2 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 This work is dedicated to

Dr. Byron Alexander

with appreciation for his inspiring talent and dedication PUBLICATIONS COMMITTEE to teaching, research, and scientific illustration. of THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1998

Thomas J. Henry Wayne N. Mathis Gary L. Miller, Book Review Editor David R. Smith, Editor

Printed by Allen Press, Inc. Lawrence, Kansas 66044

Date issued: 5 March 1998 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 LINGAFELTER: GENERA OF

TABLE OF CONTENTS Micranejus ...... , ...... , Micranoplium ...... , ...... , Micropsy rassa ...... , ...... , Abstract ...... Miltesthus ...... , ...... , Introduction ...... ,...... Minipsyrassa ...... ,...... Taxonomic History ...... -...... , ...... Miopteryx ...... , ...... , . . , ...... Disuibution and Diversity ...... ,...... MorphaneJlus ...... Special Problems Associated with Monotypic Taxa ...... , . . . ..,.. Neaneflus ...... , ...... Biology and Natural History ...... Neomallocera ...... ,...... , ...... Materials and Methods ...... Neoperiboeum ...... , .. . .. Cladistic methods ...... Nephaliodes ...... - ...... , . . . . . Choice of taxa ...... Nesanoplium ...... , ...... Unavailable and fossil taxa ...... Nesiosphaerion ...... Specimen preparation ...... Nesodes ...... :...... , ...... Character descriptions and states ...... ,...... Nyssicostylus ...... Characters not used in analyses or key ...... ...... ,...... Characters used in key only ...... ...... Characters used in analyses ...... Pantonyssus ...... , ...... Implied Weighting Method of Parsimony ...... , . . Paramallocera ...... ,...... Phylogenetic analyses ...... Parasphaerion ...... , ...... , . .. , ...... Results of Phylogenetic Analyses ...... ,...... Parastizocera ...... Discussion of Implied Weighting Consensus Tree ...... Parelaphidion ...... Discussion of Subset of Obtained Trees: The Five Shortest, Fittest Trees of the Implied Weight- ...... ing Analysis ...... , ...... Piezophidion ...... , ...... Problematic Taxa and Discussion of Outgroup ...... Pilisphaerion ...... Summary of Classification Changes ...... ,...... Poecihmllus ...... , ...... Generic Treatment ...... , . . Protomallocera ...... , ...... Tribe Elaphidiini ...... , ...... ...... Adiposphaerion ...... , ...... Pseudomallocera ...... ...... , ...... Pse~doperiboe~...... ,...... Amethysphaerion ...... ,...... , ...... ,. . PSYrasm ...... ,...... ...... Psyrassafona ...... ...... ,...... Rbmb0idedere.r ...... ,...... ...... , , ...... Romulus ...... , ...... ,. . . Anopliomorpha ...... Sphaerioeme ...... , ...... Anoplocurius ...... S~hacrion...... , ...... Apoclausirion ...... Sp~~aerjonillun~...... ,...... , ...... Aposphaerion ...... , ...... Stenelaphus ...... Appula ...... ,...... - ...... Stenos~henus...... , , ...... ,...... Astromula ...... , ...... ,...... , ...... ...... Terpnissa ...... Tricho~horoides...... Tropimerus ...... Cenrrocerum .. . ,...... Key to Genera of Elaphidiini ...... ,. . Clausirion ...... ,...... Conclusion ...... , ...... , ...... ,. . Conosphaeron ...... ,...... Ac~owledgments...... ,. . .. . ,. . ...... Literature Cited ...... ,...... , ...... ...... :,...... Appendix 1: Taxa Used in Phylogenetic Analyses ...... Elaphidionopsis ...... , , ...... Appendix 2: Provisional Ingroup Genera Not Represented in Phylogenetic Analyses...... ...... Index ...... , ...... , . . . . , , . , Etymosphaerion ...... ...... Eustromula ...... Gymnospyra ...... ,...... Hemilissopsis ...... , ...... , . . ...... ,...... ,. . . . . ,...... Linsleyonides ...... , . . ...... Meganejus ...... , ...... , ...... , ...... Megapsyrassa ...... ...... Metironeus ...... 6 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 LINGAFELTER. GENERA OF ELAPHIDIINI

Abstract.-A generic-leve1 phylogenetic analysis of the tribe Elaphidiini Thomson 1864 elaphos, meaning deer, and latinized with (Coleoptera: Cerambycidae) is presented. Cladistic methods using morphological features The Elaphidiini is among the most ge- the nominative singular termination, -on. and implied weighting parsimony were employed. The monophyly of the tribe is weakly nerically diverse tribes in Cerambycidae. Thus the stem is Elaphidi and the tribal suf- supported by presence of antennal carinae, tibial carinae, and an abmptly rounded anterior Seventy-seven genera and over 520 fix, -ini must be added, forrning Elaphidiini margin of the mesonotum. Based on the results of the implied weights phylogenetic are currently known for this group of wood- (following Zlion = Iliidae exarnple from analysis and classificatory decisions, the following taxonomic changes are made: Four boring . Most species are nocturnal, ICZN, page 21 1). Until this study only two genera are transferred into other tribes and six genera are synonymized leaving 77 genera and in some regions of México at certain of these original genera (Orion Guérin- in Elaphidiini. Phoracanthini Lacordaire 1869 is removed from Elaphidiini and resurrected times of the year, elaphidiines are the most Méneville and Elaphidion Audinet-Ser- as a tribe with Newman 1840b as the type genus. Serville abundant cerarnbycids attracted to lights. ville) were still in the tribe. The others are 1834 and Allotraeus Bates 1887 are not elaphidiines and are tentatively retumed to Phor- As currently defined, elaphidiines occur currently distributed arnong other tribes in- acanthini. Championa Bates 1880, and ORon Guérin-Méneville 1844 are removed from from Canada to South America, with the cluding Hesperophanini, Callidiopini, and Elaphidiini and placed as incertae sedis in . The following six new generic greatest species diversity in the warmer lat- Methiini. Thomson further characterized synonymies are proposed: Eutrichophoroides Linsley 1961b and Neotrichophoroides itudes. Characters to define and diagnose the tribe Eburitae, originally proposed by Linsley 1961b both = Linsley 1935a. Nesostizocera Linsley 1961b = genera have rarely been found or applied in Blanchard (1845), as including many other Stizocera Audinet-Serville 1834. Hemistizocera Linsley 1961b = Pascoe 1866. this group, and the morphological similarity currently recognized elaphidiine taxa in- Peranoplium Linsley 1957b = Anelaphus Linsley 1936. Axestinus LeConte (1873) = among genera has precipitated its confused cluding Atylostagma White, Centrocerum Aneflus LeConte (1873). The following 29 new combinations are proposed: Aneflus ob- taxonomic history. With this study, I pres- Chevrolat, Ambonus Gistel, Au- scurus (LeConte 1873), Anelaphus eximium (Bates 1885), Anelaphus hoferi (Knull ent an overview of what is known on the dinet-Serville, Periboeum Thomson, Appu- 1934b), Anelaphus inornatum (Chemsak and Linsley 1979), Anelaphus maculatum , diversity, and biology of Ela- la Thomson, Stizocera Audinet-Serville, (Chemsak and Noguera 1993), Anelaphus piceum (Chemsak 1962), Anelaphus simile phidiini. I developed a list of explicitly de- Mallocera Audinet-Serville, and Eurysthea (Schaeffer 1908), Anelaphus subdepressum (Schaeffer 1904), Anelaphus tuckeri (Casey fined characters and states for a11 elaphidi- Thomson. Thomson (1864) defined the Ela- 1924), Anelaphus undulatum (Bates 1880), Psyrassa cribricollis (Bates 1885), Stizocera ine genera and potentially closely related phidiini as having body convex, eyes atiaia (Martins and Napp 1983), Stizocera caymanensis (Fisher 1941), Stizocera dozieri taxa and coded these characters in an ex- coarsely faceted, femora slightly clavate, (Fisher 1947), Stizocera floridana (Linsley 1949), Stizocera insulana (Gahan 1 895), Sti- tensive matrix. I provided the first cladistic and elytral apices spinose. These characters zocera jassuara (Martins and Napp 1983), Stizocera phtisica (Gounelle 1909), Stizocera analyses of genera in this tribe in an attempt separated Elaphidionitae from his Eburitae poeyi (Guérin-Méneville 1838), Stizocera punctiventris (Cazier and Lacey 1952), Stizo- to discover the evolutionary history and de- which had body subdepressed, elytra with- cera submetallicus (Chemsak and Linsley l968), Stizocera suturalis (Martins and Napp cipher their relationships. For each genus, I out apical spines, and femora clavate. La- 1992), Stizocera vanzwaluwenburgi (Fisher 1932), Stizocera wagneri (Gounelle 19 13), provide a diagnosis, description (in the ma- cordaire (1869) proposed the groups Hes- Trichophoroides albisparsus (Bates 1872), Trichophoroides jansoni (Bates 1885), Tricho- trix), comments on distribution and diver- pérophanides (including taxa which are cur- phoroides aurivillii (Linsley 1961), Trichophoroides decipiens (Bates 1880), Trichopho- sity, and a discussion of relationships and rently in Elaphidiini and Hesperophanini, roides pilicornis (Fuchs 1961). Diagnoses of a11 genera are presented with notes on dis- similarities to other taxa. A key to a11 the characterized by the non-globose anterior tribution, diversity, and relationships. A key to genera of Elaphidiini is presented. genera of Elaphidiini (and similar taxa with coxae, externally open intermediate coxal mesally spined antennae) is provided for cavities, and generally unspined antennae), their identification. Éburiides (mainly consisting of taxa with ebumeous elytral calli but identified pri- marily on the basis of externally-closed in- termediate coxal cavities, globose anterior Thomson (1864) proposed the "Divi- coxae, and unspined antennae), Phoracan- sion" Elaphidionitae to include ten genera. thides (including taxa currently in Elaphi- This farnily group name was based on the diini and Phoracanthini, characterized by genus Elaphidion Audinet-Serville. Elaphi- spined but non-carinate antennae, interme- dionini has been used since 1930, but Ivie diate coxal cavities open externally, procox- (1985) indicated Elaphidiini should be the al cavities not angulate externally), and appropnate name of the tribe. The basis of Sphérionides (including nearly a11 of Thom- Ivie's suggestion rests on the actual stem of son's Eburitae, characterized by spines and Elaphidion. The ICZN (Article 29a) states carinae on antennae, anterior coxal cavities that the appropriate farnily group suffix is angulate externally, and intermediate coxal added to the stem of the narne of the type cavities open externally). genus. Elaphidion was based on the Greek The variability of the characters above 8 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 LINGAFELTER: GENERA OF ELAPHIDIINI

