'^'^.X ISOi HARVARD UNIVERSITY Library of the Museum of Comparative Zoology IviUS. COT/.P. ZOOL. LIBRARY Brigham Young University Science Bulletin NOV 29 19b6 UNIVERSITY MITES OF UTAH MAMMALS by DORALD M. ALLRED AND D ELDEN BECK BIOLOGICAL SERIES — VOLUME VIII, NUMBER 1 OCTOBER, 1966 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Dorald M. Allred, Department of Zoology and Entomology, Brigham Young University, Provo, Utah Associate Editor: Earl M. Christensen, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: J. V. Beck, Bacteriology C. Lynn Haywahd, Zoology W. Derby Laws, Agronomy Howard C. Stutz, Botany Wn-MER W. Tanner, Zoology, Chairman of the Board Stanley Welsh, Botany Ex officio Members: RtTDGER H. Walker, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications The Brigham Yoimg University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numibers and back volumes can be purchased from University Publications, Brigham Yoimg University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah. Brigham Young University Science Bulletin MITES OF UTAH MAMMALS by DORALD M. ALLRED AND D ELDEN BECK BIOLOGICAL SERIES — VOLUME VIM, NUMBER 1 October 1966 MUS. COMP. 200L LIBRARY NOV 29 19b6 MMrtVMaU UNIVERSITY. TABLE OF CONTENTS Page INTRODUCTION 1 Topographical and Faunal Features 1 HISTORICAL REVIEW 3 MEDICAL AND ECONOMIC IMPORTANCE 3 ACCOUNTS OF THE SPECIES 4 Pachylaelaptidae 8 Gamasolaelaptidae 8 Neoparasitidae 8 Halarachnidae 9 Zurnptiella bakeri 9 Spinturnicidae 9 Parasphiturnix <:^lohosus 9 Spinturnix orri 9 Parasitidae 9 Macrochelidae 9 Macrocheles sp 9 Haemogamasidae 10 Brevisterna tiwntaiitis 10 Brevisterna utahensis 11 Eulaelaps stahularis 11 hchyropoda funnani 12 Ischi/ropoda armatus 12 Hiwmogtimasus uhiskctisK 13 H aemo<^(ima.sus pontiger 13 H aemogumtuius lipom/ssokles occidentalis- 14 Ilaeinoguiuuxu.'i nri\l>ul(n\)s 14 Hucmogamiisus hmgitarsus 15 Phytoseiidae 15 Klccnumia sp 16 Dermanyssidae 16 Myonyssus ntontuniis 17 Hirstionysiux staff onii .„ 18 Hirst ionysiu-t tursalis 18 Hirst ionysstis uffiiii.s 19 Hir.tlioiiyssus ptihistris 19 Hirstionijssus punctalus 20 Hirstionyssus iniaginatiis 21 Hirst ionyssiis riitiimiiic 21 tlirstionysstis utiilu-iisis 22 Hirstioni/ssus aiigustus 24 Hirstioiiysstis feniuralis .. 25 Hirstiouyssiis lotigichelac 25 Hirstiomjssus tlwtnotnys 26 Hirstionysstis torus 27 Hirsliduyssus iwotoniae 29 Hirstiouyssiis hisetnstts 29 Hirstiouyssiis isiil)cllinus 29 Hirstionysstis triticunthiis 30 Hirstionyssiis hitti 30 Hirstiouyssiis inconiptiis , 30 Hirstionysstis hiicoti 31 Hirstionysstis carnifcx 31 Hirstionys.sus p,ivmijdi.s 31 Hirstiotiyssus ohsoletus 31 IchoTonijssiis robustipes 31 Dernmntjsnus sanguineus 32 Dermcmyssus hechl 32 Dcrnmmjssus gallinae 32 Steatonyssus antrozoi 32 Ornithonyssus aridus 33 OrnHhonyssus hacoti 33 Ornithonyssus sylviurum 33 Laelaptidae 33 Androlaehips leviculus 34 LaeUips kochi 35 Laelaps multispinosus 35 Laelaps incilus 36 Laelaps nuttalli 37 Euhruchylaelaps crowei 37 Euhrachijhelaps hollisteri 38 Euhmchiilaelaps circuhiris 38 Euhrachi/hiclaps dehilis 38 Hypoaspis luhrica t 39 Hypoaspis gurahensis 39 Haemolaelaps casalis 40 Haemolaelaps ghsgowi 40 Haemolaelaps geomys 41 Listrophoridae 41 Myocoptes sp 41 Listrophorus sp 41 Myobiidae ■ 4- Myobia sp 42 Radfordia bachai 42 Radfordia lemina 42 RadfordUi sidiuligcr 42 Trombiculidae 4^ Bernia, new genus 4o Berniti marita 4J Odontacarus linsdalei 44 Odontacarus niicheneri 44 Odontamrus hirsutus 45 Leeuwenhuekia americana 45 Whartoniu pcrph'xti 4o Chutui setosa 45 Chatia ochotomi 45 Guhrlk'pia inncriciina 4d Trotnbicida calif ornica 47 Tromhicula belkini 47 Trombicula hoplai 4o Troml>ictda niyotis 48 Tromhicula potusina 4o Trotnliicula pauamensis 4o Trombicula suhsignata 48 Trombicula liar peri 49 Trombicula jewetti 49 Trombicula sargenti 4 J Tromhicula csocnsis - Tromhicula uniiciri ■"* Tronihiculd hanlosi 49 Tromhicula hakeri i 49 Trotnhiculii urcnitiAci __ 50 Trot)il>wula ttHinUtm nsis 50 Troniliiculu durcini 50 Trumhicuhi altrcdi 50 Chcldihrnla iTo.\.\i 50 EuschocHjjl^axtUi laccrtci 52 Euschi)cnij,a.stui Itofjiiuiiwc 52 Euschoenfj^dstiii jurmani 52 Euschocn'^iistui utulicnsi.s- 52 Euschocni^aslid soricimis 52 EuscltiH-nj^ii.stUi s 95 75 1 80 43 RABBITS AND HARES Lepus sp. L. americanus L. caHfornicus L. toxcnsendii 1 4 561 13 2 50 6 4 18 8 1 2 4,54 2 Sylvihigus sp. S. auduhonii S. idahoensis S. nuttallii 79 48 60 33 16 11 20 16 BlllGIlAM \t )UN(; L'NivEHsriv SciENC:ii BUL .LKIIN 1 43 P. niunicuUitus 3077 1821 723 1 12.54 805 .3 41 P. tnwi 184 63 33 103 23 4 11 Rcithrodontoiiii/s 2 17 nicPalotus 122 37 6 20 11 SQUIRRELS AND RELATIVES MICROTINE RODENTS CU'thrioiwttnis rinis- 8 1 6 1 MURID RODENTS Glaucomys salirinus Murmota fUwiventris 4 52 2 40 1 29 2 9 2 16 Mus inusculus 23 7 6 Sciurus ciherti 1 1 1 1 Ruttus norvegicus 151 25 98 73 6 Spennophilus armatus 2.W 170 132 98 116 R. rati us 9 4 - 3 S. heldingi 4 3 2 S. lateralis 130 76 18 54 82 JUMPING MICE S. richardsonii 7 7 4 1 4 Xapus princcps 96 64 1 25 30 S. spilosoma 2 2 1 S. townsendii 8 5 2 1 2 HVSTRICOMORPH RODENTS S. tridecemlinecitus 5 2 3 1 1 S. variegatus 60 49 7 17 18 Ercthizon dorsatum 36 1 1 22 Tamiasciurus hudsonic us 56 33 11 14 8 FOXES AND COYOTES POCKET GOPHERS Caitis liitraris 4 1 4 Thomomys sp. 12 3 2 6 1 Urocyon cinereoargcnteus 1 1 T. talpoides 151 64 54 37 19 Vutpes mucrotis 4 3 1 T. umhrinus 73 30 47 29 2 RACOONS AND ALLIES HETEROMYIDS Bassuriscus astutus 1 1 Dipodomys sp. 15 1 2 6 Procyon lotor 1 1 D. deserti 7 4 2 5 2 D. merriami 1.52 68 13 63 49 MUSTELIDS D. microps 87 6 11 48 32 Martcs sp. 1 1 1 D. ordii Microdipodops 820 293 222 610 351 Mephitis mephitis Mustek erminea 1 1 1 megaccphahis 6 6 M. freuata M, vis'iiin 11 10 4 5 5 Perogiuithus sp. 63 3 5 24 9 2 2 P. apache P. formosus 12 96 3 1 27 10 55 10 Spilogale gracilis Taxidva taxus U 6 7 1 2 8 3 P. intermedius 1 P. longimemhris 72 1 2 14 30 P. parvus 224 46 19 82 86 CATS Lynx riifus 6 5 2 BEAVERS Castor caiuiden.tis 4 CERVIDS Ceniis caiuidenMs I 1 CRICETID RODENTS Davui heniionus 28 1 25 Neotuma sp. N. albigula 9 6 1 4 4 3 2 1 1 DOMESTIC ANIMALS N. ciiierea 29 25 5 7 6 Cat 4 2 2 N. lepida 149 80 20 56 20 Cou 3 1 N. mexicana 1 1 Dog 16 4 9 Onychomijs sp. 9 7 2 3 Hog 1 1 2 O. leucogastcr 32 27 7 19 9 Horse 7 1 6 O. torridus 4 3 2 2 1 Rabbit 2 1 2 Peromyscus sp. 20 6 1 10 1 Sheep 2 P. hoylii P. crinitus 13 4 1 6 128 49 14 72 20 •M,,„ .■,„„,;„•. ,..a f,.ll,,%v \IM ^m 1 KeUon 1 ( li7.ii„.ii,.ii ,iiT.uiKi.Mii.iii ..( (;ci M«'< '•■- Mites of Utah Mammals HISTORICAL REVIEW Until recent years little was published on p;irasitic mites from Utah. Keegan ( 1949, 1953, 1956b) described a new species, erec-ted a new genus based on Utah material, and listed a num- ber of new records for the state. Allred and Beck ( 1953b ) described a new species of chig- ger { which was later synonymized by Brennan and Beck, 1955), and studied mites of woodrat nests in Utah (1953a). Allred (19.54a, 19.54c, 1957b, 1957c, 1957d, 1958) studied mites found on mice of the genus Feromijscus in Utah, de- scribed a new species (19.57a), discussed mor- phological variations and bionomics ( 1954b, 1957e, 19571, 1957g, 1965), and listed additional host records (1961). Furman and Tipton (19.55) named a new species from Utah and composed a key to the known species of Mi/om/sstis. Bren- nan and Beck ( 19.55 ) ' listed host and distribu- tion records, named new species, and included a key to the chiggers of Utah. Lipovsky, Crossley and Loomis ( 1955 ) also named a new genus and species of chigger from Utah. Woodbury (1956a, 1956b) composed a checkUst of mites taken in ecological studies at Dugway and in- dicated host relationships. Strandtmann and All- red (19.56) discussed mites of the genus Brevi- sterna, listed additional host and distribution records, and composed a key to the species. Howell, Allred and Beck (1957) studied mites foimd in woodrat nests and listed parasitic as well as free-living species. Allred and Marchette (19.57) studied the feeding habits of Brevisterna utahensis in Utah. Talley ( 19.57 ) studied the nasal mites of blackbirds. Allred and Roscoe ( 1957 ) studied p;irasitic mites in woodrat nests. Howell and Elzinga (1962) described a new species of fiu" mite and composed a key to the species of Radfordici. Elzinga (1960) listed mites of rodents from one specific area in Utah, and Ho (1962) listed mites from two areas. Ash (1963) studied the mites of deer mice in a chaparral community. Jenkins (1965) listed records of mites from squirrels taken in 12 counties. Other workers have published data on mites from Utah principally as incidental records asso- ciated with studies of other areas or fauna. These are listed in the species discussions that follow. MEDICAL AND ECONOMIC IMPORTANCE Dermamjssus americanus and D. guUiiuic have been involved with encephalomyelitis, but their exact relationship with the virus is not entirely clear (Baker et ah, 19.56). Transmission of trypanosomes and avian spirochaetosis has been demonstrated for D. gallinae ( Macfie and Thompson, 1929; Manwell and Johnson, 1931; Seddon, 1951). Rickettsialpo.x transmission has been related to D. sanguineus ( Huebner ct al.. 1946). Tlie implication of these species with disease suggests a similar potential for D. becki which also occurs in Utah. Ornithomjssus hacoti has been involved with murine typhus, rickettsialpo.x, tularemia, plague and nematode parasites (Baker et ah, 1956). Transmission of encephalomyelitis and New- castle disease has been related to O. si/lvkirum (Baker et al, 1956). Laelaps jettmari has been involved with hemorrhagic fever (Asanuma, 19.52), and L. •.