal Distribution. This species is known only from the Yucatan lobe; surface minutely foveolate, each foveolar seta arising Peninsula. from center; prosternum short, not separating coxal apices; mesosternum truncate apically, postmesocoxal sulcus Biology. The only host association known is with Lysiloma strongly angulate. Scutellum quadrate, deeply emarginate. latisiliqua. Collections were made in December and March. Elytra together (fig. 75) about as long as wide; striae shallow, serially punctate, normal in course except 3 and 4 Discussion. Merobruchus lysilomae, the species most simi- slightly deflected laterally toward paired basal denticles; lar to chetumalae, is compared in the previous description. intervals flat, finely imbricate. Metacoxal face densely, dis- See M. sonorensis for similarities in the 5 genitalia. cretely punctulate proximally, punctures crowded distally; metafemur as in figure 76, pectén with one long and three Placidus Subgroup shorter denticles, metatibia as in figure 76, carinae complete except lateroventral abbreviated apically, coronal den- Characteristics of this subgroup are pronotum and elytra ticles as figured. Abdomen with first segment longer than convex; lateral margins arcuate or sinuate; dorsal pattern remaining segments combined in both sexes; $ with broad, moderately contrasting; ventral valve rounded; internal sac shallow emargination on terminal margin; 2 anal notch nar- with paired, elongate basal spines, median sclerite broad or rowly emarginate, lateral limits obtusely angulate, projecting narrow. Two species. perceptibly. Pygidium as in figures 77 and 78, basal triangle absent to moderately prominent, sometimes with me- Merobruchus placidus (Horn) dian pale line extending to apex. Male genitalia: Median lobe 4 X as long as width of apex; ventral valve variable in (Figs. 75-85) form (figs. 79 and 81-85); armature of internal sac with paired sclerites of variable form near base of sac, Bruchus placidus Horn, 1873:341, 1894:345; Fall, 1910:184; subtriangular median "wishbone" sclerite and paired thorn- Cushman, 1911:498. like sclerites near apex of sac; extreme apex of sac lined Bruchus limpidus Sharp, 1885:456; Pic, 1913:31 (NEW with minute, acute denticles, closure valve circular. Lateral SYNONYMY). lobes (fig. 80) variable in width. Mylabris placidus: Leng, 1920:306. Acanthoscelides placidus: Blackwelder, 1946:760. Body length 1.8-2.8 mm, width 1.1-1.8 mm; pronotal Acanthoscelides limpidus: Blackwelder, 1946:760. length 0.5-0.9 mm, width 0.9-1.3 mm. Merobruchus placidus: Johnson, 1967:264, 1968:1269; Bot- timer, 1968:1024; Johnson, 1969a:55, 1979:123, Type locality. Of Bruchus placidus, Texas; of Bruchus lim- 1981b:251; Johnson and Kingsolver, 1982:418. pidus, Guatemala, Capetillo. Merobruchus limpidus: Johnson and Kingsolver, 1977:154. Type depository. Bruchus placidus (lectotype 8210) in Mu- Color. Integument dark red to dark brown with some sternal seum of Comparative Zoology, Cambridge, Massachusetts, areas piceous; legs and antennae sometimes with piceous designated by Johnson (1968). Bruchus limpidus, lectotype suffusion; pubescence of short, grayish yellow, grayish 5 here designated, marked with small red cross on card white, and brown hairs, latter inserted on maculations of mount of three cotypes, in British Museum (Natural His- elytra; grayish white hairs on ventral areas of body and tory), London. legs; scutellum with white hairs; pattern of vestiture on pronotum variable from nearly uniformly clothed with yellowish Distribution. U.S.A.: Arizona, Texas, Oklahoma, Kansas, hairs to having distinct median and lateral lines cf hairs; Missouri. MEXICO: Baja California, Chihuahua, San Luis pattern on elytra variable from nearly unicolorous to Potosí, Nuevo Leon, Durango, Sinaloa, Jalisco, Michoacán, vaguely maculate at middle of lateral margins and at apex, Veracruz, Mexico, Morolos, Chiapas, Guerrero, Oaxaca, and third interval with prominent, elongated yellowish spot; Campeche, Yucatan. GUATEMALA: Guatemala, Escuintia, pygidium yellowish with variable basal triangular spot. Sacatepéquez. BELIZE: Cayo. COSTA RICA: Puntarenas, San José, Alajuela.