(antennal carinae, openlclosed intermediate characters or combinations of characters in bark of dead branches. The larvae either cies. Many taxa in the southem United coxal cavities, procoxal cavities angulatel which endlessly split natural groups of spe- feed for some time under the bark, or im- States and México are associated with le- not angulate, and antennal spines) was ac- cies. These are the same attributes expected mediately enter the wood, feeding and de- gurninous plants of the genera Prosopis and knowledged by Linsley (1936). He noted of a phylogeny. The carefully defined genus veloping within the heartwood. Larval de- Acacia (Linsley 1963, and references cited that the distinction of Sphaerionini from taxon (or clade) should allow one to make velopment most often takes one to three therein). Much additional information is be- Phoracanthini (including Elaphidiini) was predictions or generalizations about other years, correlated with the size of the . ing discoverd by G. Tavakilian in French not satisfactor y. In typical Phoracanthini species included (Clayton 1972, Gauld and A pupal cell is created in the region be- Guyana. (including Elaphidiini), he indicated that Mound 1982). No group of taxa should be tween the bark and sapwood at the end of the anterior coxal cavities may be either removed from a clade if it will render either larva1 development. Pupation occurs in ei- MATERIALS AND METHODS closed or open in closely related species of paraphyletic. If this happens, one loses the ther late surnmer to early fall, or early Cladistic methods.-The goals of this the same genus, and therefore was not a potential predictive attributes (for host plant spring. Adults do not emerge until spring study are to: recharacterize the tribe Ela- useful character. He felt that the most reli- associations, biological compound pros- or surnmer, regardless of when pupation oc- phidiini and the included genera based on able characters were the non-carinate anten- pecting, adaptative characters, rates of spe- curs. exarnination of morphological characters of nae and tibiae in Phoracanthini, but recog- ciation, etc.) of the phylogeny and the clas- It is assumed that adults of most elaphi- the adults; perform a phylogenetic analysis nized that even those characters were not sification becomes meaningless. However, diines (like many longhomed beetles) feed in order to propose a hypothesis of generic consistent in species of Elaphidion. Linsley there will always be subjectivity in deter- very little or not at all, but few references relationships and reveal needed changes in (1963) included the sphaerionine and phor- mining the amount of difference necessary to feeding behavior exist. Specimens in classification; and develop a key to genera. acanthine genera in his new concept of Ela- to erect a new taxon in cases where this will some genera (Anelaphus, Elaphidion, Par- I have used the principles and tenets of cla- phidiini, rendering Phoracanthini and not render one paraphyletic with respect to elaphidion Skiles, Enaphalodes Haldeman) distics to guide the methods described here Sphaerionini as junior synonyms. the other. are attracted in great numbers to brown sug- (Forey, et al. 1994, Hennig 1966, Wiley The Elaphidiini have a history of generic ar bait solutions (Lingafelter and Homer 1981, and Wiley, et al. 1991). DISTRIBUTION AND DIVERSITY concepts based on presumed unique com- 1993), indicating their natural attraction to Choice of taxa.-I have used the type The Elaphidiini as recognized herein, binations of widespread character states and sap flows or other natural high-sugar species for each provisional elaphidiine ge- have a widespread distribution, but with such genera do not convey the information sources of nutrition. Adults of at least two nus in the analyses when possible. If the distinct concentrations of diversity. The or allow the predictions or generalizations diurna1 genera including Tropimerus Gies- type species was not available, I hypothe- "elaphidiine" group is most concentrated that would be preferred. It is my intention bert (Giesbert 1987), and sized that the chosen taxon was a compa- in México and the West Indies, extending to lessen this problem by redefining genera Haldeman (Giesbert and Chemsak 1989) rable representative for the genus based on into South America and Canada. The on the basis of detailed morphological ex- are comrnonly encountered on flowering knowledge of the type species and at least "sphaerionine" group is most concentrated amination and results of the phylogenetic trees. AneJlomorpha tenuis (LeConte) adults superficial similarity to it by the chosen tax- in South America, extending into Central analyses. have been reported feeding in large num- on. I also included additional representa- America and México. The number of de- bers on Karwinskia blossoms (Tumbow and tives of large, diverse genera to better allow scribed Elaphidiini in the Westem Hemi- Wappes 1981). Adults of Anelaphus albo- tests for their monophyly. sphere exceeds 500 species (Monné 1993). Very little is known of the biology of fasciatus Linell have been reported feeding Because the Elaphidiini have had no phy- In México alone the number is over 200 Elaphidiini. As nearly all of them are noc- on new growth of Opuntia (Raske 1972). logenetic investigations previously, it is im- species (Chemsak 1991, Monné 1993, tumal as adults, over 95% of the specimens Twig-girdling-cutting off the flow of portant to avoid a restricted outgroup Monné and Giesbert 1993). This tribe has are taken at lights and thus are collected nutrients or chemicais to a portion of the choice. In fact, a11 potentiaily closely relat- a very unusual ratio of species to genera. without any host or association records. plant, thereby killing part of it-is a behav- ed taxa should be included in the analyses There are 86 genera of which 32 are mono- Their abundance at lights can be remark- ior most comrnonly associated with the dis- to allow for the most rigorous test of mono- typic (37%), 15 are bitypic (17%), and only able. In westem México, 13 nights of col- tantly related Onciderini. In this group the phyly for the ingroup (Nixon and Carpenter 39 (46%) have more than two species lecting at one illuminated roadside sign pro- girdling is performed by the adult female 1993). I have included a broad representa- (many have only three species). duced 1700 specimens of longhom beetles. prior to oviposition. Girdling has also been tion of additional, potentially closely relat- Elaphidiines represented 63% of a11 speci- noted to occur in at least three genera of ed tribes within the subfarnily Cerambyci- SPECIAL PROBLEMS ASSOCIATED WITH mens and 35% of a11 species (Chemsak, et Elaphidiini including Psyrassa Pascoe nae. MONOTYPIC TAXA ai. 1988)! (Champlain, et al. 1925); AneJlomorpha Unavailable and fossil taxa.-Due to the The existence of small and monotypic The following is a generalized life cycle Casey (Craighead 1923); and Anelaphus number of rare, monotypic genera in this genera is an important problem in this of elaphidiines (for temperate species, sum- Linsley (Craighead 1950), although in these tribe, some genera could not be represented study. Genenc taxa should be erected to marized and generalized from Solomon groups the girdling is intemal and done by in the analyses either because the specimens convey information on characteristics that 1995). Adults emerge in spring or surnmer, the larvae. were not available for exarnination, or be- bind sets of species together, not unique mate, and females lay the eggs in notches Larval hosts are not known for most spe- cause the few rare specimens could not be . . .. . -. --. . . - - --.- ..-