\lthough the printed date on this publication is il Deienilier lOj-'i. i for purposes of priority this reference should be considered as I'l'i'i, 1 paper il is listed as 1955. kuchi has a suspected implication with tularemia (Baker, et ah, 19.56). Nothing is known of the potential of other species of Laelaps which oc- cur in Utah. Tularemia has been isolated from Eulaelaps stabularis and Haemolaehips mohrae (Baker, et al., 1956). Other species of Haemolaelaps, including the cosmopolitan H. glasgowi, have not as yet been implicated with disease trans- mission. Ornithosis virus of turkeys has been isolated from Haemolaelaps cusalis, Haemoga- masiis pontiger, and Ornithonijsstis sijlvianim (Eddie, ef a/., 1962). Hirstiomjssus Isahellinus has been implicated with tularemia in the laboratory (Francis and Lake, 1922). In addition to their disease transmission po- tential, mites occasionally are pests of man by their bites or invasion of his tissues. Derviamjssus Bhigham Young University Science Bulletin gallinae, D. sanguineus, O. bacoti, O. bursa, O. sylviarum and H. casalus have been known to attack man, and there is some circumstantial evidence to incriminate E. stabularis and H. pontiger. Still other parasitic mites may bite man on occasion. The high populations of rodents in the temperate desert areas of Utah, and the presence of many mites which have been implicated with disease transmission as shown above, create a potential reservoir of diseases in nature com- municable to man, which necessitates an in- creased knowledge of their natural history and other aspects of ecology. ACCOUNTS OF THE SPECIES In the discussion that follows, a brief state- ment is given about each family represented, usually followed by a listing of each species known to occur in Utah. However, several fami- Ues are listed for which mites were not identified beyond the family level. These were occasionally found on mammals but are not considered regu- lar consortes with them. Their inclusion is for convenience in separating them from mites which have more common occurrence and which for the most part are parasitic on their mam- malian hosts. Following the list of each species, comments on taxonomic or morphological variations are in- cluded when applicable. Under the heading "Distribution," published records for the United States are noted. In most cases only the earliest published record is given. Recently published hterature which duplicates previous records is not indicated. Where more than ten hosts are known for a state other than Utah, only a gen- eral statment on host relationships is given. Names for host records were taken verbatim from literature references. In Table 3 the equiva- lent names listed by Hall and Kelson ( 1959) are indicated in parentheses. For more specific lo- cality references, records for Utah are hsted by county. Following the heading "Other Utah Records" or "Utah Records," heretofore unpublished data are listed as represented by our surveys over the past sixteen years. For the most part the only records given are those which are new host or county records for Utah. In most instances where our collections duplicate previously listed rec- ords, we have not repeated them. Each of our records is represented bv detailed collection data in our files, but it is not practical to list such details at this time. Information on these data may be obtained by re(]uest. "Seasonal Occur- rence" is a summary of montlis over the total sixteen-year period. Under "Comments" when sufficient data are available, the geographic distribution in Utah (northerly or southerly) and occurrence in the Great Basin or Upper Colorado River Basin are indicated (Fig. 615). Apparent host preferences and other mite species associations are also listed. Keys to families, genera and species ;u-e in- serted where appropriate. Keys to tlie nymphs and males of some groups of Mesostigmata are lacking because of insufficient descriptions, or because these stages are unknown for many species. Key characteristics selected to separate the families, genera and species of mites known to occur on mammals in Utah are for the most part those which are m.ost easily seen. These may not be valid when dealing with species not in- cluded here. A properly cleared and mounted specimen is a necessary prerequisite to its iden- tification when deahng with the dorsal plate, chelicerae, legs, and specialized seta on the palpal tarsus. Such characters are best seen with a phase-contrast microscope. Where a specific feature Ls not visible or for the purpose of veri- fication, structures other than those listed in the keys may be used, and drawings of these for most species or a representative of each genus or family are appended. Setae usually are not shown except on the ventral plates of the meso- stigmatids and the scuta of the trombicidids. The keys to the males were composed using some specimens from areas other than Utah, and from literature descriptions. Relating the male or nymph to the female \vith which it Ls found is an aid to its identity, but discretion should be used. Where possible, verification should be made by comparing the specimen with one whose identity is known, and with the description of the type where such is adequate and available. In the descriptions of new species, measure- ments given are in microns. For purposes of structural orientation and to avoid duplication, figure references in the keys relate to Figures 1 to 5 on pages 5 to 7, and other illustrations be- ginning on page 71. Mites of Utah Mammals COXA I TRITOSTERNUM STERNAL PLATE STERNAL PORE 3rd STERNAL SETA PERITREME STIGMA GENITOVENTRAL PLATE COXA IV GENITOVENTRAL SETA METAPODAL PLATE ACCESSORY SETA ANAL PLATE ANUS ADANAL SETA POSTANAL SETA Fig. 1. Neopanisitidae, female ventral. KEY TO FAMILIES OF FEMALE AND MALE MITB5 AND L.AKVAL TROMBIC:ULLDS FOUND ON MAMMALS IN UTAH 1. With a lateral stigma situated dorsal to and at level between coxae 111 and IV ( Fig. 1 ) Without lateral stigma as above 2. With tritosternum (Fig. 1); ventral and dorsal plates well-developed (Fig. 1) Without tritosternum; dorsal and ventral plates much reduced or absent (Fig. 8) 3. Coxae arranged essentially in two parallel lines, situated on anterior half of body (Fig. 7); poorly sclerotized mites with few small setae Halarachnidae, page 9 Coxae somewhat radially arranged, occupying anterior three-fourths of body (Fig. 8); heavily sclerotized mites, densely covered with setae (from bats) Spinturnicidae, page 9 12 4 3 Bhicham Young University Science Bulletin EMPOOIUM PARASUBTERMiNALA- TIBIALA^ SUBTERMINALA SPUR MASTITARSALA CHELICERAL TEETH CHELICERAL BLADE GALEAL SETA CHELICERAL BASE TIBIAL CLAW CHELICERA Figs. 2-5. Chiggcr larva. 2, bocK dorsd; 3, left half gnathosoma %entral; 4, right chclicera dorsal; 5, body ventral. Mites of Utah Mammals PALPAL CLAW DORSOTIBIAL SETA LATEROTIBIAL SETA VENTROTIBIAL SETA GENUAL SETA TRICUSPID CAP GALEAL SETA CHELICERAL BLADE GALEA FEMORAL SETA CHELICERAL BASE Fig. 6. Chigger larva, right lialf gnathosoma dorsal. 4. Specialized seta on palpal tarus with three tines ( Fig. 407 ) Complex of Gamasolaelaptidae, Macrochelidae, Neoparasitidae, Pachylaelaptidae, Parasitidae, pages 8-9 Specialized seta with two tines 5. Anterior edge of sternal plate with circular structure and/or posteriorly e.xtend- ing tube, the genital opening ( Fig. 3.53 ) ; ventral plates fused into one holo- ventral plate (Fig. 355) or sometimes two plates ( sternogenital and ventro- anal, Figs. 362, 171, or stemo-genitoventral and anal, Figs. 351, 170) (males in part) Anterior edge of sternal plate without such genital opening; ventral plates sepa- rate as three distinct plates (Fig. 1) (females, in part) 10 6 6. Genitoventral plate truncate posteriorly, usually almost touching enlarged anal plate which is fre iif> San luan Co.: D. ordii, O. leucosaster. Utah Co.: P. Fits. 156, 262, 319 . i ^ ,,, r^ t, ■ i ^ " tuaniciilatus. Wayne Co.: r. mamcuUitus. Distribution. NEVADA: Dipodomys merriami, tu oo r i Dipodomys microps, Onychomys torridus, Peroguathus Seasonal occurrence. The 88 females were formosus, Perognathus hmgimemhris, Peromyscus crini- taken from May through September, mostly in tus, Peromyscus numiculutus, Peromyscus truei (Allred, June. 1962, 1963; Goates, 1963; Allred and Goates, 1964a). UTAH: Tooele Co.: Dipodomys ordii, Onychomys leuco- Comments. Mites of this species are distri- gaster, P. formosus, P. longimemhris, P. crinitus, P. buted over the state in 12 counties. maniculaUiS (Woodbury, 1956b). Utah Co.: ? host (El- j^ ^^^^^ ^j 33 eollection.s Klecmunia sp. was zinga, 1960 . ^u i -.. ,. i ^ ^ the only mite on its host. Other Utah records. Bo.\ Elder Co.: D. ordii. There is some question as to the parasitic Perogmithus parvtis Civrhon Co.: Eutamias minimus. ^^^^^^^ ^j: ^^^^^ ^^j^^^ Likely they are pre- Daeeett Co.: D. ordii. Duchesne Co.: CiteUus lateralvi. , 1.1 r .1 • Emery Co.: D. ordii, P. crinitus. Juab Co.: D. microps, daceous, but because of theu" common oc-cur- D. ordii, P. parvus, P. maniculatus. Kane Co.: D. orrfti, rence on rodents, they are included here. DERMANYSsmAE Kolcuati, 1859 This family represents a diverse group of mites parasitic on a variety of animals. They are worldwide in distribution and mav be commonly found on mammals. KE'^ TO GENERA OF FEMALE UEHMANVSSLDAE 1. Anal plate more than half as wide as body, concave on anterior margin (Fig. Ill) Mijom/.^stts, page 17 Anal plate narrow, conve.x or rarely truncate on anterior margin (Fig. 113) 2. Coxa III with one or more spurs or slight semi-circular protuberance on pos- teromedian side (Fig. 468) 3 Coxa III lacks spur or protuberance 4 3. Coxa III with well-formed spur (Fig. 468) Hir,sYion(/.s.s(/.v, page 17 Coxa III with rounded protuberance Ichoronyssus, page 31 4. Chelicera filamentous, very long; chela so minute as to be almost indiscernible (Fig. 389) Dcnnumis.ms, page 31 Chelicera not as above, with distinct and well-formed chela (Fig. 386) 5 5. With two dorsal plates (Figs. 32, 35) 6 With one dorsal plate (Fig. 40) 7 6. Posterior dorsal plate large, longer than anal plate (Fig. 35) Stccitoni/.yiii.'i, page 32 Posterior dorsal plate small, shorter than anal plate (Fig. 37) Ornithomissus, page 32 7. Anterior pair of sternal pores situated against lateral margins of plate resulting in lateral projections of plate for their accommodation (Fig. 248); third sternal setae on separate, small, distinct plates (Fig. 248) Ichoronyssus, page 31 Anterior pair of pores not against edge of plate which is without lateral projec- tions for their accommodation; third sternal setae, if not on plate, not on distinct separate plates (Fig. 257) Oruifliont/.'isus, page 32 Mites of Utah Mammals 17 MlJ0m/.s.sus montunus princcps (Furman and Tipton. 1955), Fiirman and Tipton, 1955 Other Utah records. Summit Co.; O. princeps. Figs. Ill, 219, 2.55, 564 Seasonal occurrence. Three females were Distribution. UTAH: San Juan Co.; Ochotomi taken in August. KEY TO SPECIES OF FEM.