27 Specific records. U.S.A.: Missouri, Jasper Co., Alta, ex and the apex of the spines is straight or curved (figs. 82 Acacia angustissima van hirta. Oklahoma, (no locality). Kan- and 83). The median sclerite is always broad-based but sas, Cowley Co., ex Acacia "hirta." Texas, Dallas, ex Acacia variable in shape. The apical sclerites are always thornlike angustissima var. Iiirta; Bangs; Big Bend N.R, El Paso, ex but differ in the outline of the apex. The lateral lobes offer Acacia texensis (now a var. of angustissima)', Chambers Co.; no consistent characters for specific differentiation. Further Rio Pecos. Arizona, Santa Rita Forest, ex Acacia angustis- collections in Mexico and Central America with longer se- sima var. suffruticosa; Palmerlee; Sierra Ancha Mts.; Pima ries are needed to decipher this problem. Co., Kitt Peak, CDJ 91-72; Huachuca Mts.; Ramsey Can- yon, ex Acacia angustissima. MEXICO: Baja California, San Merobruchus limpidus is here synonymized based on the Jose del Cabo; La Higuera, ex Acacia goldmani. Chihua- male genitalic (fig. 85) and external characters falling within hua, nr. Morita, ex Acacia cuspidata; 35 mi N Chihuahua, the range of variation of placidus. San Luis Potosí, Tamazunchale. Veracruz, 13 mi N Jalapa; Lake Catemaco; Cordoba. Guerrero, Acapuico, ex Acacia Capotillo is an estate near Dueñas, Guatemala. bicolor, 8 mi SE Iguela. Chiapas, 11 mi S Tapilula; El Rin- cón; 7 mi SE Teopisca; 13 km W Ocozocoautia, CDJ Merobruchus solitarius (Sharp) 361-78. Oaxaca, S Juchatango; 116 km SE Oaxaca, CDJ 570-79. Durango, R Buenos Aires, 37 mi El Salto; 11 SW (Figs. 106-110) El Salto; 30 mi W Durango. Sinaloa, Culiacán, CDJ 185-73; Mazatlan, Acacia rosei and A. goldmani; 15 mi W El Pal- Bruchus solitarius Sharp, 1885:456; Pic, 1913:50. mito. Jalisco, Guadalajara. Mexico, Temascaltepec. Morolos, Acanthoscelides solitarius: Blackwelder, 1946:761. Tepoxtlan; Cuernavaca. Yucatan, 23 mi W MotuI, ex Acacia l\Aerobruchus solitarius: Johnson and Kingsolver, 1977:154, angustissima, CDJ 487-78; 20 km S Progreso, ex Acacia 1982:418. angustissima, CDJ 484-78. Campeche, 15 km S Tenabo, CDJ 524-79; 13 km E Merida, CDJ 1854-80. GUATEMALA: Color. Integument dark red to piceous. Head with vertex, Quetzaltenango, Cerro Quemado, ex Acacia filicoides, Sa- eyes, and base of clypeus piceous, frons and antenna red, catepéquez, Dueñas; Capotillo. Escuintia, El Zapote, 20 mi pronotum evenly dark red or medium red with paired pic- SE Guatemala City, in Acacia angustissima, CDJ 1975-80; eous maculae, thoracic sterna piceous, abdomen with 42 km ENE Rio Hondo, Zacapa, CDJ 1891-80. BELIZE: basal sternum piceous along basal margin and sterna 2-5 Cayo, Augustina, ex Acacia angustissima. COSTA RICA: red to black with lateral parts red; elytra varying from all red Puntarenas, 16 mi N Pulperia on P-A Hwy., ex Acacia to red with contrasting black maculae (fig. 106); foreleg and angustissima. San José, ex Acacia angustissima. Alajuela, midleg reddish yellow to red, hindleg dark red with base of San Ramon, ex Acacia angustissima. femur piceous; pygidium red. Vestiture of white or yellowish gray and brown hairs above, all gray hairs ventrally; macu- Biology. The principal host for placidus is Acacia angustis- lations on pronotum and elytra of darker forms accented sima and its varieties suffruticosa, hirta, and texensis, but A. with brown hairs; pronotum with vestiture patchy, more con- bicolor, cuspidata, filicoides, goldmani, and rosei are also trasted in darker specimens; scutellum white; elytra in ten- listed on specimen labels. Acacia filicoides is listed as a eral forms with maculae marked by brown hairs, in speci- synonym of angustissima in Isely (1973), but as a valid spe- mens with developed pattern, third interval with series of cies name in Standley (1922). elongated whitish spots alternated with quadrate brown spots, transverse bars of white or yellowish hairs outlining Discussion. Merobruchus placidus as treated here ranges humeral, lateral, and apical maculae; contrasting dark and from Oklahoma and Missouri to Costa Rica with little varia- light areas less obvious in teneral specimens; pygidium yel- tion in external appearance but with considerable diversity lowish, basal triangle indistinct, paired dark spots of brown in male genitalic details. With the specimen material avail- setae varying in size and shape, 5 usually with only two able to me, I am unable to find any consistency in these spots (fig. 109), 2 often with scattered additional brownish genitalic characters that would indicate a further breakdown areas (fig. 110). of placidus. I am assuming here that the variation in placidus genitalia is random and represents no more than geo- Structure. Body ovate, convex above. Head short, vertex graphic variation. Several forms of the male genitalia are densely punctulate, frons longitudinally rugose, frontal ca- illustrated. rina prominent, transverse sulcus connecting dorsal limits of eyes prominent; pubescence located along frontoclypeal The diversities in the median lobe are as follows: The ven- suture shaggy; postocular lobe narrow, setose; antenna ex- tral valve ranges from a short, broad, arcuate form (fig. 85) tending past humérus, clávate, segments 5-10 eccentric, to a more elongate type (figs. 82 and 84). The basal scle- 11 subtriangular. Pronotum campaniform (fig. 106), lateral rites are with or without auxiliary spines (figs. 79 and 82), margins slightly to moderately convex, disk almost evenly