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 1 o LINGAFELTER: GENERA OF ELAPHIDIINI 11 dissected for exarnination of the full range sclerites. Male genitalia, labia, maxillae, Table 1. Institutions and private collections that provided material for this study of phylogenetically important characters. and mesonota were then transferred to the Ornission of taxa is probably very common glycerin vial. The remaining body parts Institution Visited? Loanlgift in phylogenetic analyses since most extinct were partially cleared in 10% hydrogen per- received? taxa have an undiscovered or absent fossil oxide solution for 1-10 minutes depending Academy of Natural Sciences, Philadelphia, Pennsylvania (ANSP, Donald Azuma) record, or insufficient information can be on structure and amount of melanization. American Museum of Natural History, New York, New York gleaned from their fossil record. The Ela- When cleared, remaining structures were (AMNH, Lee Herman) phidiini have a poor fossil record with transferred to the glycerin vials. Australian National Collection-CSIRO, Canberra, Australia (ANIC, Tom Weir) North American examples known only Types of characters used in phylogenetic Bernice P. Bishop Museum, Honolulu, Hawaii (BPBM, AI Samuelson) from Florissant, Colorado (Linsley 1942). I analysis and key.-The majority of char- California Academy of Sciences, San Francisco, California examined in the MCZC two of the three acters used were exoskeletal features or cu- (CASC, David Kavanaugh, Roberta Brett) ticular processes visible through dissection Canadian Museum of Nature, Ottawa, Ontario, Canada described fossil species of Elaphidiini, Ane- (CMNC, Bob Anderson) laphus extinctus (Wickham) and Steno- microscopy with fiber optic illumination Canadian National Collection, Ottawa, Ontario, Canada sphenus pristinus Wickham (Fig. 11). I and magnification less than 20X. Addition- (CNCI, Ed Becker, Jean McNamara) have compared my observations with those a1 characters could only be revealed effec- Carnegie Museum, Pittsburgh, Pennsylvania (CMNH, John Rawlins) Daniel Heffern private coilection, Houston, Texas (DHPC) of Wickham (1914) and have concluded tively through the use of Nomarski inter- David Marqua private collection, Davis Mountains, Texas (DMPC) that these fossils convey insufficient infor- ference compound microscopy. Presump- Edward Riley private collection, College Station, Texas (ERPC) mation for inclusion in the phylogenetic tive homologous characters were compared Edmund Giesbert private collection, Beverly Hills, California (EGPC) and discretely variable states were identi- Essig Museum of Entornology, Berkeley, California analyses. These taxa, along with Elaphidion (CISC, John Chemsak, Cheryl Barr) fracticorne Wickham, should be designated fied and coded in the data maüix. Wilkin- Estación de1 Biología Chamela, Jalisco, Mexico (EBCC, Felipe Noguera) incertae sedis at the leve1 of genus. son (1992) presents an argument for and Field Museum, Chicago, Illinois (FMNH, AI Newton, Philip Parrillo) Sumrnary of terminal taxa.-I included against ordering of characters. I have treat- Frank Hovore private collection, Los Angeles, California (FHPC) ed a11 characters as unordered according to Henry Stockwell private collection, Panamá City, Panamá (HSPC) 89 provisional ingroup species and 10 pro- Instituto Nacional de Biodiversidad, Santo Domingo, Herédia, Costa Rica visional outgroup species as terminal taxa the principle of indifference (Keynes 1921) (INBC, Angel Solís) (Appendix 1) in the phylogenetic analyses. summarized in Wilkinson (1992). This as- Jim Wappes private collection, Bulverde, Texas (JWPC) Specimens used in the analyses were ob- sumption proposes that transfonnation be- Marlin Rice private collection, Ames, Iowa (MRPC) Montana State University Entomology Collection, Bozeman, Montana tained from the collections listed in Table tween states is equiprobable, and there is no (MTEC, Mike Ivie) 1. Listed in Appendix 2 are provisional ela- information available to suggest otherwise. Museu de Zoologia da Universidad de São Paulo, São Paulo, Brazil phidiine genera which were excluded from Some characters were found to be contin- (MZSP, Ubirajara Martins) uously variable. Thiele (1993) presents jus- Museu Nacional Quinta da Boa Vista, Rio de Janeiro, Brazil the analyses because no specimens were (QBUM, Miguel Monné) available for dissection. tification and a procedure for using contin- Muséum National d'Histoire Naturelle, Paris, France Specimen preparation.-For the phylo- uou~morphometric data in phylogenetic (MNHN, Jean Meniér) genetic analyses and key, I used characters analyses. However, due to the limitations of Museum of Comparative Zoology, Cambridge, Massachussets (MCZC, David Furth, Cleone Graham) of the adult morphology. Since the larvae software to deal with a11 potential states and National Institute Agro-Env. Sciences, Kannondai, Tsukuba, Ibaraki Pref., are unknown for most elaphidiine species, the artificial state construction formula of Japan (ITLJ, T. Matsumura, Akiko Saito) this potential data set could not be used. Thiele (1993: 284), I chose to exclude con- Natural History Museum, London, England (BMNH, S. Shute) Specimens were prepared by relaxing them tinuously variable characters from the phy- Smithsonian Tropical Research Institute, Panamá City, Panamá (STRI, Annette Aiello and Donald Windsor) in hot water for 5-10 minutes, depending logenetic analyses. These characters were Snow Entomological Museum, Lawrence, Kansas (SEMC, J. Steve Ashe) on size. Then, the head, prothorax, abdo- used in the key and coded multiple ways Steven Lingafelter private collection, Washington, D. C. (SLPC) men, elytra, hind wings, and genitalia were for taxa with ambiguous states. Texas A&M University, College Station, Texas Character descriptions and states.-A to- (TAMU, Horace Burke, Ed Riley) disarticulated. The wings, and in some National Museum of Natural History, Washington, D. C. cases, genitalia, were placed directly into tal of 102 derived character states for 70 (NMNH, Terry Erwin and Gloria House) vials of 50% glycerin/50% of 80% ethanol. characters was used in the phylogenetic Universidade Federal do Viçosa, Minas Gerais, Brazil (UFVB, Dr. Fiuza) The remainder of the beetle was transferred analyses. These were entered and main- Universidade Federal do Paraná, Curitiba, Paraná, Brazil (DZUP, Solange Napp) to a vial containing a 5-10% KOH solution, tained using MacClade software (Maddison Universidad Nacional Autonomia de México, D. F. and was carefully heated for 10-30 rnin- and Maddison 1992). As these are present- (UNAM, Silvia Santiago Fragoso) utes, depending on size. This procedure ed in an extensive matrix (Table 2), and the University of Nebraska State Museum, Lincoln, Nebraska character states are thoroughly discussed (UNSM, Brett Ratcliffe) caused digestion of the muscle tissue which University of Colorado Museum, Boulder, Colocado otherwise would obscure characters of the below, the matrix substitutes as a presen- (UCMC, Virginia Scott) MEMOIRS O1F THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. LINGAFELTER: GENERA OF ELAPHIDIINI 13 tation of detailed descriptions for each ge- found, however, that these ratios did not nus. Additionally, male genitalia were ex- adequately account for the differences in amined for a11 available taxa and (as ex- femoral shape and they also varied contin- plained below) not found to be useful for uously. Therefore, I excluded these char- the analysis or key. Leg shape was also ex- acters from the phylogenetic analyses, but amined and these data were included in the I did include leg-shape descriptions in the key but could not be coded for the phylo- key, coding them multiple ways for bor- genetic analyses. A discussion of all char- derline values. acters examined in this study follows. Characters not used in analyses or key.- Genitalia of many genera of Elaphidiines The following is a description of a11 the and related taxa were examined. In males, characters used in the phylogenetic analy- variation was discovered in the length of ses. Some of these characters were also the parameres, paramere setae, and shape of used in the key. Character and state num- the eighth tergite (Fig. 3). In females, vari- bers refer to the data matrix (Table 2). This ation was observed in the position of the matrix, combined with the detailed charac- stylus of the coxite and the length and num- ter and state discussion below, serves the ber of setae present on the stylus (termi- function of generic descriptions. Many of nology based on Saito 1989). Because the following characters and their states are males and females were not available for illustrated (Figs. 1-49). Additional morpho- a11 terminal taxa and the noted variation logical information is presented in Figs. 1- was not discrete, genitalic characters were 5A, 6, 8-10, and 49 as a reference aid for not included in the phylogenetic analyses. the characters used in the analyses. The ter- Since the key was intended to be practical minal taxa used in the analyses are meant and not require dissections, genitalic infor- to be representative of the genera and fine mation was not included. morphological details such as setae and Characters used in key on1y.-Femoral punctation are not always shown, since shape (Fig. 37) was found to vary tremen- these attributes can vary among individuals. dously among genera. Historically, the No comments are included here as to prim- terms "clavate," "pedunculate," and "lin- itive and derived states since a11 terminais ear" have been applied to qualitatively de- included in the analyses were treated as po- fine this variation. I attempted to devise tential ingroup taxa. For most characters, formulae based on six measurements of the comments are included here regarding the femur that could give a quantitative and states possessed by terminal taxa (including consistent basis for these terms. I present the provisional outgroup taxa). an example (Fig. 47) which shows and de- Character 1.-Omrnatidial size (Fig. 39): fines these measurements and how they are (0) large (coarse), Fig. 39B; (1) small (fine), used. A11 measurements and variables are Fig. 39A. Taxa with coarse ommatidia usu- defined in that figure legend. Two ratios ally have the lower eye lobe occupying are used, "CR" and "PR." I developed the >50% of the head when viewed laterally. ratio "CR" to determine presence or ab- Each coarse facet is relatively large and Figr 1-4. Morphology of Elaphidiini. 1. Lateral habitus of Elaphidion mucronatum (head pronotum. ely- sence of clavate femora. CR is x/y from convex with light reflected only from the tmn. hindwing. and legs removed). (A) metanotum; (B) mesonotum; (C) mesepimeron; (D) mesepistemum; (E) Fig. 47. I developed the ratio PR to deter- outermost point. The overall surface of the mesosternum; (F) metepisternum; (G)metasternum; (H) metacoxa. 2, Venter of head of Eluphidion mucronatum. mine presence or absence of pedunculate coarsely faceted eye appears uneven. Most (A) labium; (8)maxilla. 3 Ventral view. variation in male genitalia of (A) Phoracantha semipunctaiu (Fabncius) femora. PR is A/B from Fig. 47. If PR and taxa with fine ornrnatidia have smaller eyes, and (B) Eburia haldemani LeConte. (a) eighth tergite; (b) parameres; (c) eighth sternite; (d) sternite-8 apodeme; (e) median lobe Terminology from Fragoso (1985) and Fragoso, et a1 (1987). 4, Hind wing of Aneffurprotensur CR > 3, then I determined the femur to be with the lower lobe occupying -30% of the with terminology of Kukalová-Peck and Lawrence (1993). pedunculate. If CR > 2.5, but PR < 3, then head when viewed laterally. Each fine facet the femur is clavate. If CR < 2.5, then the is relatively small and reflects light evenly femur is gradually enlarged or linear. I from a11 points. The overall surface of the LINGAFELTER: GENERA OF ELAPHIDIINI 15 14 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 Table 2. Continued. Table 2. Character state matrix for taxa of Elaphidiini and other Cerambycinae used in phylogenetic analysis.