\LE HirstiuntJ.SSllS 1. Tarsus II has apical spurs (Fig. 606) 2 Tarsus II lacks spurs 13 2. Co.xa I has spiu- (Fig. 436); inner spurs of coxae II and III bifid (Figs. 457, 460) staff onii, page 18 Co.xa 1 lacks spur; other co.xal sptirs simple (Fig. 463) 3 3. Coxa II has two spurs (Fig. 450) 4 Co.xa II has three spurs (Fig. 442) 11 4. Coxa III has one spur (Fig. 461) /flr,s«/(.s, page 18 Co.xa III has two spurs (Fig. 463) 5 5. Coxa IV lacks spur . . 6 Coxa IV has spur (Fig. 514) 7 6. Inner spiu-s on coxae II and III rounded or truncate (Figs. 456, 462) . affmis, page 19 Inner spurs on coxae II and III shiuply pointed (Fig. 455) palustrix, page 19 7. Inner spur on coxa II broadly rounded (Fig. 454) punctatus. page 20 Inner spur on coxa II acutely or bluntly pointed (Figs. 451, 452) 8 8. Sternal plate seven times as wide as long; posterior border invaginated to level anterior to second sternal setae (Fig. 225) iiwaginatus. page 21 Sternal plate less than six times as wide as long; posterior border not invaginated beyond level of second sternal setae (Fig. 228) 9 9. Peritreme does not extend beyond anterior edge of coxa II (Fig. 555) eutamiae, page 21 Peritreme extends to middle of coxa I 10 10. Inner spur of coxa II short (not over 8 mm long), about as wide as long (Fig. 450) .. iitahcnsis, page 22 Inner spur of co.xa II long (about 18 mm long), much longer than wide (Fig. 449) . angustus, page 24 11. Femur II has blunt spur near anterior margin (Fig. 540) femuraUs. page 25 Femur II lacks spur 12 12. Inner spurs of coxae II and III broadly rounded (Fig. 447) longichidac, page 25 18 Bkic.ham Young Univehmty Science Bulletin Inner spurs of coxae II and III bluntly pointed (Fig. 446) //lOHiomi/.s, page 26 13. Coxa II has two spurs 14 Co.xa II has three spurs 17 14. Coxa IV has spur 15 Coxa IV lacks spur 16 15. Inner spiu" of coxa II represented only by slightly raised hump (Fig. 445) torus, page 27 Inner spur of coxa II distinct, pointed spur (Fig. 444) ncotomae, page 29 16. Genitoventral plate has two pairs of setae (Fig. 299) biscto.sii.s, page 29 Genitoventral plate has only usual pair of setae (Fig. 295) isahcUinus. page 29 17. Peritreme very wide for entire length, equal to almost half the thickness of legs (Fig. 569) triacanthtis, page 30 Peritreme of normal width or only slightly widened (Fig. .568) 18 18. Sternal plate less than four times as wide as long (Fig. 244) hilli variant, page 30 Sternal plate four or more times as wide as long 19 19. Peritreme ends at anterior edge of coxa II (Fig. 573); sternal plate about five times as wide as long (Fig. 242) hilli, page .30 Peritreme ends at middle of coxa I ( Fig. 575 ) ; sternal plate about eight times as wide as long (Fig. 245) incompttis, page 30 Hirstioni/sstis staff urdi Strandtmann and Hunt, 1951 Figs. 12, 122, 213, 282, 436, 457, 460, 479, 602 Distribution.GEORGI.^: Mephitis elonguta, Spilo- gale putorius; OKLAHOMA: Spilogale interrupta; TEXAS: Mephitis mesomelas, Spilogale leticoparia ( Strandtmann and Hunt, 1951 ) . Utah records. I'tali Co.: Spilogale gracilis. Seasonal occurrence. The 19 females were taken in July. Each of 12 contained an egg. Comments. This mite likely is more common and widely distributed than has been reported. An obvious reluctance on the part of scientists to collect and examine its common host likely accounts for the few records available. Hirstiontjssus tarsalis, new species Figs. 13, 123, 215, 270, 459, 461, 563, 603 Utah records. Box Elder Co.: Peromyscus muni- culatus, Lynn, two females, July, 1957. Daggett Co.: P. manicutatus. Deep Creek Camp Ground, one female, June, 1958. Type data. Holotype female, B.Y.I', collec- tion no. 445. Taken from Pcromijsctts maniculat- (w. Deep Creek Camp Croimd, Daggett Co., Utah, 23 June 1958. In the acarologv collection of the Department of Zoology and Entomology, Brigham Young University, Provo, Utah. The trivial name tarsalis refers to tarsus IV which possesses a long spur-like seta. Female Gnathosonia. Greatest width at base, 94; length to base of palpal trochanter, .58. All setae nude. Cheliceral digits moderate in length and thickness, .50 long from base of moveal)ie digit; lack teeth. Legs. Length from distal edge of co.xa to base of pretarsus: I, 218; II, 185; III, 197; IV, 271. Width of genua: I and II, .58; III, 43; IV, 41. Tarsus II has pair of apical, slightly curved, blunt spurs, 8 long. Tarsus IV has apical, spur- like seta, 22 long. Goxa I has two sube(jual setae, proximal one slightly larger. Coxa II has two MiTKs OK UiAii Mammals spurs: usual anterocaudal one of mcKlerate thick- ness and sharjilv pointed; inner spur bluntly pointed, 7 long and 7 wide. Coxa III has one inner posterior spur, sharj^ly pointed, 12 long and 7 wide. Coxa IV lacks spur. Dorsal plate. Elliptical; all sides convex; tapers to blunt point posteriorly; 480 long, great- est width 268. Has 26 pairs setae; most anterior pair 17 long, anteromedial setae 31 long, medial setae 12 long, posterior ones 21 long. Sternal plate. Five times as wide as long (width measured between outer edges of third sternal setae); anterior border only slightlv con- vex; posterior border concave, invaginated to level even with second sternal seta; lateral mar- gins moderately concave; anterior corners elon- gate and sharply pointed; posterior corners nar- rowly extend between coxae II and III. Has three pairs subecjual setae, first and third pairs about 29, middle pair about 36 long; distance between posterior pair of setae 2.7 times as great as distance between first pair; first pair of setae on anterior margin of plate, second pair set in from lateral margins, third pair on posterior edge of plate. Genitoventral plate. Width 79 (measured between outer edges of genitoventral setae); length 86 (from anterior edge of genitoventral seta to posterior border of plate); has prominent internal ridges; ha.s narrow, darkened border effect; thickly punctate; setae 24 long, situated slightlv posterior to mid-coxa IV. Anal plate. Elongate oval; evenly convex but more strongly posteriorly; with cribum. Length 65 ( from base of postanal seta to anterior edge of plate); greatest width .S5; has light border com- pletely encircling plate; lightly punctate. Adanal and postanal setae sube(jual, about 19 long; adanal setae situated opposite middle of anus. Urisclerotized part of venter. Has 13 pairs of setae posterior to genitoventral setae (not counting those on posterior borders ) ; setae about etjual, 24 long. Peritreme. Sinuous, ends at anterior fourth of coxa I; granulo-punctate for entire width and length; about equal width (5 wide) for entire length. Hirstionyssus affirm (Jameson), 1950 Figs. 14, 124, 221, 286, 456, 462, .5.56, 604 Distribution. CALIFORNIA: Eutamias minimus, Pfromijscus hoytii, Tamiasciurun dougUisii (Jame.siin, 1950b,' Strandtmann and Wharton, 1958). Utah records. Bo.x Elder Co.: E. iiiiiiimus. Kant- Co. : Pcrogiuithus formosus, Peromyscus tnici. Seasonal occurrence. Seven females and six protonymphs were taken in June and July. Comments. Little is known about the host relationsliips of this species. In two of the three collections, H. affinls was the only mite on its host. In one collection it was associated with Hirstioni/ssus iitohensis. Hirstioiiijssits pahistris, new species Figs. 15, 131, 222. 261, 455, 463, 558, 605 Utah records. Carbon Co.: Sorex pahistris, three miles south of Scofield, two females, July, 1960. Type data. Holotype female, B.Y.U. collec- tion no. 4808. Taken from Sorex pahistris, 3 mi south of Scofield, Carbon Co., Utali, 29 July 1960, by D Elden Beck and Clyde L. Pritchett. In the acarology collection of the Department of Zoology and Entomology, Brigham Young Uni- versity, Provo, Utah. The trivial name pahistris is Latin for "marshy," and refers to the habitat of the type host. Female Gnathosoma. Greatest width at base, 73; length to base of papal trochanter, 70. All setae nude. Cheliceral digits 47 long from base of moveable digit, moderate in thickness, lack teeth. Legs. Length from distal edge of coxa to base of pretarsus: I, 273; II, 225; III, 200; IV, 275. Width of genua: I and II, 53; III, 45; IV, 43. Tarsus II has apical spurs of medium size and slightly curved. Coxa 1 has two subequal setae, about 28 long. Coxa II has two spurs: usual anterocaudal spur sharply pointed, about 20 long; inner spur sharply pointed, 18 long and 11 wide. Coxa III has two spurs: outer posterior one sharply pointed, 10 long and 5 wide; inner spur sharply pointed, 18 long and 10 wide. Coxa IV lacks spur. Dorsal plate. Covers most of dorsum; sides parallel for two-thirds of length; tapers to blunt tip; 510 long; greatest width 290. Has approxi- mately 23 pairs of delicate subequal setae, 10 to 15 long, except for anteromedial and terminal ones which are slightly larger, 15 to 20 long. Sternal plate. Three and eight-tenths times as wide as long (width measured between outer edges of third sternal setae); anterior border slightly convex; posterior border concave, in- vaginated almost to level of second setae; lateral margins concave; anterior corners sharply pointed, extend between coxae I and II; pos- 20 Bhigham Young Univehsity Science Bulletin terior corners sharply pointed, extend between coxae II and III. Has three pairs of setae: an- terior pair 49 long, median pair 50, posterior pair 45; distance between posterior pair of setae slightly more than twice distance between anterior pair; first pair of setae on anterior edge of plate, second pair set in from sides, third pair situated on extreme posterior comers. Genitoventral plate. Width 88 (measured between outer edges of genitoventral setae); length 1.33 (from anterior edge of genitoventral seta to posterior border of plate); broadly rounded, almost truncate; has prominent inter- nal ridges; has distinct light border; setae 30 long, situated at level opposite middle of coxa IV. Anal plate. Sub-oval, sides broadly rounded, anterior edge almost truncate. Has narrow cribum, 13 long. Length 75 (from base of post- anal seta to anterior edge of plate); greatest width, 75; has distinct darkened border. Adanal and postanal setae subequal, 24 to 2.5 long; ad- anal setae situated at level opposite middle of aniLS. Unsclerotized part of venter. Has 15 pairs of setae between level of genitoventral seta and posterior end of anal plate, not counting peri- pheral setae; those closest to genitoventral plate .30 long, others 20 to 25 long." Peritreme. Sinuous, narrow, extends to middle of coxa I. Hirstionyssiis punctatus, new species Fif^s. 16, 12.5, 224, 2.58. 