28 convex but with slight depressions near posterolateral cor- pronotal margins of solitarius contrasted with the more ners; surface irregularly foveolate, each foveola with central smoothly arcuate outline of placidas. These characters are black seta, interspaces microrugose; prosternum barely the basis for segregation of the two species. separating procoxal apices; postmesocoxal sulcus broadly angulate. Scutellunn transverse, bilobed. Elytra together (fig. The host plant, A. angustissima, is used sparingly as an 106) slightly longer than wide, convex; striae slightly sin- ornamental. uate, 1 to 4 slightly deflected laterally near base, 3 and 4 arising from small, bidentate gibbosity on basal margin, Boucheri Subgroup striae deep, narrow; intervals 3, 5, and 7 wider than 2, 4, and 6. Metacoxal face densely punctulate except elongate Characteristics of this subgroup are elytral pattern strongly polished space near anterior border; pectén of metafemur contrasting; ventral valve rounded; internal sac lacking with wide gap between long first denticle and second or basal spines; wishbone sclerite minute with quadrate, trans- third smaller denticles following; muero of metatibia short, parent base; apical spines minute; lateral lobes slender, lateroventral carina obsolete apically. Abdomen with fifth bowed. One species. sternum of 5 broadly emarginate about one-half length of sternum, not flanged laterally, fifth sternum of 2 shallowly Merobruchus boucheri Kingsolver emarginate but with small lateral flanges; pygidium of 5 slightly convex basally, reflexed apically, of 2 oblique, not (Figs. 1-14) reflexed. Male genitalia: Median lobe (fig. 107) nearly 4 x as long as wide, ventral valve broad, arcuate; internal sac Merobruchus sp. 2: Johnson, 1979:123. with rows of short denticles in basal one-half, pair of elon- Merobruchus boucheri Kingsolver, 1980:248; Janzen, gate, fingerlike sclerites in basal one-half, slender, symmet- 1980:948; Johnson and Kingsolver, 1982:418. rical sclerite in middle (corresponding to wishbone-shaped sclerite in other species), and pair of large, thornlike scle- For detailed description, see Kingsolver, 1980:248. rites in apical one-half; apex of sac covered with minute, triangular denticles. Lateral lobes as in figure 108. Color. Integument yellowish to piceous, forelegs and midlegs yellowish red, hindlegs dark red; antenna yellowish Body length 2.2-2.4 mm, width 1.4-1.5 mm; pronotal except segments 8-10 dark brown (fig. 12). Vestiture of yel- length 0.7-0.8 mm, width 0.9-1.1 mm. lowish gray and dark brown slender hairs with scattered coppery hairs on disk in pattern shown in figure 9; prono- Type locality. GUATEMALA: Zapote. tum (fig. 9) with dark median stripe; elytra with dark brown hairs on piceous spots, yellowish gray and coppery on light Holotype depository. In British Museum (Natural History), colored areas; pygidium (fig. 14) with basal triangular spot London. yellowish white, median and lateral dark spots brown, median spot yellowish; forelegs and midlegs yellowish red, Distribution. MEXICO: Veracruz, Michoacán, Puebla, Mex- hindlegs dark red; antenna yellowish except segments 7-9 ico, Oaxaca, Chiapas. dark brown in most specimens.

Specific records since 1980. MEXICO: Veracruz, nr. Rinco- Salient characters. Pronotum elongated with dark median nada, W Clark. Michoacán, Pitzcuaro. Puebla, nr. Acatepec. stripe; antenna with segments 8-10 dark brown; pygidium Mexico, Atlahuaca, O'Brien. Oaxaca, 17 km N Oaxaca, in with basal triangle and median diamond spot prominent, Acacia angustissima, CDJ 282-78; 81 km SE Oaxaca, anal notch of 2 deep, processes long with prominent tufts swept from Acacia angustissima, CDJ 296-78. (figs. 7 and 8); postmesocoxal sulcus strongly angulate (fig. Chiapas, 12-15 km W San Cristóbal, 29-IX-1986. 5); pronotum evenly arcuate in lateral aspect; mesopleural suture meeting middle of anterior margin of metepisternum. Biology. The only host recorded for this species is Acacia Male genitalia as in figures 10 and 11. angustissima. The species has been collected during Feb- ruary, March, July, September, and December. Body length 3.2-3.7 mm, width 1.5-2.2 mm; pronotal width 0.8-1.3 mm, width 1.1-1.5 mm. Discussion. Sclerites of the internal sac of solitarius are similar to those of placidus except that the median sclerite Type locality. COSTA RICA: Guanacaste Prov. in solitarius is always slender (fig. 107). This seemingly triv- ial difference is borne out by the strongly mottled dorsal Holotype depository. In National Museum of Natural History, pattern of solitarius (fig. 106) contrasted with the more Washington, DC. muted pattern of placidus (fig. 75), and the strongly sinuate