1 2 3 4 5 6 7 1 2 3 4 5 6 7 1234567890 1234567890 1234567890 TAXON 1234567890 1234567890 1234567890 1234567690 1234567890 1234567890 1234567890 TAXON 1234567690 1234567890 1234567890 1234567890 Neotrichophoroides p. 010002101 1 01 10000012 120001 1101 40001 0001 1 0000200001 2221000000 1000210000 Ambonus d. Nephaliodes r. 000002101 1 o1 12001012 1100001 100 000010201 1 001000101 1 221 1000000 1000100000 Aneflomorpha r. Nesodes i. O010121011 1011102012 1200010100 6000101011 0000010011 2220010000 1000210000 Aneflomorpha I. Nesosfizocera f. O00002101 1 01 12000010 10001 11101 1000100011 0010010011 1111000000 1000200000 Aneflomorpha p. Nesostizocera v. 0000021011 01 1200201 1 10001 11101 1000100011 001000001f 1201000000 1000210000 Aneflomorpha f. Nyssicostylus a. 002002101 1 01 10101001 120001 0100 400010001 1 0001201 001 2221000200 1000210000 Aneflus p. Nyssicus t. 0000021010 11100~0000 1201010100 4000100001 0001001001 0001000200 1000210000 Aneflus b. Orion p. 0000021011 O011 102010 1000010101 0000101012 0000001001 2220010100 10001 10100 Anelaphus m. Orwellion g. O110121010 0112100012 1200010100 6000101002 0000012011 2220000100 1000210000 Anelaphus p. Panfonyssus n. 0000021011 0112001012 1100010100 0000101011 0010000011 1011000000 1000210000 Anelaphus S. paramallocera c. 000002101 1 00101 00010 1200010100 0000102001 0000000001 1 11 101 0000 1000210000 Anopliomorpha r. parastizoeera p. 000002101 1 01 1000001 1 100001 1100 0000101 01 1 0010?10011 101 1 O00000 1000210000 Anoplocurius a. Parelaphidion i. 0010121010 1012100012 1200010100 6000101011 000020201 1 2220010100 1000210000 Aposphaerion I. Peranoplium u. 0011121011 1011012012 120001010? 6000101001 0010012011 2220000100 1000210000 Appula I. Peranoplium S. 0011121011 1011012112 1300010100 6000102001 0010012011 2220000100 1000210000 Astromula n. Peranoplium h. O01 112101 1 10110121 12 1300010100 6000102001 001001201 1 2220000100 1000210000 Atylostagma p. Periboeum a. 0000021010 1110000001 1200011100 100010001? 0000000011 2211000200 1000?10000 Axestinus o. Phoracanfha S. 0000021010 0012000010 100001010? 0000102012 0010000001 0001010000 1000210000 Castiale e. Poecilomallusp. 00001?1010 101 1002012 1000010101 2000101001 001021 101 1 2221000100 1000210000 Centrocerum e. Protornallocera h. 002002101 1 01 12100001 1200010100 000010101 1 0000201001 2221000000 000021 0000 Championa e. Protosphaerion V. 000002101 1 0010101012 1200010101 000010101 1 00000000?? 22210101 00 1000210000 Clausirion c. Pseudomalloceraa. 0000021011 1112100011 100001010? 2000101011 0002011001 1111002000 0000210010 Conosphaeron S. Pseudoperiboeums. 0000021011 0112001010 1200011101 100010?011 0010001011 2220000000 1000110000 Curtomerus f. Psyrassa si. 0000001011 0111002112 1100000100 1000101011 0010000011 2221000000 1000000000 Elaphidion m. Psyrassa b. 0000011011 0111012112 1100000100 1000101011 0010010011 2221000000 1000000000 Elaphidion I. Psyrassa st. 000001 101 1 o1 11002112 1100000101 100010101 1 0010010011 2221000000 1000100000 Elaphidion e. Psyrassaforrna n. 00?0001011 1101002012 1200011 101 100010101 1 0010000011 2220000200 1000000000 Elaphidion p. Rhomboldederes o. 000002101 0 01 10001 000 101 001 01 00 400010201 1 0000010001 2221 000200 1000210000 Elaphidion S. Sphaerioeme r. 0000??1011 0110102002 1000010101 4000100011 0000200001 2221000000 1000000000 Enaphalodes a. Sphaerion ca. 0020021011 0110000000 1200011100 4000101011 0001201100 2221000200 1010210000 Eurysthea o. Sphaerion cy. 0020021011 0110001000 1200011100 4000101011 0001201100 2221000200 1010210000 Eustromula v. Sphaerionillumc. 0000021011 0112002112 1000010101 1000102012 0010201011 2111000000 1000211000 Eutrichophoroides a. Stenelaphvs a. 00000?1021 0012100012 1200010100 1000112001 0000010011 2220000200 1000210000 Eutrichophoroldes i. Stenosphenus S. 1000021011 0112012012 1000011101 1000101011 0010100011 2210000000 1000200000 Gymnospyra m. Stizocera I. 0000021011 O1 12001012 1100101 101 100010001 1 001000001 1 0101000000 1000210000 Ironeus S. Stizocera p. 0000021011 0112002012 0000101101 1000101011 0010000011 0101000000 1000210000 Ironeus d. Stizocera a. 0000021011 O1 12001012 1000101100 100010001 1 0010000011 1101000200 1000210000 Mallocera g. Terpnlssa I. 0000021011 0110112002 1000010000 4000101011 0001101000 2221000100 0000210100 Meganeflus f. Trichophoroidesn. 0100021011 1110000012 1200010100 4000102011 0000200001 2220000200 1000210000 Megapsyrassa x. Ceresium sp. 0000101001 1012000012 1200010101 ?00010101? 00?0000071 2221010000 1000110000 Mephritus d. Ectenessa v. 0000021001 1112001012 1200010000 4001100011 00?1210010 2221000000 1000210000 Metironeus h. Allofraeus S. 0000021011 0010001012 1200010100 0000101002 0000212001 2221000000 1000210000 Micraneflus b Neocompsa t. 0010021001 0112102112 1100010101 4000100011 0011010010 222?010000 1000210000 Micropsyrassa b. Eburia q. 0010020001 0012011012 1000011310 0000102011 0010000011 0101010000 1000210000 Miltesthus m. Achryson S. 0010021001 0010002012 1200010000 1000101001 0001201001 2221010000 10001 1 O000 Miopteryx S. Batyle S. 1000000000 0011012012 1100011000 4000101012 0010200000 2221010000 1001210000 Morphaneflus p. Hesperophanes sp. 0010020000 001 1101002 1200010000 4000101001 00031 10000 2221010200 10001 10000 Neaneflus f. Coleoxestia n. 00?0101000 2012012011 1000010300 000010101? 0010001000 2221010200 1000010000 Neornallocera o. Cordylomera sp. 0000020010 101 1001 112 110001 1101 400120101 1 00001 10000 2221010200 1001010000 Neoperiboeum j. Neotrichophoroides d