4.54. 467, 483, 562, 6(X) Utah records. Box Elder Co.: Eutaniitis mmimi/.v, Lynn, one female, July, 1957. Daggett Co.: Glauconu/s suhritius. Deep Creek Campground, .six female.s ( each of three with an egg). June, 19.58. Kane Co.: Peromysctis maniculatus, head of Cottonwood Creek, .six females, two deutonymphs, June, 19.58. Summit Co.: Eutamkis quuilrivittatus, Bald Mtn., 14 females. August. 1957. Type data. Holotype female, from Etttamias Cjuadrivittatuf!, Bald Mtn., Summitt Co., Utah, 25 .A.ugust 1956. In the acarology collection of the Department of Zoology and Entomology, Brigham Young University, Provo, Utah. The trivial name punctcitus refers to the punctation of the anal plate. Female Gnathosoma. Greatest width at base, 115; length to base of palpal trochanter, 77. All setae nude. Cheliceral digits moderate in length and thickness, 60 long from base of moveable digit; lack teeth. Legs. Length from distal edge of coxa to base of pretarsus: I, 370; II, 320; III, 285; IV, 406. Width of genua: I, 58; II, 62; III, 46; IV, 43. Tarsus II has pair of apical, slightly curved, blunt spurs, 9.6 long. Tarsus IV has apical, spur- like seta, 17 long. Coxa I has two subequal setae, pro.ximal one slightly longer. Co.xa II has two spurs: usual anterocaudal one long and sharply pointed; inner spur thumb-like, broadly rounded, almost truncate, 14 long, 12 wide. Coxa 111 has two spurs: outer posterior one sharply pointed, 12 long and 7 wide; inner posterior one blimtly pointed, 14 long and 12 wide. Co.xa IV has sharp- ly pointed spur, 13 long and 7 wide. Dorsal plate. Sides neither concave nor con- vex, but slightly sinuous; tapers to blunt point; 558 long; greatest width, 3.30. Has 26 pairs of setae; most anterior pair 16 long, anteromedial setae 31 long, medial setae 16 long, posterior ones 31 long. Sternal plate. Three and nine-tentlis times as wide as long ( width measured between outer edges of third sternal setae); ;mterior border convex; posterior border concave, invaginated to level almost to second setae; lateral margins moderately concave; anterior corners elongate but bluntly pointed; posterior comers narrowly extend along anterior margins of co,xae III. Has three pairs of subequal setae, third pair slightly longer than first and second; distance between posterior pair of setae 2.8 times as great as dis- tance between first pair; first pair of setae on anterior margin of plate, second pair set in con- siderably from lateral margins, third pair set in from posterior margin. Genitoventral plate. Width 98 (measured between outer edges of genitoventral setae ) ; length 127 (from anterior edge of genitoventral seta to posterior border of plate ) ; has conspicu- ous internal ridges; has narrow, darkened border effect; lightly punctate. Setae .34 long, situated at level opposite mid-coxa IV. Anal plate. Elongate oval; evenlv convex but more stronglv tapered posteriorly. \Vith cribum. Length 77 (from base of postanal seta to an- terior edge of plate); greatest width 72; has dark border completely encircling plate; lightly but thickly pimctate. Adanal and postanal setae sub- equal, about 31 long; adanal setae situated op- posite middle of anus. Unsclerotized part of venter. Has 15 pairs of setae posterior to genitoventral setae ( not coimt- ing those on posterior borders); setae about equal, 31 long. Mites of Utah Mamma Peritreme. Slightly sinuous; ends at anterior fourth of coxa II; granulo-punctate for entire width and length; 9.6 wide at base, gradually tapering to 4.8 at tip. Hirstionyssus invagmatiis, new species Figs. 17, 18, 127, 130, 225, 227, 267, 276, 452, 453. 464, 468, 484, 485, 560, .561, 607. 608 Hir.s-tionyssus occidentalis (Ewing) of Utah records (in part). Utah records. Cache Co.: Citellus ar/iiutus, Blacksmith Fork Junction, one female, June, 19.53. Rich Co.: C. tinnatus; Laketown. .seven females, June, 1953. San(x-te Co.: C. urnuitus, 11 miles east of Mt. Pleasant, one male, one female, August, 1951, Summit Co.: C. tirmutus. Lake Creek Summit, 19 females, August, 1953. Utah Co.: C. armatus, Provo Canyon. 12 females. |une, 1951 and 1957; five females, July, 1957; Mus musculus, Lelxi, two females, one deutonvmph, January, 1951; Tlwmomys tulpoides, Provo Canyon, one female. JuK, 1956. Wasatch Co.: C. armatus. Currant Creek, two males, five females, June. 1953; Soldier's Summit, two females. June, 1951; Strawberry Reservoir, nine females, June, 1951. Comments. Apparently this mite prefers the squirrel CiteUus armatus as its host. In five of 17 collections, H. invagimitits wa.s the only mite on its host. In five collections it was associated with other species of Hirstioni/.s- .si/.v. Type data. Holotype female, B.Y.L'. collec- tion no. 4117. Taken from CiteUus armatus 3 mi west of Big Tree Camp, Provo Can\on, Utah Co., Utah, 22 July 1956, by Dorald .M. Allred. In the acarology collection of the Department of Zoology and Entomology, Brigham Young Uni- versity, Provo, Utah. The trivial name invaginat- us refers to the extreme invagination of the pos- terior border of the sternal plate. , Female Gnathosoma. Greatest width at base, 92; length to base of palpal trochanter, 73. All setae nude. Cheliceral digits 48 from base of moveable digit, moderately developed, lack teeth. Legs. Length from distal edge of coxa to base of pretarsus: I, 345; II, 298; III, 275; IV, 375. \\'idth of genua: I and II, 55; III and IV, 43. Tarsus II has pair of medium-sized, slightlv curved apical spurs. Coxa I has two setae: distal one 22 long, proximal one 30. Coxa II has two spurs: usual anterocaudal spur sharplv pointed, 2.5 long; inner spur shaqilv pointed, 2.3 long and 20 wide. Coxa III has two spurs: outer posterior one sharplv pointed, 10 long and 6 wide; inner spur sharply pointed, 20 long and 10 wide. Coxa IV has posterocaudal spur, 12 long and 6 wide. Dorsal plate. Elliptical, covers most of dor- sum except sides and posterior corners; lateral margins parallel, straight or slightly concave, with_ distinct thickened border; tapers to blimt tip; 554 long; greatest width 304; has 25 pairs of setae, those on periphery largest; peripheral setae 20 to 25 long, terminal pair .30 long, medial setae 10 to 13 long. Sternal plate. Almost seven times as wide as long (width measured between outer edges of third sternal setae); anterior border sHghtlv con- vex; posterior border concave, invaginated to level slightly anterior to second pair of setae; lateral margins straight; anterior and posterior borders of plate slightly darkened; anterior cor- ners sharply pointed, extend between coxae I and II; posterior corners sharply pointed, extend laterally between coxae II and III; has three pairs of subequal setae, 37 to 40 long; distance between posterior pair of setae two and one- half times as great as distance between first pair; first pair of setae on anterior edge of plate; sec- ond pair set in from sides; third pair near edge of posterior corners. Genitoventral plate. Width 93 (measured be- tween outer edges of genitoventral setae); length 120 ( from anterior edge of genitoventral setae to posterior tip of plate ) ; slightly expanded behind genitoventral setae, tapers abruptly (al- most straight) to blunt, rounded tip; has distinct internal ridges and light border edged inwardly with darkened line; setae 25 long, situated at level about midwav opposite coxa IV. Anal plate. Suboval, sides and anterior edge broadly rounded; tapers to broadly rounded tip. Cribum 17 long. Length 68 (from base of post- anal seta to anterior edge of plate); greatest width 72; has distinct dark border. Adanal setae 28 long; postanal seta heavier, 30 long; adanal setae situated at level opposite middle of anus. Unsclerotized part of venter. Has 12 pairs of setae between level of genitoventral setae and posterior tip of anal jilate, not counting peri- pheral setae; setae subequal, 20 to 30 long. Peritreme. Sinuous, curved, wider at base, tapers graduallv to moderate width; extends to middle of coxa I. Hirstionyssus eufamiae, new species Figs. 19, 128. 228. 279, 451, 469, 482. 555, 609 Hirstionyssus occidentalis (Ewing) of Utah records ( in part ) . Utah records. Utah Co.: Eutamitis quadrivittatun, Rock Canyon, Provo. one female, October, 1956; Sco- 22 BlUGHAM VoUNC UnINKHMIV SciliNCK BvLLKl IN field Reservoir, eight females, June, 1957; Reithrodon- tomys megalotis, Cedar Vailev. one female. November, 1952. Type data. Holotype female, B.Y.U. collec- tion no. 4778. Taken from Eutamias sp.. Aspen Grove, Utah Co., Utah, 16 October 1956, by Lawrence Tseu. In tiie acarology collection of the Department of Zoology and Entomology, Brigham Yoimg University, Provo, Utah. Tlie tiivial name cutamiae refers to the generic name of the type host. Female Gnathosoma. Greatest widtli at base, 90; length to base of palpal trochanter, 72. All setae nude. Cheliceral digits 78 from base of move- able digit, strongly developed, lack teeth. Legs. Length from distal edge of coxa to base of pretarsus: I, 308; II and III, 264; IV, 352. Width of genua: I, 48; II, 50; III, 40; IV, 38. Tarsus II has small pair of apical claws. Co.xa I has two subequal setae, 22 long. Coxa II has two spurs: usual anterocaudal spur sharply pointed, about 22 long; inner spur sharply point- ed, 10 long and 8 wide. Co.va III has two spurs; outer posterior one sharply pointed, 10 long and 5 wide; inner spur sharply pointed, 12 long and 10 wide. Coxa IV has posteromarginal spur, 5 long and 5 wide, which may be branched or double in some specimens. Dorsal plate. Covers all of dorsum except posterior corners; sides parallel for two-thirds length; tapers to blunt tip; 524 long; greatest width 334; has distinct hght border; has about 26 pairs of delicate setae, those on periphery and posterior tip of plate largest: medial setae 12 to 18, peripheral setae 20 to 30. Sternal plate. Slightly more than three and one-half times as wide as long (width measured between outer edges of third sternal setae ) ; an- terior border convex; blunt projection tapers an- teriorly from first pair of setae; posterior border concave, invaginated to level almost etjual with .second pair of setae; lateral margins straight, with darkened borders; anterior corners shaqilv pointed, extend between coxae I and II; posterior corners sharply pointed, extend between coxae II and III. Has three pairs of subecjual setae, 3.5 to 38 long; distance between posterior pair two and one-half times as great as distance between first pair; first pair of setae on anterior edge of plate, second pair set in from edge, third pair slightly in from edges of posterior corners. Genitoventral plate. Width 95 (measured be- tween outer edges of genitoventral setae ) ; length 113 (from anterior edge of genitoventral setae to posterior tip of plate ) ; slightly expanded behind genitoventral setae; tapers abruptly to blunt tip; has prominent internal ridges; setae 25 long, situated at level near posterior border of coxae IV. Anal plate. Suboval, sides and anterior edge broadly rounded; tapers to blunt tip. Cribum 21 long. Length 73 (from base of postanal seta to anterior edge of plate); greatest width 67; has distinct dark border. Adanal and postanal setae subequal, 28 long; adanal setae situated at level opposite middle of anus. Unsclerotized part of venter. Has 1 1 pairs of setae between level of genitoventral setae and posterior end of anal plate, not counting peri- pheral setae; setae subequal, 25 to 27 long. Peritreme. Sinuous, narrow, extends to an- terior edge of co.xa II. H irstiom/ssus utahcn.ii.s, new species Figs. 20, 88. 