29 Literature Cited

Distribution. MEXICO: Baja California Sur, Sonora, Sinaloa, Anonymous. 1940. List of intercepted plant pests, Jalisco, Guerrero, Nayarit, Quintana Roo. HONDURAS: 1939. U.S. Department of Agriculture, Bureau of Plant Comayagua, Olancho, Yoro. COSTA RICA: Guanacaste. Quarantine, Washington, DC. 41 pp. PANAMA: Canal Zone, Veraguas. CUBA: Cayamas. BRA- ZIL: Rio de Janeiro^ Anonymous. 1942. List of intercepted plant pests, 1941. U.S. Department of Agriculture, Bureau of Plant Specific records since 1980. MEXICO: Baja California Sur, Quarantine, Washington, DC. 50 pp. nr. Miraflorae; nr. Burrera; nr. San Jose del Cabo. Sonora, nr. Alamos, in Pithecellobium undulatum, CDJ 175-76 (new Anonymous. 1943. List of intercepted plant pests, host record). Sinaloa, nr. Rosario, in Pithecellobium undula- 1942. U.S. Department of Agriculture, Bureau of Plant tum, CDJ 549-79. Jalisco, Chamela Biological Preserve, in Quarantine, Washington, DC. 41 pp. Pithecellobium mangense. Guerrero, 39 km NW Zihuata- nejo, in Pithecellobium undulatum, CDJ 175-76. Nayarit, nr. Anonymous. 1944. List of intercepted plant pests, Acoponeta, in Pithecellobium mangense, CDJ 482-73. 1943. U.S. Department of Agriculture, Bureau of Plant Quintana Roo, 49 km S Cancun, in Pithecellobium Quarantine, Washington, DC. 35 pp. mangense, CDJ 1783-80. HONDURAS: Comayagua, Si- guatepeque. PANAMA: Veraguas, nr. Santiago. CUBA: Ha- Anonymous. 1945. List of intercepted plant pests, vana, Cayamas. BRAZIL: Rio de Janeiro, Barra de Marica. 1944. U.S. Department of Agriculture, Bureau of Plant Quarantine, Washington, DC. 37 pp. Biology. Host plants for boucheri include Pithecellobium mangense and P undulatum in North America and P tortum Blackwelder, R.E. 1946. Checklist of the Coleóptera of in Brazil. It has been collected in December, February, and Mexico, Central America, the West Indies, and South Amer- March. ica. U.S. National Museum Bulletin 185, part 4, pp. 551- 763. Discussion. A combination of characters separates this species from others in the insolitus group, specifically the Blackwelder, R.E., and R.M. Blackwelder. 1948. Fifth elongated, conical pronotum, dorsal median stripe of the supplement (1939 to 1947, inclusive) to the Leng catalogue pronotum, pattern of pubescence on the elytra and pygid- of Coleóptera, north of Mexico. J.D. Sherman, Jr., Mount ium, and digitate apical processes of the female terminal Vernon, NY. 87 pp. sternum. Associating characters are the median and two lateral thornlike sclerites in the internal sac, bowed lateral Borowiec, L. 1987. The genera of seed beetles (Coleóp- lobes, and basal triangular pad of white pubescence on the tera, Bruchidae). Polskie Pismo Entomologiczne 57:3-207. pygidium. Bottimer, L.J. 1962. Bruchidae. In Arnett, R.H., Jr., The The Brazilian record is a significant extension of geographi- beetles of the United States, part 105, pp. 951-958. cal range for this species known previously no farther south Catholic University of America, Washington, DC. 1112 pp. than Panama. Bottimer, L.J. 1968. Notes on Bruchidae of America north of Mexico with a list of world genera. The Canadian Entomologist 100:1009-1049.

Bottimer, L.J. 1969. Bruchidae associated with Mimosa with the description of a new species. The Canadian Ento- mologist 101:1186-1198.

Bôving, A.G. 1927. On the classification of the Myla- bridae larvae (Coleóptera: Mylabridae). Proceedings of the Entomological Society of Washington 29:133-143.

Bradley, J.C. 1946. Contributions to our knowledge of the Mylabridae, seu Bruchidae (Coleóptera) with especial reference to the fauna of northeastern America. Psyche 53:33-42.

30 Bridwell, J.C. 1946. The genera of beetles of the family Janzen, D.H. 1975. Interactions of seeds and their insect Bruchidae in America north of Mexico. Journal of the predators/parasitoids in a tropical deciduous forest. In Washington Academy of Sciences 36:52-57. Price, RW., ed.. Evolutionary strategies of parasitic insects and mites, pp. 154-186. Plenum Press, New York. Burks, B.D. 1956. The species of Chryseida (Hymenop- 224 pp. tera, Eurytomidae). Bulletin of the Brooklyn Entomological Society 51:109-116. Janzen, D.H. 1977. Intensity of prédation on Pithecello- bium saman (Leguminosae) seeds by Merobruchus colum- Cushman, R.A. 1911. Notes on the host plants and para- binus and Stator limbatus (Bruchidae) in Costa Rican decid- sites of some North American Bruchidae. Journal of Eco- uous forest. Tropical Ecology 18:162-176. nomic Entomology 4:489-510. Janzen, D.H. 1980. Specificity of seed-attacking beetles Fall, H.C. 1910. Miscellaneous notes and descriptions of in a Costa Rican deciduous forest. Journal of Ecology North American Coleóptera. Transactions of the American 68:929-952. Entomological Society 36:89-197. Janzen, D.H. 1982. Cenizero tree (Leguminosae: Pithe- Fall, H.C. 1912. A new Tetropium, two new bruchides, cellobium saman) delayed fruit development in Costa Rican with brief notes on other Coleóptera. Entomological News deciduous forests. American Journal of Botany 23:320-323. 69:1269-1276.

Forister, G.W., and CD. Johnson. 1970. Bionomics of Janzen, D.H. 1983. In Janzen, D.H., ed., Costa Rican Merobruchus julianus (Coleóptera: Bruchidae). Coleop- natural history. University of Chicago Press, Chicago. terists' Bulletin 24:84-87. 816 pp.

Gibson, L.R 1972. Revision of the genus Urosigalphus of Johnson, CD. 1967. Notes on the systematics, host the United States and Canada. Miscellaneous Publica- plants, and bionomics of the bruchid genera Merobruchus tions of the Entomological Society of America 8:83-157. and Stator. Pan-Pacific Entomologist 43:264-271.

Hastings, J.R., R.M. Turner, and D.K. Warren. 1972. An Johnson, CD. 1968. Bruchidae type-specimens depos- atlas of some plant distributions in the Sonoran Desert. ited in United States museums, with lectotype designa- University of Arizona Technical Report on Meteorology and tions. Annals of the Entomological Society of America Climatology of Arid Regions, No. 21, 255 pp. 61:1266-1272.