Zum Bates, Stenosphenus Haldeman, and levels which contain many taxa of differing hair as in Orwellion Skiles to large, contig- finely faceted eye appears smooth. This Tropimerus Giesbert. behaviors. My studies show ornmatidial uous patches as in Linsleyonides Skiles. character has been used since Lacordaire Character 2.-Supraocular pubescence: size to be consistent within genera, how- Character 3.Right mandibular tooth (1869) for classification of Cerambycidae. (O) absent or not differentiated from sur- ever. Virtually a11 traditional elaphidiine (Fig. 30): (0) elongate, planar to indistinct Because ornmatidial facet size is related to rounding head region; (1) present and dense taxa have coarsely faceted eyes. Those with (Fig. 30C); (1) short, distinct and often bi- the diel activity of the beetle (finely faceted (Fig. 19). Patches, when present, range in finely faceted eyes include: Championa dentate (Fig. 30A); (2) elongate and mesa- in diurnal adults, coarsely faceted in noc- sim from small separated regions of dense like (Fig. 30B). Mandibles in Cerambycidae turnal adults), it cannot be used at higher Bates, Ironeus Bates (in part), Sphaerionil- 16 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, NO. 20 LINGAFELTER: GENERA OF ELAPHIDIINI 17 are asymmetrical, with notches on one cor- 5B); (1) wide, apically positioned patch responding to teeth on the other (Fig. 29). with indistinct edges (Fig. 5A); (2) narrow, To avoid inconsistency in designating these apically positioned patch with defined edg- Digitiform sensilla region character states, I compared only right man- es (Fig. 5C, D); (3) digitiform patch absent; dibles among taxa. State O is recognized by (4) narrow, distinct, basally positioned not having any indentation in the incisor re- patch. The digitiform sensilla, which can Galea gion towards the apex of the mandible, occur on the terminal labial palpomeres when viewed from the ventral side. State 1 (see Character 5) and maxillary palpo- meres, are widespread and variable in bee- is recognized by having an apical and basal Palpifer indentation, creating a raised plateau which tles (Lawrence and Newton 1995) and par- Lacinia also helps to define the prosthecal region. ticularly so in Chrysomeloidea (Mann and Stipes States O and 1 are widespread among ter- Crowson 1984). This character can only be Cardo minal taxa. State 2 is recognized by having seen effectively with Nomarski interferente two apical indentations separated by a small compound microscopy or scanning electron tooth so that the apex appears bidentate. rnicroscopy. In a11 Cerambycidae examined This state is much less widespread, occur- except Championa, the digitiform patches nng only in Appula Thomson, Mallocera occurred towards the outer apex of the ter- Audinet-Serville, Miltesthus Bates, Nyssi- minal palpomeres. This supports the obser- costylus Melzer, Protomallocera Martins vation of Mann and Crowson (1984) who and Napp, and Sphaerion Audinet-Serville. noted this fundamental difference between Morphological terms for mandible are from Chrysomelidae and Cerambycidae, with Lawrence, et al. in Stehr (1991) and Napp most Chrysomelidae (except some Sagra (1994). and Timarcha) having basally positioned Character 4.-Subapical incisor region: sensillum patches. The well-defined narrow, (0) narrow; (1) wide. State O is the general apical patch (State 2) is most widespread in condition among a11 Elaphidiini and related the terminal taxa of this study. taxa examined and is characterized by a Character 7.-Apex of terminal maxil- narrow width between the dorsal and ven- lary palpomere: (0) not expanded, width Fig. 5. Left maxillae of Elaphidiini (ventral view). (A) Centrocerum exornatum; (B) Curtornerus$avus; (C) Elaphidion mucronatum; @) . tral margins of the subapical incisor region much less than half length; (1) moderately (much less than one-third width of base of expanded (Figs. 5A, C, D); (2) very ex- mandible viewed from mesal, biting sur- panded, greater than three-fourths length absent (Fig. 13); (1) present and acute shape: (0) linear (Fig. 22,45A, B); (I) grad- face). State 1, shared by Peranoplium and (Fig. 5B). The moderate apical expansion (Figs. 8B, 9B, 45,46); (2) present and blunt ually widened at apices (Fig. 45C); (2) fully Anelaphus, is characterized by having of the terminal maxillary palpomere occurs (Fig. 28). Nearly all traditional elaphidiine appendiculate (Fig. 13). The majonty of widely separated dorsal and ventral planes in all traditional elaphidiine genera except taxa possess mesa1 antennal spines on at taxa examined have antennomeres which of the subapical incisor region (distinctly Curtomerus Stephens, which has greatly least the third antennomere. A11 preliminary are approximately parallel-sided for the en- greater than one-third width of base of man- expanded palpi. Most taxa in other Cer- outgroup taxa lacked these spines except tire length, i.e. linear or gradually widened dible viewed from mesal, biting surface). ambycinae tribes (and other subfarnilies) for Allotraeus and Cordylomera. Anoplo- at the apices. Only AneJlus, Axestinus, Mi- Character 5 .-Digitiform sensillum patch have unexpanded terminal maxillary pal- curius Fisher, Axestinus (Fig. 45), Curto- craneJus, Neanejlus Linsley, and Coleo- on terminal labial palpomere: (0) present; pomeres. merus, Elaphidion elegans, Eurysthea xestia Aurivillius have the apices greatly (1) absent. See descriptive discussion of Character 8.-Maxillary palpomeres 3- Thomson, MicraneJlus Linsley, and Mor- expanded. this feature in next character. Digitiform 5: (0) short and thick (each less than 1.5X phaneJus Martins and Napp lack these Character 12.-Antennal carinae: (0) ab- sensilla are absent from the labial palpi in longer than wide) (Fig. 5A-D); (1) elongate spines. Some AneJlomorpha Casey species, sent; (1) single or double carinate (Fig. the speciose, primarily North American and narrow (each much greater than 1.5X Centrocerum Chevrolat, Micropsyrassa 45B, C). Antennal carinae are narrow ridg- genera including Anelaphus, Peranoplium, longer than wide). The unique slender con- Linsley, Neoperiboeum Linsely, and most es where the cuticle appears to be abruptly and Elaphidion. Most other taxa examined dition of this character in Miopteryx spini- Stenelaphus alienus Linsley specimens pinched in a long, straight line. Sometimes possess these sensilla on the labial palpi. ger Blanchard was not seen in any other have blunt spines on the third antennomere. antennomeres are sulcate (Fig. 45A) and su- Character 6.-Digitiform sensillum patch taxa. A11 other exarnined terminal taxa have Character 10.-Lateral antennal spines: perficially appear carinate, however, this is on terminal maxillary palpomere (Fig. 5): shorter palpomere dimensions. (0) present (Figs. 8A, 9A); (1) absent. a result of a broad depression and shadow (0) diffuse patch with sparse sensilla (Fig. Character 9.-Mesa1 antennal spines: (0) Character 11. -Lateral antennomere which sometimes occurs along its edge. LINGAFELTER: GENERA OF ELAPHIDIINI MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, NO. 20 ~hischaracter, like ornmatidial structure, has been used historically to Separate Sphaerionini (carinae present) from the rest of Elaphidiini (carinae absent) (Lacordaire 1869; Linsley 1936; Linsley 1961b). Lins- ley (1936) acknowledged, however, that this character was not absolute in that it oc- curred in some species of Elaphidion. I have found antennal carinae to be wide- spread in Elaphidiini, with no regional pat- terns. Carinae were absent from a11 provi- sional outgroup taxa examined, except Neo- compsa Martins. Character 13.-Antennal segmentation: (0) 12 antennomeres; (1) 11 antennomeres. Most species of Anefus and all Axestinus, Etymosphaerion Martins and Monné, and Psyrassafoima Chemsak have antennae dis- tinctly 12-segmented. Many additional taxa (including some AneJlus species) have the eleventh segment constricted, superficially appearing 12-segmented (Fig. 48), but this is not the case (see discussion of Character 15). Character 14.-Length of third anten- nomere of male: (0) approximately length of pronotum or greater (Fig. 46C); (1) about half length of pronotum (this state also present in females) (Fig. 46A); (2) approx- imately two-thirds length of pronotum (Figs. 8, 46B). This character is sexually dimorphic. Generally, males have longer antennae than females in Elaphidiini. This Fig. 11. Fossils from Florissant, Colorado (MCZC) is particularly evident in the length of the placed in Elaphidiini by Wickham (1914). (A), (B) Ane- third antennomere. For this reason, com- laphus extinctus (Wickham); (C) Stenosphenus pristinus paring females of genus "a" with males of Wickham. genus "b" would be inappropriate. Character 15.-Pseudosegmental setae on terminal antennomere: (0) present (Fig. (1929: 118-1 19), who coined the term 48); (1) absent. The setae, if present, occur "pseudo-segment" to refer to the modified in a poorly defined post-medial ring around eleventh antennomere. Most taxa examined antennomere 11 and are often associated have these pseudosegmental setae present. with a small constriction. I hypothesize that Character 16.-Median pronotal callus: these setae are serially homologous to those (0) present (Fig. 8, 12D, E); (1) absent (Fig. at the apex of the other antennomeres and 12A-C, F-I). A median pronotal callus is a hgs. 6-10. Lateral closure of mesocoxa and spination terminob, in Elaphidiini. 6. Mesocoxa closed laterally provide evidence of (1) fusion of two an- well-defined slightly raised region in the in Nesostirocem~ondana.(A) mesosternum; (B) mesepistemum; (C) metastemum; (D) mesepimeron; (E) closun tennomeres, or (2) incomplete expression of center of the pronotal disc with little or no separating mesocoxa from mesepimeron; (F) mesocoxa; (G) metepistemum. 7, Mesaoxa open laterally in Mesap- a gene coding for a twelfth antennomere. syrusro xestioder 8. Dorsal habitus of Elaphidion spinimm. (A) lateral antennal spine; (B) mesal antennal spine; punctation and pubescence. In some taxa, a (C) lateral femoral spine; (D) mesa1 femoral spine; (E) sutural elytral spine; (F) apicolateral elytral spine. 9. This character has been discussed in Carter central region is defined, but this is due to Enlargement of Fig. 8, showing apex of left elytron. (A) laterd antennal spine; (B) mesa1 antennal spine; (C) laterd femoral spine; (D) mesa1 femoral spine 10, (E) sutural elyaal spine; (F) apicolateral elytral spine.