129, 187, 229, 264, 334, 355, 450, 470, 486, 489, 495, 501, 516, 524, 557, 610 Hirstioni/ssus occidcntalis (Ewing) of Utah records (in part). Distribution. UTAH: Everv county except Cache, Davis, Grand, Juab, Millard, Rich, Salt La]ccki was the onlv mite on its liost. The mites reported bv Keegan (1953), Woodbun,' (1956]j), and Allred and Roscoe (19.57) as Dcrmani/.ssus sp. from N. lepida and P. crinitus likely are D. hecki. Dernuini/ssus gallinae (DeCeer), 1778 Figs. 29. 145, 247, 302 Distribution. UTAH: Tooele Co.: Neotoma lepida, Onychomys leucogaster, Peromyscus crinitus (Ho, 1962). Ut;i]i Co.: O. leucogaster ( Elzinga. 1960). Other Utah records. None. Comments. Mites of this species are usual parasites of birds, although mammals occasional- ly are attacked. Steatomjssus antrozoi Radoysky and Fumian, 1963 Figs. 32. 35, 150, 2.53, 312, 421, .578 Distribution. ALABAMA, ARIZONA, CALI- FORNIA, GEORGIA, ILLINOIS, NEW YORK, OKLA- HOMA, OREGON, SOUTH CAROLINA. TEXAS: Various bats (Radovsky and Furman. 1963). Utah records. San Juan Co. -.Corynorhinus rafin- esipie. Seasonal occurrence. Fiye females, each of three with an egg, were taken in Mav. KEY TO SPECIES OF FEM.\LE Omithoni/SSUS 1. With two dorsal plates (Fig. 37) ariilus. pagi' 33 With one dorsal plate (Fig. .54) 2. Sternal plate \\'ith three pairs of setae (Fig. 2.54) hacoti. page .3.3 Sternal plate with two pairs of setae (Fig. 2.57) st/Iciaruin. page .33 KEY TO spe;c:ies oe m.\le Ornitlnxiiissiis 1. Dorsal plate narrow , coNcrs less tliaii lialf of dorsal sinlace of body ( Fig. 86) hacoti. page 33 Dorsal plate co\frs lialf or more of dorsal surface of bod\' Mites ok Utah Mammai.> 33 2. Dorsal plate abruptly invaginated near postiTior tip .sylvkiitim Dorsal plate evenly tapered posteriorly from about midpoint ( Fig. 85 ) a rid us Ornithuniis.su.'i aridu.s Furman and Radovsky, 19f>3 Figs. 37, 85, 94, 152, 256, 314, 369, 401, 427, 539, .581 Distribution. CALIFORNIA, NEVADA: Am- inospermuphilus leucurus; UTAH: Washington Co.: A. leucurus (Furman and Radovsky, 1963). Other Utah records. None. Ornithomjssus hacoti (Hirst), 1913 Figs. 54, 86, 95, 146, 176, 254, 303, 346. 372, 587 Distribution. Soutiieastem United States: Do- mestic rats (Pratt and Good, 1954). ALABAMA: Didelphis sp., Peronnjscus nuttallii, Rattus norr^egicus, Sigmodon sp. (Hays ;md Guyton, 1958). MARYLAND: Microtus peniistjlvanicus, Peromyscus leucopus (Dnim- mond, 1957). OKLAHOMA: Peromyscus spp. (Elks, 1960). TEXAS: Didelphis sp., Liomys sp., Neotorrui floridutui, Mus musculus, Neotomu micropus, R. tior- vegicus, Sigmodon .sp. (Randolph and Eads, 1946; Eads, Menzies and Miles, 1952; Eads, Trevino and Campos. 1965). UTAH: Beaver Co.: Peromyscus maniculatus; Carbon Co.: P. irumiculatus; Daggett Co.: Peromyscus crinitus; Duchesne Co.: P. numicuhtus, Peromyscus truei; Emery Co.: P. maniculatus; Garfield Co.: P. manicuUitus; Iron Co.: Peromyscus eremicus ( Allred, 1957c). Juab Co.: Ncotoma lepida, P. mimicu- latus (AUred, 1957c; Howell, Allred and Beck, 1957). Kane Co.: P. maniculatus, P. truci; MiUard Co.: P. mtiniculatusi S;in Juan Co.: Peromyscus hoylii, P. maniculatus; Sanpete Co.: P. maniculatus (Allred, 1957c). Tooele Co.: Citellus townsendii (Ho, 1962); Eutamias minimus (Woodbury, 1956b); N. lepida (All- red and Roscoe. 1957); P. crinitus, P. maniculatus, P. truei (Woodbury, 1956b). Uintah Co.: P. maniculatus (Allred, 1957c); Utah Co.: C townsendii, P. manicu- latus (Elzinga, 1960); P. truei (Allred, 1957c); R. norvegicus (Myklebust, 1951). Washington Co.: P. eremicus, P. maniculatus (Allred, 1957c). Other Utah records. Beaver Co.: Perognathus sp. Daggett Co.: P. maniculatus. Duchesne Co.: E. minimus. Garfield Co.: P. truei. Juab Co.: Perognathus )>arvus, P. truei. Kane Co.: P. parvus. San Juan Co.: Dipodomys ordii, P. crinitus, P. truei. Seasonal occurrence. Seventeen males, 56 females and 115 protonymphs were taken. Males were found in March, June and August; females in May, June, August, September and November; protonymphs from F'ebruary through November, e.vcept in April and October. Comments. Mites of O. bacoti are distri- buted over the state, altliougli apparently they are more common in the southern parts in the Upper Colorado River Basin. They are known from 15 countless. This species apparentiv prefers P. truei as its host. Its fre(|uency of occurrence was greatest on P. truei and R. norvegicus. Population indices were three to four for Perorm/.^cus spp., seven for Rattus, and one for other hosts. In 24 of 56 collections, O. hacoti was the only mite on its host. Figs Ornithonyssus stjlviarurn (Canestrini and Fanzago), 1877 30, 31. 151, 257, 259, 304, 311, 386. 415, 577, 583 Some variations were noted in the Utah series. The Utah specimens have a small pair of penultimate setae on the dorsal plate contrary to Furman's and Radovsky s (1963) diagnosis of the genus. The third pair of sternal setae are off the plate more frequently than on. Tliey vary in position from the edge of the plate to a con- siderable distance from it. Distribution. ? locality, mostly birds; Eutamias sp., Mus sp. ( Stnuidtmann and Wharton, 1958). MARYLAND: Mus musculus (Dnimmond, 1957). TEXAS: Eptesicus fuscus, Myotis velifer (George ;ind Strandtmann, 1960). Utah records. Beaver Co.: Marmota flaviventris. Utah Co.: Sylvilagus nuttallii. Seasonal occurrence. One female was taken in April and five in June. Comments. Principally a parasite of birds, O. sijlviarum infrequently attacks other animals in its environs. L.M-;L.\i>TiDAb; Berlese, 1892 Most mites of this fainilv are par;isitic on both invertebrates and vertebrates. They are world- wide in distribution, and are commonlv found on mammals. IJmOllAM ^OUNO UnIVEHSITV SciKNC'K BULLETIN KEY TO GENERA OK FEMALE LAELVPTIDAE 1. Femur II with large thumb-like spur (Fig. 541) Aiidioliiclap. Femur II laeks spur 2. Genitoventral plate with more than one pair of setae ( Fig. 309 ) _ _ Laclups. page 35 Genitoventral plate with only usual pair of setae (Fig. 313) 3 3. Body circular; legs I and II subequal; setae of sternal plate and coxae thick, al- most .spine-like (Fig. 271) Eul)racliylaela})s, page 37 Body oval; leg I distinctly longer and thinner than II; setae of sternal plate and coxa moderately slender (Fig. 280) 4 4. Sternal plate about as long as wide, distance between first and third setae about same as between third pair (Fig. 277) . Hypoasjiis. page 39 Sternal plate wider than long, distance between first and third setae about half as great as between third pair (Fig. 280) Haemolaelaps, page 39 KEY TO GENEHA OF MALE LAELAPTIDAE 1. Dorsal setae expanded distally, blade-like (Fig. 424) Hijpouspis, page 39 Dorsal setae normal 2 2. Femur II with large spur (Fig. .541) Androluclaps Femur II lacks spur — 3 3. Leg I more slender and much longer than leg II Hacmolaclaps, page 39 Legs I and II subequal 4 4. Setae of holoventral plate thick, heavy, almost spine-like ( Fig. 368 ) Laclaps. page 35 Setae of holoventral plate of normal dimensions (Fig. 36.5) Euhrachijlaelap.s, page 37 Till (1963) considers Hacmohelaps as a synonym of Androlaehps. However, for the present we recognize these as separate genera. AndroUielups Icvicuhis Eads, 1951 1964a). TEXAS: C). kucogaster, Verognuthus hispidus, T- ICO n,n nnK KAi coc Sipmodoti Impidtis (Eads, 1951). UTAH: Box Elder Figs. 158, 260, 305, 541, 585 /-.^ r. ■ 1 ^ / ah i incuN -r i ° Co.: Peroiinjscus mamcuuitus ( Allred, 1958). looele Hljpoaspis leviculus of some authors. Co.: O. Icucogaster, Perogtmthus pcirvus, Pcromyscus Mites collected in this studv varv slightlv in crimtus, P. maniculatus (Keegiin, 19,5.'}). ? lociility; size and morphology from the description of' the '^'f''"'" '«"'™''* ris, P. mtiniculatus ( Elzinga. 1960). TEXAS: Liotnys sp., Sylvilagus floridanus ( Randolph and Eads, 1946; Eads, Trevino, and Campos, 1965). Other Utah records. Tooele Co.: Perognathus formosus. Seasonal occurrence: A single mite was taken in March. Comments. Elzinga (1960) listed L. dipo- domius from Utah County, but did not designate a host, although he indicated Listrophorus sp. from two hosts (see Distribution above). It is likelv that this species is more common in Utah than siLspected. The few specimens represented may be due to superficial collecting techniques. Eads, Trevino, and Campos (1965) reported that in Texas half of the mice ( Liomi/s Bhi(;ham Yf)UNC llNiMinsnv Science Bulletin sp. ) examined were infested, one so heavily that there were several mites on most hairs on its back. Jameson (1950) indicated that Listro- phorws sp. was the most numerous ectoparasite on shrews, as abundant as 5(K) per animal. Myobildae Megnin, 1877 Myobiids are worldwide in distribution, found clinging to the hairs of small m;mima!s. Their association with hair follicles may result in local dermatitis and secondary infection. Although usually not considered parasitic, Wharton and his associates demonstrated that myobiids do suck body fluids ( Strandtmann, personal correspondence). KEY TO GENEVA OF MYOBUDAE Tarsus II with single claw Mijohia Tarsus II with paired claws Radfordia Mifobia sp. Comments. It is unusual that more speci- Distribution. UTAH: Tooele Co.: Neotoma lepi- mens of fur mites are not represented in Utah da (Alfred and Roscoe, 1957). collections. Collection techniques may be at Other Utah records. None. fault. KEY TO SPECIES OF FEMALE Radfordia 1. Dorsolateral seta I hooked near tip (Fig. 595) buchai Dorsolateral seta I not hooked 2 2. Submedian seta I as long as submedian III (Fig. 10) leinnina Submedian seta I about one-half as long as submedian III (Fig. 9) suhuUner Radfordia hachai Howell and Elzinga, 1962 Figs. 99, 595 Distribution. UTAH: Tooele Co., Utah Co.: Dipodomt/s ordii (Howell and Elzinga, 1962). Other Utah records. None. Radfordia Icrniiina (Koch), 1841 Figs. 10, 596 Distribution. MARYLAND: Microtus penmt/l- vanicus, Pitymys piiietoruni ( Dnimmond. 