Horn, G.H. 1873. Revision of the Bruchidae of the Johnson, CD. 1969a. Horn's Bruchidae type-material in United States. Transactions of the American Entomological the Ulke collection. Pan-Pacific Entomologist 45:54-56. Society 4:311-342. Johnson, CD. 1969b. The lectotype of Bruchus julianus. Horn, G.H. 1894. The Coleóptera of Baja California. Annals of the Entomological Society of America 62:676. Proceedings of the California Academy of Sciences 4:302-449. Johnson, CD. 1979. New host records in the Bruchidae (Coleóptera). Coleopterists Bulletin 33:121-124. International Code of Zoological Nomenclature, XV Interna- tional Congress of Zoology. 1961. Stoll, N.R., and Johnson, CD. 1981a. Seed beetle host specificity and others, eds. Richard Clay Co., Ltd., Suffolk, England. the systematics of the Leguminosae. In Polhill, R.M., and 176 pp. PH. Raven, eds.. Advances in legume systematics. Pro- ceedings of the International Legume Conference, Royal International Code of Zoological Nomenclature, XX General Botanic Gardens, Kew, Richmond, England, vol. 2, part 1, Assembly of the International Union of Biological Sciences. pp. 995-1027. 1985. Ride, W.D.L., and others, eds. H. Charlesworth and Co., Ltd., Huddersfield, England. 338 pp. Johnson, CD. 1981b. Interactions between bruchid (Co- leóptera) feeding guilds and behavioral patterns of pods of Isely, D. 1973. Leguminosae of the United States: I. Sub- the Leguminosae. Environmental Entomology 10:249-253. family Mimosoideae. Memoirs of the New York Botanical Garden 25:1-152.

31 Johnson, CD. 1983. Ecosystematics of Acanthoscelides Schaeffer, C 1904. New genera and species of Coleóp- (Coleóptera: Bruchidae) of southern Mexico and Central tera. Journal of the New York Entomological Society America. Miscellaneous Publications of the Entomological 12:197-236. Society of America 56:1-370. Sharp, D. 1885. Bruchidae. In Godman, F.D., and O. Johnson, CD., and J.M. Kingsolver. 1973. A revision of Salvin, eds.. Biología Centrali-Americana, Insecta, Coleóp- the genus Sennius of North and Central America (Coleóp- tera 5:437-504. tera: Bruchidae). U.S. Department of Agriculture Technical Bulletin 1462, 135 pp. Standley, PC. 1922. Trees and shrubs of Mexico, Con- tributions from the National Herbarium 23:171-515. Johnson, CD., and J.M. Kingsolver. 1977. New taxonomic combinations in Bruchidae (Coleóptera). Coleop- Teran, A.L., and S.M. de L'Argentier. 1981. terists Bulletin 31:154. Observaciones sobre Bruchidae (Coleóptera) del noroeste Argentino. IV. Estudios morfológicos y biológicos de Johnson, CD., and J.M. Kingsolver. 1982. Checklist of Ambiycerus hoffmanseggi (Gyll.), Acanthoscelides comptas the Bruchidae (Coleóptera) of Canada, United States, Mex- Kingsolver y Merobruchus bicoloripes (Pie). Acta Zoológica ico, Central America, and the West Indies. Coleopterists Lilloana 36:61-84. Bulletin 35(1981 ):409-422. Wenzel, H. 1912. (Notes at Dec. 11, 1911, meeting, Kingsolver, J.M. 1970. A study of male genitalia in Bru- American Entomological Society.) Entomological News chidae (Coleóptera). Proceedings of the Entomological 23:140. Society of Washington 72:370-386. Wheeler, W.H., J. Hunt, and E.P Reagan. 1950. List of Kingsolver, J.M. 1975. New synonymies and combina- intercepted plant pests, 1948. U.S. Department of Agricul- tions in North America Bruchidae (Coleóptera). ture, Bureau of Entomology and Plant Quarantine, Wash- Proceedings of the Entomological Society of Washington ington, DC. 58 pp. 77:60. White, B.E. 1941. A new species of Bruchus with notes Kingsolver, J.M. 1980. Eighteen new species of Bruchi- on Bruchus major Fall and julianus Horn. Pan-Pacific Ento- dae, principally from Costa Rica, with host records and dis- mologist 17:189-190. tributional notes (Insecta: Coleóptera). Proceedings of the Biological Society of Washington 93:229-283. Whitehead, D.R., and J.M. Kingsolver. 1975a. Megasen- nius, a new genus for Acanthoscelides muricatus (Sharp) Leng, CW. 1920. Catalogue of the Coleóptera of Amer- (Coleóptera: Bruchidae), a seed predator of Cassia grandis ica, north of Mexico. J.D. Sherman, Jr., Mount Vernon, L. (Caesalpiniaceae) in Central America. Proceedings of NY. 470 pp. the Entomological Society of Washington 77:460-465.

Pic, M. 1913. Coleopterorum catalogus. Pars 55, Bru- Whitehead, D.R., and J.M. Kingsolver. 1975b. Biosystem- chidae. Junk, Berlin. 74 pp. atics of the North and Central American species of Gibbobru chus (Coleóptera: Bruchidae: Bruchinae). Transactions of Pic, M. 1927. Coléoptères du globe. Mélanges exotico- the American Entomological Society 101:167-225. entomologiques. Moulins 50:1-36. Zacher, F. 1952. Die Nährpflanzen der Samenkäfer. Pic, M. 1934. Neue Bruchidae (Col.). Arbeiten über Zeitschrift für Angewandte Entomologie 33:460-480. morphologische und taxonomische entomologie aus Berlin- Dahlem 1:116-117.