LINGAFELTER: GENERA OF ELAPHTDIINI MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 23

Character 28.-Prosternal projection variously modified with regard to the an- shape between procoxae: (0) linear, without terior margin and mesostemal process be- expansions (Fig. 14); (1) broad and ex- tween the mesocoxae. State 6 occurs only panded apically only (Fig. 18); (2) broad in the North Amencan genera, Anelaphus, and expanded laterally only; (3) broad and Nesodes Linsley, Orwellion, Parelaphidion, expanded apically and laterally. The linear and Peranoplium. It is like state 2 (Fig. prostemal projection which occurs only in 35F) except for the anterior margin being Anoplocurius, Terpnissa Bates, and Ecte- abruptly directed posteriorly at the sides. nessa Bates, is recognized as a vertical pla- State 3 occurs only in Aposphaerion Bates. nar projection lying between very narrowly The other states are widespread among the separated procoxae. All other states have terminal taxa. the prosternal projection broad and flattened Character 32.-Mesosternum between horizontally between the procoxae. Most coxae: (0) planar or slightly angled; (1) taxa have state 1, with a slight or strong abruptly vertical. The abrupt vertical ori- expansion at the apex. Eburia Lepeletier entation of the anterior margin of the me- and Coleoxestia have state 3, with expan- sostemum is found only in Championa. sions between the procoxae as well as api- Character 33.-Mesosternal process cally. width: (0) very wide; (1) narrow. The very Character 29.-Abruptly declivous pro- broad mesostemal process, separating the stemal projection (Fig. 44): (0) absent (Fig. procoxae by a distance greater than their 44A); (1) present (Fig. 44B). State 1 is diarneter, is found only in Anopliomorpha. present only in Elaphidion and Eburia. In Character 34.-Mesostemal notch (Fig. some Elaphidion there is also a posteriorly 35): (0) present; (1) absent. The notch at directed process extending into the mesos- the posterior margin of the mesostemum is ternum. absent only in Ectenessa and Cordylomera. Character 30.-Lateral mesocoxal cavi- Character 35.-Mesonotal lateral projec- ties: (0) open to mesepimeron (Fig. 7); (1) tion orientation: (0) anteriorly oriented (to- closed to mesepimeron (Fig. 6). The states wards head); (1) medially oriented between of this character are interpreted slightly dif- anterior and posterior margins of mesono- ferently from Linsley and Chemsak (1984, tum (Fig. 36A-F); (2) postenorly oriented Fig. 1). When the open state is present, the (towards scutellum). A11 taxa examined Figs. 13-22. Anatomical characters in Elaphidiini. 13, Aneffu, obscurus, dorsal view of Ieft antenna and elytron, mesostemum and metastemum do not con- have the lateral projections of the mesono- (A) laterdly appendiculate antenna (lateral is to the Ieft); (B) "typicd" recumbent pubescence. 14, Terpnissa tact one another laterad to the mesocoxae. tum medially oriented except for Cham- 1istropteM. oblique ventral view of cleared prothorax, showing linear pmsternal projection. 15, Elaghidion por- piona and Astromula Chemsak and Linsley, toricensi.~,dorsi view of elytron and pronotum, showing epipleurd tooth. 16. Stizocem poeyi, latml view of This leaves an open space which allows elytron, showing "flying hairs." 17, Nyssicus topographicus, ventrd view of pmthora showing latedly open contact between the mesepimeron and the which have them anteriorly oriented, and procoxal cavities 18, Anelaphus moestus, posterior view of cleared prothorax. showing postefiorly open procoxal mesocoxae. When the lateral mesocoxal Cordylomera which has the arms posteri- cavities and moderately expanded procoxd process apically 19, Linsleyonides chemsaki, dorsal view of head and cavities are closed, the coxae are complete- orly oriented. pronoNm. showing post-ocular pubescence patches. 20. Sphaenon sp., venlral view of cleared pterofhorax, hohounng ly surrounded by the meso- and metaster- Character 36.-Mesonotum lateral arm me~~coxalprojections from mesostemum absent. 21. Castiale elegann

Figs. 23-28. Morphological characters in Elaphidiini. 23, Tropimerus hovorei, mesofemoral carina. 24, gen- eralized laterally closed procoxal cavity. 25, metafemoral apiccs of Pantonyssus nigriceps. (A) spinose laterally; (B) dentiform rnesally. 26, metafemoral apices of Anelaphus moestus. (C) rounded laterally; (D) rounded mesally. 27, Megapsyrassa xestiodes, ventral view of prothorax; arrow shows procoxal cavities closed posteriorly. 28, Neoperiboeum juanitae, base of antenna (first antennomere on left), showing blunt spine on antennomere 3.

Figs. 29-34. Morphological characters in Elaphidiini. Arrows indicate anterior. 29, Elaphidion mucronatum, 36A). The rounded scutellum is widespread both by rounded protuberances (condyles) dorsal view of head showing mandibular asymtry. 30, Mandibular states. dorsal views of nght mandible. (A) among most taxa examined; the acute state of the mesosternum between the mesocoxae Astromula nitidum; (8) Protomallocera hilairei: (C) Nephaliodes rutilur 31, Lateral view of mesothorax, showing and corresponding concave depressions (ac- mesepisternal states (mwpoints to anterior). (A) Anelaphus moestus; (B) Pseudomallocera auri@a; (C)Miopteryx less so. The truncate scutellum occurs only spiniger. 32, Lateral view of pterothorax, showing metepisternal-elytral junction in Megapsyrassa huberi (heavy in Curtomerus and the notched state occurs etabular excavations) on the coxae. As with stippling indicates region of metepistemum ventral to keel). 33, Lateral view of pterothorax, showing metepistemal- only in Meganeflus Linsley. most characters, clearing is necessary to ac- elytral junction in Psyrassa sp. (heavy stippling indicates region of metepisternum ventral to keel). 34, Metacoxal Character 41.Basal width of scutellum: curately determine the state present. Both ndges in Anelaphus moestu.~moestw. (A) view of the anterior margin; (B) view of the posterior margin. (0) approximately equal to width of meso- states are widespread in the examined taxa. notum; (1) much wider than mesonotum. Character 44.-Mesepisternal carina (Fig. 31): (0) abruptly angled carina (Fig. mallocera Zajciw. State 3 occurs only in (see Fig. 49) that determines the shape of The wide scutellar base occurs only in Neo- Atylostagma White, Miopteryx, and Hes- periboeum. 31A); (1) carina absent; (2) rounded carina this notch. If both sclerites come together perophanes Dejean. Character 42.-Junction of mesonotum (Fig. 3 1B); (3) straight, incomplete carina at an angle of less than 45 degrees, then the Character 45.-Shape of metasternal and scutellum: (0) free dorsally; (1) com- (Fig. 31C). The states of this character can notch is acute or blunt. An acute notch has only be seen with moderate clearing of the notch: (0) acute; (1) rounded; (2) blunt. The a pointed apex; a blunt notch has a rounded pletely fused. The fused condition occurs metastemal notch is the region of the meta- only in Miopteryx. cuticle. State O is the most widespread. apex. If both sclerites come together at an State 1 occurs only in Anopliomorpha, sternum that receives the antero-medial ex- angle greater than 60 degrees, then the Character 43.-Lateral projections into tension of the first abdominal ventrite. It is mesocoxae from mesosternal process: (0) Championa, Mephritus, Nyssicostylus, Nys- metastemal notch is rounded. The acute sicus, Sphaerion, Terpnissa, Ectenessa, and the angle of the sclerites on the inner, an- state is most widespread among the taxa ex- absent (Fig. 20); (1) present (Fig. 21). The terior margin of the posterior coxa1 cavities presence of lateral projections is defined Achryson. State 2 occurs only in Pseudo- amined. Only Clausirion Martins and Napp, 26 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON, No. 20 LINGAFELTER: GENERA OF ELAPHIDIINI 27