1957). UTAH: Garfield Co.: Pcromi/sctis manicultitus (AUred. 1957d). Other Utah records. None. Radfordia ■std)ulii^cr Ewing, 1938 Fig. 9 Distribution. .MARYLAND: Pcroinyncu.'i leuco- pus (Dnimmond, 1957). UTAH: Tooele Co.: Pero. ini/.\cus numicuhitus (\Voodl)iir\', 19.56b). Utah Co.: P. maniciilatus (AUred, 1957d); Reithrodontomi/s megalotis (Elzinga and Rees. 1964). Other Utah records. None. Trombk:ulidae Ewing, 1944 Mites of this family in the larval stage are important parasites of vertebrates the world over. They are commonly found attached to the integument of mammals where they appear as a single engorged mite or as small red, yellow, orange, or white patches in the ears, around the vibrissae, genitalia, thighs and axillary regions. Mites of Utah Mammals 43 KEY TO GENERA OF LARVAL TROMBKULIDAE 1. Leg I with seven segments; coxa I with one seta 2 Leg I with six segments; coxa I with two setae 3 2. Two anteromedian scutal setae (Fig. 4.3.5) Bernia One or no anteromedian scutal seta (Figs. 381, 4.37) 6 3. Scutum with anteromedian projection (Fig. .373) 4 Scutum without anteromedian projection (Fig. 379) 5 4. Cliehceral blade witli dorsal and/ or ventral row of teeth ( Fig. 4 ) ; spiracles and tracheae pr&sent Odontacariis, page 44 Cheliceral blade without dorsal or ventral teeth; spiiacles and tracheae absent Leemccnhockia, page 45 5. Cheliceral blade with distinct row of teeth ( Fig. 4 ) ; spiracles and tracheae pres- ent WhoitonUi, page 4.5 Cheliceral blade with tricuspid cap ( Fig. 6 ) ; spiracles and tracheae absent Chatia, page 45 6. Anteromedian scutal seta present (Fig. .381) 7 Anteromedian scutal seta absent (Fig. 4.37) Gahrliepia, page 46 7. Sensilla flagelliform (Fig. 381) Trombiciila, page 46 Sensilla expanded (Fig. 409) 8 8. Legs II and III with six segments Cheladonta. page 50 Legs II and III with seven segments Euschoengastki, page 51 Bernia, new genus of the Apoloniiiiae are known principally from Eyes present. Chelicera has tricuspid cap. the southern hemisphere from reptiles and birds. Palpal claw trifurcate. Each tarsus has two ^"^^ ^^^ ^^ly known North American representa- 1 1 1 i-i A- ^ n^ iiiL,. tive is a species from a pelican in lexas. On the claws and a claw-like empodium. Co.xa 111 has ,■ " ■, , \ ■ i i .1 . c » „ I \u • „t ,f,„ basis of its only known liost, it seems reasonable tliree setae. Scutum has three pairs of setae ,-', „ ..,, ^ ,. pairs ot setae ,-', „ .'.,,„ ,. . -i ■ r . I- t 1 t 1 to tentatively place Bernia with the Irombicu- simiuu- in fonn: anteromedians, anterolaterals, ,. ' " and posterolaterals; anteromedians surrounded iiWP- by distinct sutm-e isolating the pair from rest of We realize the inadvisability of designating a scutum; no anterior median projection; sensilla new genus and species on the basis of only one flagellifomi, situated near posterior margin of .specimen, but we feel that it is distinctive enough scutum. to warrant separation from other known genera Genotype: Bernia nuirita, new species. ^" ^P^ This distinctive genus is named in honor of r, ■ .1 . ti • r D Bernia manta, new species the senior authors wife, Berna. '■ rr.! . , 1. 1 Fig- 435 I he paired, anterior median scutal setae re- late this genus to the subfamilv Apoloniiiiae Type data. Lar\al holotype, L'niversity of which includes the genera Womersia, Apolunia Utah Institute of Environmental Biological Re- and Sauracarella. However, some members of search specimen no. 136.5:OOX. Taken from a the subfamily Trombiculinae show considerable female Perognathus longimemhris, specimen no. variation, especially in the .\M setae. Members ER2865, five miles N of Wig Mountain, Tooele Brigham Young University Science Bulletin Co., Utah, in sand dunes associated with grasses, 24 June 19.53, by Dale Parker and John Smith. In the acarology collection of the Department of Zoology and Entomology, Brigham Young Uni- versity, Provo, Utah. The trivial name marita is Latin for "wife," referring to the source of the generic name. Larva Body. Almost round; small, width 204; length (including capitulum) 228; one pair eyes, ocular plate lacking. Cnathosoma. Cheliceral blade slightly curved, with prominent tricuspid cap; basal seg- ment of chelicera longer than wide. Punctae absent. Palpal setae: femoral seta heavily branched; genual seta branched; ventral tibial seta branched on left side, forked on right side; lateral and dorsal tibial setae nude. Tarsus has tarsala, two nude and three or four branched setae; palpotibial claw trifurcate. Galeal seta with four or five branches. Scutum. Roughly trapezoidal, three times as wide as long. Anterior margin concave, pos- terior margin convex, lateral margins slightly convex with posterior lateral corners somewhat pointed. Scutum impunctate. Sensillary bases situated on posterior edge of scutum, posterior to level of PL's; separated little wider than dis- tance from SB to PL. Sensilla with 10 to 14 branches on dLstal half, proximal half with short barbs. Scutal setae plumose; AM posterior to AL; AM equals AL but shorter than PL; AM surrounded by prominent ridge. Prominent ridge along posterior border posterior to and ending lateral to SB. Legs. Coxae punctate. Setae on legs as fol- lows: I — coxa: 1 branched; trochanter: 1 branched; basifemur: 1 branched; telofemur: 5 branched; genu: 2 genualae, 1 microgenuala, 4 branched; tibia: 2 tibialae, 1 microtibiala, 8 branched; tarsus: tarsala, microtarsala, subter- minala, parasubterminala, pretarsala, empodium, and about 14 branched. II — coxa: 1 branched; trochanter: 1 branched; basifemur: 2 branched; telofemm-: 4 branched: genu: 1 genuala, 4 branched; tibia: 2 tibialae, 8 branched; tarsus: tarsala, microtarsala, pretarsala, empodium, and about 14 branched. Ill — coxa: right side 4 branched, left side 3 branched; trochanter: 1 branched; basifemur: 2 branched; telofemur: 3 branched; genu: 1 genuala, 3 branched; tibia: 1 tibiala, 6 liranched; tarsus: 1 mastitarsala, em- podium, and about 11 branched. Body Setae. Single humeral seta on each shoulder; about 26 dorsal setae, the first post humeral row with 8 setae; two pairs sternal setae; about 10 ventral setae posterior to sternals. KEY TO SPECIES OF LARVAL Oclontucariis 1. Sensilla nude (Fig. 378) ... Sensilla branched (Fig. 376) Uiudalei 2. Two genulae I (Fig. 2) hirsutus, page 45 One genuala I niicheneri Odontacanis liiisclalci ( Brennan and Jones), 1954 Fig. 376 Distribution. CALIFORNIA; Citellus heccheiji, Dipodomys agilis, Perognathus ccilifornicus ( Brennan and Jones, 1954; Loomis and Bunnell, 1962). NEV- ADA: Dipodomijs merriami, Dipodonu/s microps, Nco- toma lepida, Omjchonnjs torridus, Perognathus for- mosus, Perognathus longimemhrUs, Perognathus parvus, Peromijscus matticulatus (Allred, 1962. 1963; Goates, 1963; Allred and Goates, 1964a, 1964b), UTAH: Beaver Co.: D. microps (Brennan and Beck, 1955); Dipodomys ordii, P. parvus (Brennan and Jones, 1954), Juab Co.: P. formosus ( Brenniin and Beck, 19.55); P. parvus ( Brenniui and Jones, 19.54). Tooele Co.: D. ordii (Woodburv. 1956b); P. parvus (Brennan and Jones, 1954); P .' maniculatus (Woodbury, 1956b). Utali Co.: P, parvus (Brennan and Jones, 1954), Washington Co.: N. lepida (Brennan and Beck, 19.55), Other Utah records. Utah Co.: D. ordii. Seasonal occurrence. Fourteen mites were taken in April, July, August, September and December. Comments. In three of the seven collections, O. linsdalci was the onlv mite on its host. Odontacanis niicheneri Greenberg, 1952 Fig. 375 Distribution. COLORADO: Neotoma cincrea (Greenberg, 1952), NEVADA: Lizards (Allred and Mites of Utah Mammals Beck, 1962). UTAH: Box Elder Co.: Neotomti lepiila. Sylvilagus sp.; Garfield Co.: N. lepidti, Pemgnathus parvus; Grand Co.: N. lepida; Piute Co.: iV. lepida; Tooele Co.: N. lepidti, N. cinerea; Washington Co.: N. lepida ( Brennan and Beck, 1955). Other Utah records, t^ane Co.: N. lepida. Wash- ington Co.: Onychoviys torridus. Seasonal occurrence. Fifty-four mites were taken, mo.stly in June, but also in July and August. Comments. Mites of O. micheneri were more common in the southern part of Utah, and are known from seven coimties. Tliis species apparently has a preference for wood rats {Neotoma spp. ). In ten of 13 collections, O. micheneri was the only mite on its host. Odontacarus hirsutus (Ewing), 1931 Fig. 378 Distribution. CALIFORNIA: Aplodontia rufa, Citellus hecchciji, CiteUus lateralis. Dipodomijs venus- ttis, Eutainias sp., Microtus califurnicus, Neotoma ftis- cipes, Perognathus californicus, Stjlvilagus sp., Thonw- mys hottae (Brennan and Jones, 19.54; Gould, 1956; Jameson and Brenniin, 1957). NEVADA: Bassariscus astutus (Allred and CKJate.s, 1964a). UTAH: Grand Co.: Neotoma sp. (Brennan and Beck. 19.55). Tooele Qi.: ? host (Woodbury-, 1956b). Other Utah records. Grand Co.: Neotoma lep- ida. Kiuie Co.: N. lepida. Tooele Co.: Neotoma cinerea. Utah Co.: Mus musculus, N. cinerea. Seasonal occurrence. Fortv-eight mites were taken in February, May and July. Comments. In the six collections, O. hirsutwi- was found alone twice, and with Chatia ochotona once. The variations of the palpal claw, galeal and laterotibial setae in this species indicates a close relationship with Odontacarus chiapanensis. Leeiiwenhoekia atnericana (Ewing), 1942 Fig. 373 Distribution. ALABAMA: Cotton mouse; CALI- FORNIA: Citellu.i heecheyi, Microtus californicus, Peromifscus maniculatus, Sorex pacificus, Sorex trow- hridgii, Sorex vagrans, Thomomys hottae ( Brennan and Jones, 19.54; Gould, 19.56). COLORADO: Neotoma cinerea, Neotoma mexicana; KANSAS: P. maniculatus, Reitlirodontomys megalotis ( LoomLs, 1956). NEVADA: Neotoma lepida (Alked and Goates, 1964b). ORE- GON: Western mole (Gould, 1956). UTAH: Utah Co.: P. maniculatus (Ash, 1963). Other Utah records. None. Whartonia pcrplexa (Brennan), 1947 Fig. .379 Distribution. ARIZONA: Bats ( Bradshaw and Ross. 1961). CALIFORNIA: Antrozous pacificus, Pipistrellus hc^perus (Gould, 1956). MONTANA: Eptesicus fuscus (Brennan, 1947). NEVADA: An- trozous pallidus (Alh-ed and Goates. 