Pic, M. 1938. Nouveautés diverses, mutations. Mélanges Exotico-Entomologiques 70:1-36.

Polhill, R.M., and PH. Raven, eds. 1981. Advances in legume systematics. Parts 1 and 2. Royal Botanic Gar- dens, Kew, Richmond, Surrey. 1049 pp.

32 Appendix 1. Morphological Terms

A number of morphological terms in this and other reports Lateral carina of the pronotum—a distinct to obsolete ca- on the Bruchidae have been used with little or no explana- rina, or ridge, marking lateral limits of the disk of the prono- tion or illustration. Johnson and Kingsolver (1973) estab- tum. It extends from the posterolateral corner to the anterior lished a nomenclature for the hindleg of the genus Sennius corner of disk. In most of the Acanthoscelidini, the anterior that generally applies to other genera in the tribe Acantho- one-half of the carina is bent ventrad and loses its identity scelidini. I (1970) characterized several forms of male geni- as a carina. The cervical boss (q.v.) is probably the relic of talia in the family, and the nomenclature proposed here has the anterolateral corner. been used in subsequent reports on the Bruchidae. Lateral denticle—the short denticle on the lateral, apical Basal denticle of the elytra—a recumbent, toothlike process margin of the metatibia. at the base of a stria. In most species of Merobruchus, the denticles of striae 3 and 4 are elevated on a basal Lateral margin of the pronotum—configuration as seen in gibbosity. dorsal aspect.

Basal lobe—a broad, antescutellar lobe of the posterior Lateroventral carina of the metatibia—a carina on the lateral margin of the pronotal disk. face of the metatibia lying between the lateral and ventral carinae, sometimes evanescent or missing. Cervical boss—a small, polished elevation on the lower lateral margin of the anterior foramen of the pronotum. It Mesepisternum and mesepimeron—pleural sclerites of the bears two long setae each set deeply into a puncture. This mesothorax. In more generalized bruchid groups, these two may be the marker of the anterolateral corner of the primi- sclerites are nearly equal in size. In the Bruchinae to which tive pronotum. Merobruchus belongs, however, the posterior margin of the mesepisternum has overridden the mesepimeron, narrow- Cervical sulcus—a vertical, lateral sulcus demarcating the ing the latter so that in some genera the mesepimeron is cervix parallel to the margin of the anterior foramen of the reduced to a small, triangular sclerite isolated from the pronotum and dorsad of the cervical boss. mesocoxal cavity.

Coronal denticles—small denticles (usually three) on the Muero—the acute fixed spine at the apex of the ventral ca- dorsoapical margin of the metatibia opposite the muero. rina in more derived groups of bruchids.

Dorsomesal carina of the mesotibia—a carina lying on the Ocular sinus—the deep, usually setose emargination of the mesal face of the tibia near the dorsal margin. eye.

Fifth sternum—the terminal sternum of the abdomen, mor- Pectén—a row of flat denticles set on the mesoventral, dis- phologically the seventh but visibly the fifth. To avoid confu- tal margin of the metafemur. sion, I number the sterna 1 to 5. Postmesocoxal sulcus—a fine sulcus usually running paral- Fovea—a setal insertion enlarged to form a round or oval, lel to the posterior margin of the mesocoxal cavity and ex- flat-bottomed depression, with the seta emerging either tending laterad to the pleurosternal suture. In some species from the center or from the anterior margin of the fovea of Merobruchus, the sulcus is angulate caudad of the coxa. (diminutive foveola). Postocular lobe—a narrow to broad crescentic lobe, usually Frontal carina—a vertical ridge extending from a line con- setose, attached to the posterior margin of the eye. necting the upper limits of the eyes (usually the transverse sulcus, q.v.) to the frontoclypeal suture. Pygidium—terminal (seventh) tergum usually fully exposed caudad of the elytral apices in the Bruchidae. Incrassate—thickened. In bruchid literature, the term is used to describe a metafemur that is expanded Subbasal gibbosities—small elevations near the posterior dorsoventrally. margin of the pronotal disk; present in several genera of Neotropical bruchids, especially Caryedes, Gibbobruchus, Lateral carina of the metatibia—a carina extending the full and Ctenocolum. In Merobruchus, the gibbosities are much length of the metatibia in the middle of the lateral face. reduced or absent. Only in Merobruchus major (Fall) are they apparent.

33 Appendix 2. Synonymical List of Merobruchus Species

Transverse sulcus—a vague to well-defined transverse de- 1. boucheri Kingsolver pression on the head separating the vertex from the frons. 2. chetumalae, new species 3. columbinus (Sharp) Ventral carina of the metatibia—the hindleg is described as 4. cristoensis, new species though it were open and extended. The morphologically 5. hastatus Kingsolver ventral margin with the ventral carina and apical muero ap- 6. insolitus (Sharp) pears to be dorsal when the leg is closed. 7. ¡ulianus (Horn) ochreolineatus Fall 8. knulli (VJhWe) 9. lineaticollis (Sharp) 10. lysilomae, new species 11. major (Fall) flexicaulis Schaeffer 12. paquetae Kingsolver 13. placidas (Horn) limpidus Sharp 14. politus, new species 15. porphyreus, new species 16. sanfarosae Kingsolver 17. so//far/iys (Sharp) 18. sonorensis Kingsolver 19. terani Kingsolver 20. triacanthus, new species 21. \/ac/7/afor (Sharp) 22. xanthopygus, new species