Eurysthea, Stenosphenus, Terpnissa, Hes- the cuticle is usually membranous, and be- perophanes, and Cordylomera have the low which it is more heavily sclerotized. rounded state. Both states are widespread among exam- Character 46.-Metasternal sulcus: (0) ined taxa, although the dorsally-positioned complete; (1) incomplete. The metasternal carina is more common among provisional groove is an invagination corresponding to outgroup taxa. the interna1 metendosternite. The typical, Character 50.-Metepisternal sclerotiza- complete condition is present when the tion: (0) even sclerotization above and be- groove runs anteriorly to near the plane at low keel; (1) Highly sclerotized ventrally the posterior margin of the metacoxae (Fig. below keel and membranous dorsally (Fig. 49). The incomplete condition does not at- 38A-D). State 1, differential sclerotization, tain the coxae and typically extends ante- is widespread among most taxa examined. riorly only half the length of the metaster- Those taxa with even sclerotization above num. The complete state is widespread. The and below the keel include: Appula, Atylo- incomplete condition occurs in many West- stagma, Championa, Clausirion, Sphae- D Indian genera not included in the analyses: rion, Terpnissa, Ectenessa, Neocompsa, notch Linsleyonides, Nesanoplium Chemsak, Ne- Batyle Thomson, Hesperophanes, Coleox- siosphaerion Martins and Napp, and Ela- estia, and Cordylomera. phidionopsis Linsley. Character 5 1 .-Mesal mesofemoral api- Character 47.-Posterior notch of met- ces: (0) spinose (as in Figs. 8D, 9D); (1) episternum (Fig. 38): (0) absent or subtle dentiform (Fig. 25B); (2) rounded (Fig. indentation (Fig. 38A); (1) narrow, shallow 26D). Spinose femoral apices are acute, indentation (Fig. 38C); (2) wide and deep pointed processes extending beyond the tib- (usually extending at least one-third the ial insertion and having an angle about the 35 width of the metepisternum at the posterior apex of less than 45 degrees. Dentiform edge, below the keel) (Fig. 38B, D). This femoral apices are pointed at the apex, do notch, if present, occurs on the posterior not extend much beyond the tibial insertion, margin or postero-ventral margin of the and have an angle about the apex of greater metepisternum, and is most easily seen with than 60 degrees. Rounded femoral apices some clearing of the cuticle. A11 states are have a rounded apex and do not extend widespread among the exarnined taxa. much beyond the tibial insertion. Dentiform Character 48.-Metepisternal notch pro- and rounded states are widespread among jection: (0) absent; (1) small lateral bump the exarnined taxa. Those with spinose mes- present. If present, the projection is located a1 mesofemoral apices include: Atylostag- in the region of the metepistemal notch, on ma, Castiale, Elaphidion, Nyssicus, Phor- scutellum the postero-ventral margin of the metepi- acantha, Stizocera, and Eburia. median projection sternum. This is best seen by looking down Character 52.-Lateral mesofemoral api- the metepisternum from the anterior end. Of ces: (0) spinose (as in Figs. 8C, 9C); (1) the specimens exarnined for the phyloge- dentiform (as in Fig. 25B); (2) rounded netic analysis, this projection was found (Fig. 26C). See state descnptions for Char- only in Sphaerion. Subsequent exarnination acter 51. Dentiform and rounded states are of non-dissected Mephritus species has re- widespread among the examined taxa. vealed a small protuberance in the same re- Those with spinose lateral mesofemoral gion of the metepisternum. apices include: Appula, Castiale, Neomal- Character 49.-Metepisternal keel posi- locera, Pantonyssus Bates, Parastizocera, tion: (0) dorsally positioned (Fig. 38B); (1) and Phoracantha. 36 approximately midway positioned (Fig. Character 53'-Mesa1 metafemoral Figs. 35-36. Mor~hologicalcharacters in Elaphidiini. Arrow indicates mtenor. 35, Mesostemd "uiation, arrow indcates mfenor. (A) Miopter-yx spiniger; (B) Conosphaeron concolor; (C) ,yaphjdjon mucroMtum; (D) caStiaze 38A, C, D). The keel is the longitudinal (O) (Figs- 8D' 9D); denu- ezegantuk (E) A~osphaerionlongicolk (F) &pula hferali.7. 36, ~~~d "uiation. (A) ~~~~~~ffw~~~~~~~~~~; ridge along the metepisternum above which Iorm (as in Fig. 25A) ; (2) (Fig. (B) f~kerioninus;(c) Parelaphid~onaspersum; (D) Trjchopbroider njVeUS; (E) paraStiZoCeraprocera; (F) Penboeum acumimtum. LINGAFELTER: GENERA OF ELAPHIDIINI MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON. No. 20