1964a). UTAH: Salt Lake Co.: A. pallidus; San Juan Co.: Mtjotis cali- fornicus (Brennan and Beck, 1955). Other Utah records. Millard Co.: Bat. Seasonal occurrence. The 30 mites were taken in August. KEY TO SPECIES OF LARV.'VL ChotiO Genuala III and empodium present (Fig. 2) ochotona Genuala III and empodium absent .setosa Chatia .seto.su Brennan, 1946 Fig. 384 Distribution. CALIFORNIA: Eutumias towit- scndii, Peromt/scus maniculatus (Gould. 1956). IDAHO: Citellus lateralis, P. maniculatus (Brennan, 1946b). MONTANA: C. lateralis, Neotoma cinerea, Ochotona princeps, P. maniculatus, Tamia.iciurus hud.sonicus (Brenn;in, 1946b). UTAH: Cache Co.: P. maniculatus ( Brenn;in and Beck, 1955). Utah Co.: P. maniculatus (Ash, 1963). WASHINGTON: ? host (Brennan ;ind Beck, 1955). Other Utah records. Utali Co.: O. princeps. Seasonal occurrence. Nine chiggers were taken in June and August. Comments. Cliiggers of C. .setum were taken most commonly from deer mice {Peromijscus nmniculatus) from two counties in Utah. This mite was associated with other cluggers in two collections, and was foimd alone twice. Chatia ochotona (Radford^ Fig. 382 1942 Shun.sennia ochotona (Radford) of authors (Trauh and Nadchatram, 1966). Distribution. CALIFORNIA: Clethrionomys cali- fornicus, Microtus longicaudus, Ochotona schl^tieeps, Peromyscus hoi/lii, Peroini/sciis maniculatus. Sorex trow- 46 Bhicham Vounc I'nivebsity Science Bulletin hridgii (Gould, 1956; Jameson ^uid Brennan, 1957). Galirliepia umericana Ewing 1942 IDAHO: ? host (Brennan and Betlc, 1955), MON- p. .3- TANA; Ochotona ,sp, (Gould, 1956), NEVADA: ? ^' host (Brennan and Beck, 1955), UTAH: Cache Co,: Walchiu americana of authors ( Brennan and p. manicuUitus (Brennan and Beck, 1955), Millard Co,: Jones 1959) P. manicuUitus (Allred, 1957d), Tooele Co.: Neotoma ^ cinerea (Brennan and Beck, 19,55), Utali Co,: P. mani- Distribution. CALIFORNIA: "Gray squirrel"; culatus (A.sh, 1963). FLORIDA: "Cotton mouse," (Gould, 1956), IOWA: Other Utah records. None. Pewnnjscus leucopus; KANSAS: Neotoma micropus, c i„ Aiiifon .. f- leucopus, Sciurus carolineiisis, Sciurus niPer, StiL Seasonal occurrence. A total ol 39 mites was , f ' ' ,,.pvi Imi^ p ., r \«.i i» . i.T vilaiius floridanus ( Loomis. 1956), MAKYLANU: r, taken from May through August, and m January ^^J^^^^ (Dnunmond. 1957), NEBRASKA: S, caro- and November. Hnensis, S. niger; OKLAHOMA: Neotoma floridana. Comments. In seven of 18 collections, C. P- leucopus (LoomLs, 1956), UTAH: Garfield Co,: ochotom was the only mite on its host. It was Eutamias umhrinus; WISCONSIN: ? host (Brennan associated with chiggers of other species in si.x ^"^^ Beck, 1955), collections. Other Utah records. None, KEY TO SPECIES OF LARVAL Tromhictila 1. Sensilla nude (Fig. 385) caUfornica, page 47 Sensilla branched (Fig. 387) 2 2. Palpal claw trifurcate (Fig. 6) 3 Palpal claw bifurcate; accessory prong inner and ventral belkini, page 47 3. One pair of himieral setae (Fig. 2) 4 Two pairs of humeral setae Iwplae, page 48 4. Palpal femoral, genual, laterotibial, and ventrotibial .setae nude; galeal setae branched; two genuala I 5 Without this combination of characters 7 5. Palpal dorsotibial setae branched (Fig. 6) mijotis, page 48 Palpal dorsotibial setae nude 6 6. Spur on tarsus I longer than on tarsus 11; palpal genual, laterotibial, and ventro- tibial setae occasionally forked potosina, page 48 Spur on tarsus 1 equal to one on tarsus II paiuiincmi.s. page 48 7. Mastitibiala 111 present (Fig. 2) - 8 Mastitibiala 111 absent 11 8. Two mastitarsalae III; mastifemorala present (Fig. 2) 9 Three mastitarsalae III; mastifemorala III absent subsipmta, page 48 9. Galeal seta nude (Fig. 6) .. harperi, page 49 Galeal seta branched 10 10. Palpal femoral and genual setae branched (Fig. 6) jcuefli. page 49 Palpal femoral and genual setae nude . harperi, page 49 Mites of Utah Mammals 47 11. Scutum pentagonal with acute posterior angle (Fig. 398); .sensillae branclied or t)arhed the entire length (Fig. 398); coxa III with two or more setae; with tliree genualae 1 12 Without this combination of characters 13 12. Coxa 111 with two setae saigenti, page 49 Co.xa 111 with five setae esoensis, page 49 13. Ma.stitarsala 111 present (Fig. 2) 15 Mastitarsala 111 absent 14 14. Sensillae heavily branched and rebranched (Fig. 393) tinivuii, page 49 Sensillae simply branched (Fig. 403) kardosi, page 49 15. Two mastitarsalae 111 (Fig. 2) 18 One mastitarsala 111 16 16. Coxa 111 with one seta (Fig. 5); piUpal dorsotibial seta branched (Fig. 6) bakeri, page 49 Coxa III with three or four setae; palpal dorsotibial seta nude 17 17. Dorsal formula begins 2-8-8; cheliceral bases punctate arenicola, page 50 Dorsal formula begins 2-6-6; cheliceral bases impunctate montancn.'iis, page .50 18. Palpal dorsotibial and laterotibial setae nude; about 100 dorsal setae; no distinct humerals doremi, page 50 Palpal dorsotibial and laterotibial setae branched; about 28 dorsal setae, one pair distinct humerals alhedi, page 50 A number of workers have proposed that none of the Utah species of cliiggers belong to the genus Trombicida, senstt stricto, as cmrently defined. However, for the purposes of oiu- study we have retained in Trombicida, sensu Into, those species which belong to the subfamily Trombi- culinae and possess a flagelliform sensilla. Trombicida californica Ewing, 1942 Comments. Mites of this species were found Fig. 385 only in five comities in northwestern Utah in Distribution. CALIFORNIA: Variety of rodents **" .^""f/** Basin . (Brennan and Wharton, 1950; Brennan' and Jones. I" three of its six collections, T. cahformca 1954; Gould, 1956; J;uiieson and Brennan, 19.57). was the only mite on its host. In one collection IDAHO: Citellus lateralis; MONTANA: SykiUigus it was associated with a chigger of another nuttallii, Tamiasciurus hudsonicus ( Brennan iind Whar- snecips ton, 1950). UTAH: Cache Co.: Peromyscus manicidu- '^ tus; Rich Co.: Clediriojwmys gapperi, Microttis longi- caudus (Brennaii ;uid Beck, 1955). Salt Lake Co.: P. Trombicida belkini Gould, 1950 maniculatus ( AUretl, 1957d). Sanpete Co.: Citellus tn- Pig 3^1 matus (Brennan and Beck, 1955). Distribution. ARIZONA: ? ho.st; CALIFOR- Other Utah records. Utah Co.: C. anmitus, Xu- NIA: Citellus heecheyi, Mtirinota flaviventris, Mus mus- pus princeps. eulus, Pcrognuttius ealifornicus, Peromyscus iruci, re\>- tile.s (Cknild, 1950, 1956; Brennan and Jones, 19.54). Seasonal occurrence. Ninety-four mites were NEVADA: Neotoma lepidu, Perogruithus longimemhris, taken in June and August. reptiles (AUred and Beck, 1962, 1964; AUred, 1963; 48 Bhigham Young Univehsity Science Bulletin AUred and Goates, 1964b). UTAH: Duchasne Co.: Citellus leucurus; Emery Co.: Crotaphijtus coUaris; Garfield Co.: C. coUaris; Grand Co.: Cnemidorphoms tigris, C. eollaris; Juab Co.: Gamhelia tcMizenii, P. truei (Brennan and Beck, 1955): Piluophis catenifcr (Gould, 1956); Uta stansburuina; Millard Co.: Scehpor- us graciosus; Sevier Co.: Citellus lateralis; Utah Co.: S. graciosus (Brennan and Beck, 1955). Other Utah records. Duchesne Co.: Dipodomijs ordii. Rich Co.: Clcthrionomtjs gapperi. Seasonal occurrence. Seven mites were taken in June, July and August. Comments. Mites of this species commonly are found on lizards. They occasionally infest rodents, but in such cases their population inde.x is usually low. This species is known from nine counties. In its five collections, 7'. belkini was the only mite on its host. Tromhicula hoplai Loomis, 19.54 Fig. .388 Distribution. CALIFORNIA: Pcrognuthus cali- jornicus, Peromyscus calijorniciis, Pcmmi/sciis niaiiicultit- us (Brennan iind Jones, 1954; Loomis, 19.56). COLO- RADO: Neotoma lepidu, Neotomii lucxicami, ( Loomis. 1954; Finley, 19.58). KANSAS: Aritrozous hunhcri, Cytiomys ludovicitinus, Neotoma micropus, Pewmyscu.s leucopiis, Sylvilagus jloridunus; NEW MEXICO: 'Neo- toma mexicaiui, Perogruithus fluvus; TE.XAS: Citellus tridecemlineattts (Loomis, 1954, 1956). UTAH: Jual) Co.: Peromyscus truei; San Juan Co.: Perogiuithus apache (Brennan and Be<;k, 1955). Tooele Co.: P. truei (Wood- bury, 1956b). Utah Co.: ? host ( Elzinga, 1960), N. lepidu (Ho, 1962). Other Utah records. Kane Co.: Perognathus jxir- vus. San Juan Co.: P. truei. Uintah Co.: Citellus leucur- us. Seasonal occurrence. The 14 mites were taken in June, August and September. Tromhicula nti/otis Ewing, 1929 Fig. 387 Distribution. ARIZONA: Bats (Bradshaw and Ross, 1961). ARKANSAS: Sylvilagus floridanus (Loo- mis, 1956). CALIFORNIA:' ? host (Brennan and Beck, 1955). IOWA: Microtus pinetorum, Peronii/s- cus leucopus; KANSAS: Elaphc ohsoletu, Neotoma micropus; MAINE: Myotic lucifugus; MISSOURI: Eptesicus fuscus; MONTANA: E. fuscus; NEBRAS- KA: P. leucopus, Sciuru? niger (Loomis. 1956). NEW MEXICO: Sigmodon sp.; NEW YORK: ? host (Bren- nan and Beck, 19.55). OKLAHOMA: Neotoma flori- dana; PENNSYLVANIA: £. fuscus (Loomis. 1956). UTAH: Daggett Co.: Peromyscus vmniculatus (Allred. 