34 Appendix 3. Merobruchus Species and Associated Host Plants

Merobruchus Merobruchus species Host plants species Host plants boucheri Pithecellobium mangense, tortum, sonorensis Albizia adinocephala, caribaea, lebbek, undulatum occidentalis, ortegae, sinaloensis, tomentosa; Lysiloma divaricata, chetumalae Lysiloma latisiliqua, Lysiloma sp. seemani; Pithecellobium sonorae columbinus Pithecellobium saman terani Acacia angustissima, berlandieri, gaumeri, picachensis, tenuifolia cristoensis No host known triacanthus Acacia acatlensis, coulteri, riparioides; hastatus Lysiloma desmostachys Leucaena guatemalensis; Lysiloma divaricata insolitus Acacia occidentalis, tenuifolia; Albizia adinocephala, lebbek, occidentalis, vacillator Lysiloma divaricata sinaloensis; Lysiloma acapulcensis, candida, divaricata, kellermani, xanthopygus Lysiloma acapulcensis microphylla van thornberi, seemani, Lysiloma sp.; Pithecellobium mangense, pallens, sonorae, undulatum

¡ulianus Acacia berlandieri, coulteri, greggi, roemeriana, wrightii knulli Lysiloma acapulcensis, microphylla van thornberi, watsoni lineaticollis No host known lysilomae Acacia richii, simplicifolia; Albizia lebbek, polyphylla; Lysiloma latisiliqua, polyphylla, sabicu, Lysiloma sp. major Pithecellobium flexicaule paquetae Albizia adinocephala, caribaea, lebbek, longipedata, Albizia sp.; Lysiloma sp.; Pseudosamanea guachepele, Pseudosamanea sp. placidus Acacia angustissima, bicolor, cuspidata, filicoides, goldmani, rosei politus Pithecellobium leptophyllum porphyreus Lysiloma acapulcensis, kellermani

santarosae Acacia coulteri; Albizia sp.; Lysiloma desmostachys

solitarius Acacia angustissima

35 Appendix 4. Host Plants Attacked by Merobruchus Species

Merobruchus Merobruchus Host plant species Host plant species

Acacia longipedata Britton and Rose paqaetae

acatlensis Bentham triacantlius occidentalis T.S. Brandegee insólitas, sonorensis

angustissima (P. Miller) Kuntze placidas, solitarius, ortegae Britton and Rose sonorensis terani polyphylla Fournier lysilomae berlandieri Bentham julianus, terani sinaloensis Britton and Rose insólitas, sonorensis bicolor Britton and Rose placidas tomentosa Standley sonorensis coulteri Bentham ¡allanas, santarosae, triacanthas sp. paqaetae, santarosae

cuspidata Schlectendal placidas Leacaena gaatemalensis Britton triacanthas and Rose filicoides (Cavanille) Trelease placidas Lysiloma gaumeri Blake terani acapalcensis (Kunth) Bentham insólitas, knalli, goldmani (Britton and Rose) placidas porphyreas, xanthopygas Wiggins candida T.S. Brandegee insólitas greggi A. Gray ¡allanas desmostachys Bentham hastatas, santarosae occidentalis Rose insólitas divaricata (Jacquin) Macbride insólitas, sonorensis, picachensis T.S. Brandegee terani triacanthas, vacillator

richii A. Gray lysilomae kellermani Britton and Rose insólitas, porphyreas

riparioides (Britton and Rose) triacanthas latisiliqaa (Linn.) Bentham chetamalae, lysilomae Standley microphylla var. thornberi insólitas, knalli roemeriana Scheele ¡allanas Britton and Rose

rosei Standley placidas polyphylla Bentham lysilomae

simplicifolia (Linn.) Druce lysilomae sablea Bentham lysilomae

tenuifolia F. Muell. insólitas, terani seemani Britton and Rose insólitas, sonorensis

wrightii Bentham ¡allanas wafso/7/J.N. Rose knalli

Albizia sp. chetamalae, insólitas, lysilomae, paqaetae adinocephala Britton and Rose insólitas, paqaetae, sonorensis Pithecellobiam

caribaea Britton and Rose paqaetae, sonorensis flexicaale (Bentham) Coulter major

lebbek (Linn.) Bentham insólitas, lysilomae, leptophyllam (Cavanille) politas paqaetae, sonorensis Daveau

36 Merobruchus Host plant species

mangense (Jaquin) Macbride boucheri, insolitus

pallens (Bentham) Standley insolitus

saman (Jacquin) Bentham columbinus

sonorae S. Watson insolitus, sonorensis

tortum Martius boucheri

undulatum (Britton and Rose) boucheri, insolitus Gentry

Pseudosamanea

guachepele Harms paquetae

sp. paquetae

37 ¡' ''''''mm'M'-"''''''''''''

Figures 1-8, Merobruchus boucheri Kingsolver: 1, Basal margin, right elytron, denticles at bases of striae 3 and 4; 2, lateral aspect of metasternum and left fiindleg; 3, fiead, cepfialic aspect; 4, antenna; 5, left mesocoxal cavity with postmesocoxal sulcus (arrows); 6, pronotal disk showing foveolae and pubescence; 7, apex of female pygidium showing apices of fifth sternal processes; 8, ventral aspect of female fifth sternum with digitate processes and anal notch.