jansoni Bates. Trichophoroides ...... 95 Parasphaerion Martins & Napp jassuara Martins & Napp. Stizocera ...... 93 ...... 81. 99 Parastizocera Linsley ...... 78. 81. 99 hcordairei Lacordaire Paramallocera ...... 80 INDEX . Parelaphidion Skiles ...... 78. 82. 102. 104 ...... Ihnpracantha Thomson ...... 79 acuminatum Thomson. Periboeum ...... 82 Cylindrodera Gemrninger & Harold 60 Lampracanthus Thomson 79 patagonius Guérin-Méneville. Orion ...... 39. 78 ...... Peranophm Linsiey ...... 34. 52 Adiposphaerion Martins & Napp ..... 47. 48. 100 Cyrtomerus Gemminger & Harold 60 Lampromerus Thomson ...... 60 ...... Periboeum Thomson ...... 82. 83. 100 albisparsus Bates. Trichophoroides ...... 95. 96 decipiens Bates. Tricliophoroides 95. 96 lateralis White. Appula ...... 54. 55 diabolicus Larneere. Mephritus ...... 66 Phoracantha Newman ...... 39. 104 albomaculatus Champlain & Knull. Linsleyonides 65 ...... Linsleyonides Skiles ...... 65. 66. 98 aliena Linsley. Psyrassa ...... 83 distinctus Newman. Ambonus 48 phtisica Gounelle. Stizocera ...... 93 ...... 1ippu.v Gerrnar. Ambonus ...... 49 alienus LeConte. Stenelaphus ...... 90. 92 dozieri Fisher. Siizocera 93 piceum Chemsak. Anelaphus ...... 52 duplex Bates. Ironeus ...... 65. 66 listropterina Bates. Terpnissa ...... 95. 96 Piezophidion Galileo & Martins ...... 84. 105 Allotraeus Bates ...... 39 longicolle Bates. Aposphaerion 54. 55 ..... 59. 60. 100. 104 ...... pilicornis Fuchs. Trichophoroides ...... 95 Ambonus Gistel ...... 48. 49. 101. 103 Elaphidion Audinet-Serville Iugens LeConte. Stenosphenus ...... 90 Elaphidionopsis Linsley ...... 59. 61. 95. 99 Pilisphaerio Manins & Napp ...... 84. 100 Amethysphaerion Martins & Monné ...... 49. 102 maculatum Chemsak & Noguera. Anelaphus ... 52 Elaphidium Agassiz ...... 60 pilosella Bates. Micropsyrassa ...... 70 amictus Newman. Mephritus ...... 68 magnipunctata Knull. Gymno.rpyra ...... 64 ...... 34. 48. 52. 102. 103 elegans Chemsak. Championa ...... 39. 54 plicicollis Gerrnar. Stizocera ...... 90 Anefomorpha Casey ...... Mallocera Audinet-Serville ...... 66. 67. 99 Aneftus LeConte ...... 38. 48. 51. 102. 105 elegans Chevrolat. Elaphidion 36 Poecilomallus Bates ...... 83. 85. 101 elegantula Perroud. Castiale ...... 54. 57 marginatus Bates. Miltesthus ...... 71. 72 Anelaphus Linsley ...... 34. 48. 52. 102. 103 Me~ane3u.vLinsley ...... 66. 67. 102 poeyi Guérin-Méneville. Stizocera ...... 90. 93 Elaphidionoides Linsley ...... 52 polita White. Atylostagma ...... 57 Anepsyra Casey ...... 50 Me~apsyramaLinsley ...... 66. 68. 104 erichsoni White. Pantonyssus ...... 80 portoricensis Fisher. Elaphidion ...... 36 Anopliomorpha Linsley ...... 48. 52. 101 Mephritus Pascoe ...... 66. 68. 100. 101 Enaphalodes Haldernan ...... 59. 62. 104 Anoplocurius Fisher ...... 53. 54. 98 Metironeus Chemsak ...... 66. 69. 101 pristinus Wickham. Stenosphenus ...... 10. 19 Etymasphaerion Martins & Monné .... 59. 63. 105 Apoclausirion Martins & Napp ...... 55. 99 Micraneflus Linsley ...... 66. 70. 105 procera Erichson. Parastizocera ...... 78. 81 Euristhea Lacordaire ...... 63 Aposphaerion Bates ...... 43. 55. 104 Micranoplium Linsley ...... prolixus Martins & Napp. MorphaneJlus ... 71. 74 Eurysthea Thomson ...... 59. 63. 105 66. 70. 105 Protaneflus Linsley ...... 51 Appula Thomson ...... 54. 55. 100 Micropsyrassa Linsley ...... 70. 71. 100 Eustroma LeConte ...... 63 protensus LeConte. Aneflus ...... 51 aspera Knull. Gymnospyra ...... 59 Miltesthus Bates Eustromula Cockerell ...... 59. 63. 102 ...... 71. 72. 101 Protomallocera Martins & Napp ..... 85. 100. 103 aspersum Haldeman. Parelaphidion ...... 78. 82 Minipsyrassa Martins ...... 73. 104 Eutrichophoroides Linsley ...... 38. 95 Protosphaerion Gounelle ...... 83. 86. 103 Astromula Chernsak & Linsley ...... 54. 56. 105 Miopteryx Blanchard ...... 7 1. 73. 100 ...... eximium Bates. Anelaphus ...... 52 Pseudaneflus Chemsak & Linsley ...... 68 Atharsus Bates 56. 99 moestus LeConte. Anelaphus ...... 48 ...... 93 exornatum Newman. Centrocerum ...... 54. 57 Pseudibidion Casey ...... 87 atiaia Martins & Napp. Stizocera Morphanefus Martins & Napp ...... 71. 74. 105 exoticum Martins & Napp. Pilisphaerion ...... 84 Pseudomallocera Zajciw ...... 83. 86. 98 atomarius Dmry. Enaphalodes ...... 59. 62 Moureana Zajciw 89 extinctus Wickham. Anelaphus ...... 10. 19 ...... Atylostagma White ...... 54. 57. 103 Neaneflus Linsley ...... 71. 74. 102 Pseudoperiboeum Linsley ...... 83. 87. 101. 102 fasciatipennis Linsley. Elaphidionopsis . . 59. 61. 95 Psyrassa Pascoe ...... 36. 83. 87. 102. 103. 104 Aulacoscapus Linsley ...... 72 Neomallocera Martins & Napp ...... 71. 74. 100 ...... flavus Fabricius. Curtomerus ...... 59. 60 Psyrassaforma Chemsak ...... 83. 88. 101 auriflua Klug. Pseudomallocera 83. 86 Neoperiboeum Linsley ...... 7 1. 75. 101 floridana Linsley. Stizocera ...... 93 aurivili Melzer. Nyssicostylus ...... 77. 78 Neotrichophoroides Linsley ...... 38. 95 puberulum Fieutiaux & Sallé. Nesanoplium 76. 95 fracticorne Wickham. Elaphidion ...... 10 aurivillii Linsley. Trichophoroides ...... 95 Nephaliodes Linsley ...... 71. 75. 102 punctiventris Cazier & Lacey. Stizocera ...... 93 fuchsii Wickham. Neanefus ...... 71. 74 quadriguttatus Swedems. Nyssicus 79 Axestinus LeConte ...... 38. 5 1 Newrnan 91 ...... fulvipennis Bates. Meganefus ...... 66. 67 ...... quadrisignatum Bates. Sphaerionillum ...... 92 basicornis Pascoe. Psyrassa ...... 87 Nesanoplium Chemsak ...... 76. 95. 99. 104 gibbulum Bates. Orwellion ...... 78. 79 ravidus Gounelle. Rhomboidederes ...... 83 bicolor Martins. Minipsyrassa ...... 73 Nesiophaerion Martins & Napp ... 71. 76. 95. 104 glabra Schaeffer. Atylostagma ...... 54 rectilinea Casey. Anefomorpha ...... 48 bimaculata Bates. Micropsyrassa ...... 7 1 Nesodes Linsley ...... 77. 78. 103 glauca Audinet.Serville. Mallocera ...... 66. 67 reticolle Bates. Anopliomorpha ...... 48. 52 canotiae Fisher. Anoplocurius ...... 53. 54 Nesostizocera Linsley ...... 36. 93 castaneum Chemsak & Linsley. Sphaerionillum 90 globosus Knull. Romulus ...... 89. 90 Rhomboidederes Zajciw ...... 83. 89. 99 ... 81 nigricauda Bates. Atharsus ...... 56 Castiale Pascoe ...... 54. 57. 100 granulosum Martins & Napp. Parasphaerion Romaleum White ...... 62 59. 64. 102. 103 nigriceps Bates. Pantonyssus ...... 78 caymanensis Fisher. Stizocera ...... 93 Gymnospyra Linsley ...... Romulus Knull ...... 89. 90. 104 ...... nigricorne Martins & Napp. Apoclausirion .... 55 Centrocerum Chevrolat ...... 54. 57. 104 Haplosphaerion Linsley 70 rubristerna Martins & Napp. Sphaerioeme . . 89. 90 ...... nigripes Martins & Monné. Amethysphaerion ... 49 Championa Bates ...... 39. 54. 105 Hemilissopsis Lane 64. 101 rubrum Martins & Napp. Adiposphaerion . . 47. 48 ...... nitida Chemsak. Psyrassafoma ...... 83. 88 chemsaki Skiles. Linsleyonides ...... 66 Hemistizocera Linsley 36. 87 rutilus Bates. Nephaliodes ...... 7751...... nitidum Chemsak & Linsley. Astromula .... 54. 56 Clausirion Martins & Napp ...... 54. 58. 104 hesperus Chemsak. Metironeus 69 simile Schaeffer. Anelaphus ...... 52 ...... hilairei Gounelle. Protomallocera ...... 85 niveus Linsley. Trichophoroides ...... 95. 96 sonoranus Casey. Anefus ...... 48 clenchi Lane. Hemilissopsis 64 notatus Olivier. Stenosphenus ...... 93 ..... 54. 58 hirta Kirby. Paramallocera ...... 78 Sotenus Sharp 60 comptum Martins & Napp. Clausirion Nyssicostylus Melzer ...... 77. 78. 100 ...... heferi Knull. Anelaphus ...... 48. 52 concolor Linsley. Conosphaeron 58. 59 Nyssicus Pascoe ...... 78. 79. 99 Sphaerioeme Martins & Napp ...... 89. 90. 99 Conosphaerion Linsley ...... 58 hovorei Chemsak. Metironeus ...... 66 Sphaerion Audinet-Serville ...... 90. 91. 101 ...... hovorei Giesbert. Tropimerus ...... 95 obliqua Audinet-Serville. Eurysthea ...... 59. 63 Sphaerionillum Bates ...... 90. 92. 105 Conosphaeron Linsley 58. 59. 101 ob.scums LeConte. Ane8u.s ...... 38. 48. 51 ...... Hypermallus Lacordaire ...... 62 Sphoerion Thomson Cordylomera Serville 39 ocellicollis Zajciw. Rhomboideder~ 89 ...... 91 36. 83. 87 imbellis Casey. Micraneflus ...... 66. 70 ...... cribricollis Bates. Psyrassa ...... rpinicome Dmry. Elaphidion ...... 18. 59. 60 ...... inornatum Chemsak & Linsley. Anelaphus .... 52 opulenta Newman. Neomallocero ...... 71. 74 Crocidastus Pascoe 49 Orion Guérin-Méneville ...... 39. 78. 103 rpiniger Blanchard. Miopteryx ...... 71. 73 Curtomeius Stephens ...... 59. 60. 105 insulana Gahan. Stizocera ...... 93 purcus LeConte. Anelaphus ...... 48. 52 ...... 7 1. 76. 95 Orwellion Skiles ...... 78. 79. 103 cyaneus Giesbert. Tropimerus ...... 97 insulare White. Nesiosphaerion itenelaphus Linsley ...... 90. 92. 103 cyanipenne Audinet.Serville. Sphaerion ...... 91 insularis Linsley. Nesodes ...... 77. 78 palpalis Bates. Poecilomallus ...... 83. 85 itenosphenopsis Linsley ...... 93 ...... 84 Pantonwrus Bates ...... 78. 80. 100 Cycliopleurus Hope ...... 60 intricatum Galileo & Martins. Piezophidion itenosphenus Haldernan ...... 90. 93. 105 ...... 36. 38. 65. 66. 105 Paramallocera Aurivillius ...... 78. 80. 103 Cylindera Newrnan ...... 60 Ironeus Bates itizocera Audinet-Serville ... 36. 90. 93. 100. 103