38 39 11

Figures 9-14, Merobruchus boucheri Kingsolver: 9, Dorsal habitus; 10, male genitalia, median lobe; 11, same, lateral lobes; 12, antenna; 13, hindleg; 14, pygidium.

40 Figures 15-18, Merobruchus chetumalae, new species: 15, Dorsal habitus; 16, male genitalia, median lobe; 17, pygidium; 18, same, female.

41 Figures 19-24, Merobruchus columbinus (Sharp): 19, Dorsal habitus; 20, male genitalia, median lobe; 21, same, lateral lobes; 22, head 23, hindleg; 24, pygidium.

42 26 27

Figures 25-29, Merobruchus cristoensis, new species: 25, Dorsal habitus; 26, male genitalia, median lobe; 27, same, lateral lobes; 28, hindleg; 29, pygidium.

43 32

Figures 30-34, Merobruchus hastatus Kingsolver: 30, Dorsal habitus; 31, male genitalia, median lobe; 32, same, lateral lobes; 33, hindleg; 34, pygidium.

44 Figures 35-40, Merobruchus insolitus (Sharp): 35, Dorsal habitus; 36, male geriitaiia median lobe; 37, same, lateral lobes; 38, hindleg; 39, pygidium, male; 40, same, female.

45 Figures 41-46, Merobruchus julianus (Horn): 41, Dorsal habitus; 42, male genitalia, median lobe; 43, same, lateral lobes; 44, head; 45, hindleg; 46, pygidium.

46 Figures 47-52, Merobruchus knulli (White): 47, Dorsal habitus; 48, male genitalia, median lobe; 49, same, lateral lobes; 50, hindleg; 51, pygidium, male; 52, same, female.

47 Figures 53-57, Merobruchus lineaticollis (Sharp): 53, Dorsal habitus; 54, male genitalia, median lobe; 55, same, lateral lobes; 56, hindleg; 57, pygidium, female.

48 8 II s ^ Il 11 >

Figures 58-62, Merobruchus lysilomae, new species: 58, Dorsal liabitus; 59, maie genitalia, median lobe; 60, same, lateral lobes; 61, hindleg; 62, pygidium.

49 Figures 63-69, Merobruchus major (Fall): 63, Dorsal habitus; 64, male genitalia, median lobe 65, same, lateral lobes; 66, head; 67, hindleg; 68, pygidium, male; 69, same, female.

50 Figures 70-74, Merobruchus paquetae Kingsolver: 70, Dorsal habitus; 71, male genitalia, median lobe; 72, same, lateral lobes; 73, hindleg; 74, pygidium.

51 Figures 75-78, Merobruchus placidas (Horn): 75, Dorsal habitus; 76, hindleg; 77, pygidium, male; 78, same, female.

52 85

Figures 79-85, Merobruchus placidus (Horn), male genitalia: 79, Median lobe, Brownsville, Texas; 80, lateral lobes, Brownsville, Texas; 81, median lobe, Temascaltepec, Mex., Mexico; 82, median lobe, Acapulco, Guer, Mexico; 83, median lobe, Cuernavaca, Morelos, Mexico; 84, median lobe, Yucatan Peninsula, Mexico; 85, median lobe, Bruchus limpidus paratype, Sacatetepequez, Guatemala.

53 Figures 86-92, Merobruchus politus, new species: 86, Dorsal habitus; 87, male genitalia, median lobe; 88, same, lateral lobes; 89, lateral outline of body with position of polished spot; 90 hindleg; 91, pygidium, male; 92, same, female.

54 Figures 93-97, Merobruchus porphyreus, new species: 93, Dorsal habitus; 94, male genitalia, median lobe; 95, same, lateral lobes; 96, hindleg; 97, pygidium.

55 Figures 98-105, Merobruchus santarosae Kingsolver: 98, Dorsal habitus; 99, elytron, dark phase; 100, elytron, pale phase; 101, male genitalia, median lobe; 102, same, lateral lobes; 103, hindleg; 104, pygidium, male; 105, same, female.

56 110

Figures 106-110, Merobruchus solitarius (Sharp): 106, Dorsal habitus; 107, male genitalia, median lobe; 108, same, lateral lobes; 109, pygidium, male; 110, same, female.

57 115

Figures 111-115, Merobruchus sonorensis Kingsolver: 111, Dorsal habitus; 112, male genitalia, median lobe; 113, same, lateral lobes; 114, hindleg; 115, pygidium.

58 120

Figures 116-120, Membruchus terani Kingsolver: 116, Dorsal habitus; 117, male genitalia, median lobe; 118, same, lateral lobes; 119, hindleg; 120, pygidium.

59 ^HA,.-'' 123

Figures 121-125, Merobruchus triacanthus, new species: 121, Dorsal habitus; 122, male genitalia, median lobe; 123, same, lateral lobes; 124, hindleg; 125, pygidium.

60 Figures 126-130, Merobruchus vacillator (Sharp): 126, Dorsal habitus; 127, male genitalia, median lobe; 128, same, lateral lobes; 129, hindleg; 130, pygldium.

61 ''Ay 133

Figures 131-136, Merobruchus xanthopygus, new species: 131, Dorsal habitus; 132, male genitalia, median lobe; 133, same, lateral lobes; 134, hindleg; 135, pygidium; 136, anal notch.

62 Figures 137-138, Merobruchus species from South America: 137, Male genitalia, median lobe, M. bicoloripes; 138, same, M. pickeli.

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