1957d). Tooele Co.: Bat (Woodbury, 1956b). Utah Co.: Myotis californicus (Brennan and Beck, 1955). VIRGINIA: £. fuscus; WEST VIRGINIA: E. fuscus, M. lucifugus (Yunker, 1958). Other Utah records. Beaver Co.: Myoti.t sp Box Elder Co.: Myotis sp. Rich Co.: P. maniculatus. Tooele Co.: Myotis sp. Seasonal occurrence. The 83 mites were taken in June and July, mostly in June. Comments. Mites of this species were taken commonly in the northern part of the state in the Great Basin. They are known from si.v coun- ties. This species occurs freejuently on bats, but also attaches regularly to rodents. In si.x of its seven collections, T. mi/otis was the only mite on its host. Tromhicula potosina Hoffman, 1950 Fig. 391 Distribution. COLORADO: Neotoma alhiguUi, Neotoma cinerea. Neotoma lepida, Neotoma mexicarui (Finley, 1958). UTAH: Beaver Co.: Perognathus par- vus; Iron Co.: Neotoma lepida; Juab Co.: N. lepidu; Millard Co.: N. lepida; San Juan Co.: Neotoma sp.: Sanpete Co.: Microtus longicaudus; Tooele Co.: A'. lepida; Washington Co.: N. lepida (Brennan and Beck, 19.55). Other Utah records. None. Tromhicula i)anamcnsi.s Ewing, 192.5 Fig. 390 Distribution. NEVADA: Neotoma lepida (All- red and Goates, 1964b). Utah records. Beaver Co.: Peromyscus manicula- tus. Kane Co.: N. lepida, P. maniculatus. Seasonal occurrence. The 22 mites were taken in June, August and September. Comments. In two of five collections, T. patuimcn.si.s was the only mite on its host. In one collection it was associated with Odonta- carus michencri. Tromhicula suh.nonata Brennan and Wharton, 195() Fig. 392 Distribution. CALIFORNIA: Citellus beldingi, Marmota flaviventrls (Gould, 19.56). COLORADO: Citellus lateralis (Brennan and Wharton, 1950). MIS- SOURI: Capclla gallinago ( Kardos, 19,54). MON- TANA: C. lateralis, M. flaviventer; NEW YORK: Mi- crotus pennsylvanicus, Zapus hudsonicus; NORTH DAKOTA: Sylvilagus floridanus; PENNSYLVANIA: M. pennsylvanicus, Sciurus motacilla ( Brenn;ui iuid Wharton. 19.50). WYOMING: Zapus princeps (Kar- dos, 1954). Utah records. Juab Co.: Reitlirodontomys mcga- lotK. Seasonal occurrence. A single chigger was taken in June. Mites of Utah Mammau 49 Trombicula harperi Evving, 1928 Fig. 394 Gould ( 19.56 ) indicated tliat tlie variations in Trombicula harperi and Trombicula microti are great. He di.scus.sed them a,s representatives of a single polymorpliic species, T. Jiarperi hav- ing priority. Our collection.s agree with Gould's descriptions, especiallv in the variability of the galeal and humeral setae. Consecjuently, records of T. microti are included here. Distribution. ARIZONA, CALIFORNIA, COLO- RADO, IDAHO, MAINE, MICHIGAN, MONTANA, NEW MEXICO, NEW YORK, OREGON, PENNSYL- VANIA, VERMONT, WASHINGTON, WYOMING: Variety of rodents ( Brennan and Wharton, 1950; Kar- dos. 1954; Brennan and Beck, 1955; Gould. 19.56; Jameson and Brennan, 1957; Finley, 1958). UTAH: Beaver Co.: Ochotona prmceps; Duchesne Co.: O. priu- ceps; Iron Co.: O. princeps (Brennan and Beck, 1955); Peromyscus maniculatus (AUred, 1957d). Juab Co.: Camhelia sp., Neotoma lepida: Sanpete Co.: Citellus armatus, Clethrionomys gapperi, Marmota flaviventer, Microtus longicaudus (Brennan and Beck, 1955); P. maniculattis (Alhed, 1957d); Zapus princeps; Sevier Co.: Microtus sp. (Brennan and Beck, 1955). Summit Co.: O. princeps ( K;u-dos, 1954), Tooele Co.: Melospiza lincolni (Brennan and Beck, 1955). Utah Co.: Marmota flaviventrii (AUred, 1961); O. princeps; Wasatch Co.: O. princeps, Z. princeps; Wayne Co.: O. princeps (Bren- nan and Beck. 1955). Other Utah records. Beaver Co. -.Microtus longi- caudus. Carbon Co.: Citellus lateralis. Daggett Co.: P. manicuhittis. Iron Co.: C. laterals. S;ilt Lake Co.: C. gapperi. San Juan Co.: Onychomijs leucogaster. Summit Co.: Eutamias minimus. Uintah Co.: Homo sapiens, M. longicaudus. Utah Co.: Microtus monianus, O. princeps, Perognathus parvus, Z. princeps. Washington Co.: Mi- crotus montanus. Seasonal occurrence. A total of 2,242 chig- gers was taken. Mites were most abundant in August, although small numbers were found in March, May, June, Julv, September and Novem- ber. Comments. Mites of T. harperi occur gen- erally over the state. Tliey are known from 17 counties. This species apparently prefers the pika, Ochotona princeps, as its host, although it has been commonly found on jumping mice and voles. Its population index was 107 on pikas, 6 on jmnping mice, 11 to 126 on voles, and from 1 to 59 on other rodents. In 44 of (58 collections, T. harperi was the only mite on its host. In 13 collections it was associated with chiggers of other species. Trombicula jcuctti Brennan and Wharton, 19.50 Fig. 395 Distribution. CALIFORNIA: Variety of rodents ( Brennan and Wharton, 1950; Brennan and Jones, 19.54; Jameson and Brenn;m, 19.57). OREGON: Micro- tus townsetuiii (Brennan and Wharton, 1950). UTAH: Duchesne Co. : Peromyscus maniculatus (AUred, 1957d). Other Utah records. None. Trombicula sargenti Brennan, 1952 Fig. 398 Distribution. UTAH: Juab Co.: Neotoma lepida (Brennan, 1952; Brennan and Beck, 1955). Utah Co.: N. lepida (AUred iind Beck, 1953a). Other Utah records. None. Trombicula esoensis Sasa and Ogata, 19.53 Fig. 397 The Utah record below was described as a new species, T. reesi, by Alfred (1957d). Bren- nan and Jones (19.59) placed this into synonymy with T. esoensis. Distribution. MICHIGAN, MONTANA: ? host (Brennan and Jone,s, 1959). UTAH: Salt Lake Co.: Peromyscus maniculatus (AUred, 1957d). Other Utah records. None. Trombicula univari Brennan, 1965 Fig. 393 Distribution. ARIZONA: Antrozous pallidus, Pi- pistrellus hesperus, Plecotus townsendii (Brennan, 1965). Utah records. San Juan Co.: P. hesperus. Seasonal occurrence. Two mites were taken in September. Trombicula kardosi Loomis, 1954 Fig. 403 Distribution. KANSAS: Elaphe obsoleta, Sciurtts niger; UTAH: Garfield Co.: Eutamias umbrinus (Loo- mis, 1954). Other Utah records. None. Trombicula bakeri Ewing, 1946 Fig. 399 Distribution, Apparently not known heretofore from the United States. Bhigham Young I'nivilbsitv Scienc:e Bulletin Utah records. Kane Co.: Peromyscui manicuLtii.'i. Utah Co.: Citellus amiatus, Dipodomi/s ordii. Seasonal occurrence. Only three mites were taken — one each in June, July and August. Tromhiculu arenicola LoomLs, 1954 Fig. 406 Distribution. COLORADO: Dipodomys ordii, Neotonui albigula; KANSAS: D. ordii, Muscivora for- ficata, Perognathus hispidus ( Looini,s, 19.54). NEV- ADA: Dipodomys merriami, Dipodomys microps, Onychomys torridus, Perognathus formosus, Perognathus longimembris, reptiles, Sorex ienellus, Spermophilus tereticaudus (Allred, 1962. 1963; Goates. 1963; Allred and Beck, 1964; Allred and Goates, 1964a). NEW MEXICO: D. ordii, Perognathus jlavus (LoomLs. 19.54). UTAH: Box Elder Co.: Dipodomys sp., Ncotoma lepida, Perognathus sp.; Daggett Co.: Pcrogtuithus parvus; Gar- field Co.: CynoTtujs parvidens; Grand Co.: Perognathus sp.; Juab Co.: D. microps, P. formosus; MilUird Co.: D. microps; Sanpete Co.: Peromyscus nuiniculatus; Sevier Co.: D. ordii ( Brennan and Beck, 1955). Tooele Co.: D. microps, D. ordii ( Loomis, 1954); P. formosus (Bren- nan and Beck, 1955); P. longimembris (Woodburv, 1956b); P. parvus (Loomis, 1954); P. maniculatus (Woodbury, 1956b); sn;ike,s; Uintah Co.: D. or(/ii (Brennan and Beck, 1955). Utah Co.: Citellus leucurus (Ho, 1962); D. ordii, P. parvus (Brennan and Beck. 1955); P. maniculatus, Reithrodontomi/s megalotis ( El- zinga ;md Rees. 1964). Washington Co.: P. formosus, P. longimembris; Wayne Co.: D. ordii (Brennan and Beck, 1955. Other Utah records. Beaver Co.: Perognathus sp. Box Elder Co.: Perognathus sp. Grand Co.: Pero- gnathus sp. Juab Co.: P. parvus. Kane Co.: Peromyscus truei. San Juan Co.: Citellus spilosoma, D. ordii. Wash- ington Co.: Dipodontys merriami. Seasonal occurrence. The 653 mites were taken from February through October, e.xcept in March. Most were found in August, with about half as many taken in Julv and October. Few were taken during the other months. Comments. Mites of T. arenicola are com- mon in the southern part of Utah, and about equally distributed in the Great and Upper Colorado River basins. Thev are known from 16 counties. This species apparently prefers pocket mice of the genus Perognathus as its hosts, although mites were taken frequently on kangaroo rats, particularly D. microps. Tlie population index was 11 to 14 on kangaroo rats, 5 to 21 on pocket mice, and 1 to 4 on other animals. In 53 of 68 collections, T. arenicola was the only mite on its host. It was associated with chig- gers of other species in seven collections. Trombicula nioiitancnsis Brennan, 1946 Fig. 402 Distribution. CALIFORNIA: CHettus beldingi, Citellus lateralis, Dipodomys heermani (Gould, 1956). COLORADO: ? host (Brennan and Beck, 1955; Loo- mis, 1956). KANSAS: Variety of rodents, birds, mid reptiles (Loomis, 19.56). .MONTANA: Cynomys ludo- vicitmus, Sylvilagus nuttallii (Brennan. 1946a). NE- BRASKA: ? host (Brennan and Beck. 1955; LoomLs, 1956). OKLAHOMA: Heterodon playtrhinos, Tadarida mexicana (Loomis. 1956). TEXAS: C. ludovicianus (Eads, Menzies, and Miles, 1952). UTAH: Duchesne Co.: Cynomys leucurus (Brennan imd Beck, 1955). Kane Co.: Peromyscus truei; Rich Co.: Peromyscus man- iculatus; Sanpete Co.: P. manictdatus (Allred, 1957d). Tooele Co.: Peromyscus crinitus (Woodbury, 1956b). Other Utah records. Box Elder Co.: C. lateralis. Seasonal occurrence. The single mite was taken in June. Trombicula doremi Brennan and Beck, 1955 Fig. 404 Distribution. UTAH: Kane Q).: Perognathus longimembris (Brennan and Beck, 1955). Utah Co.: ? host (Elzinga, 1960), Dipodomys ordii (Ho, 1962). Washington Co.: Dipodomi/s merriami (Brennan and Beck. 19.55). Other Utah records. Beaver Co.: Pcrogtuithus sp. Seasonal occurrence. The single mite was taken in August. Comments. According to LtK)mis (personal coiTespondence ) this specimen is aberrant, having duplicated PL setae on both sides. It agrees generally with the type description and compares favorably with a paratyjje. Trombicula allrecli Brennan and Beck, 1955 Fig. 405 Distribution. CALIFORNIA: Neotoma sp., Sig- modon hi-^pidus ( Cknild, 1956). NEVADA: Dipodomys merriami, Dipodomys microps, Neotoma lepida (Goates. 1963; AUred and Goates, 1964b). TEXAS: LUunys sp. (Eads, Trevino, and Campos, 1965). UTAH: Washing- ton Co.: N. lepida (Brennan and Beck, 1955). Other Utah records. None. Cheladonta crossi Lipovsky, Crossley and Loomis, 19.5.5 Distribution. UTAH: Cache Co.: Peromyscus maniculatus (Lipovsky, Crossley and LoomLs, 1955). Other Utah records. None. Mites of Utah Mammals 51 KEY TO SPECIES OF LARVAL Eiischocngastiu 1. Tihiala III present (Fig. 2) 2 Tihiala III absent 9 2. Subterminala and parasubterminala present (Fig. 2) 5 Siibterminala and parasubterminala absent 3 3. Genualae II and III ab.sent lacerta. page 52 Genualae II and 111 present — 4 4. One pair of humeral setae (Fig. 2) Itoffniaunac. page 52 Two pairs of humeral setae fiirmani. page 52 5. Genualae II and III present . . 6 Genualae II and III absent utahemis, page 52 6. Sternal setae 2:2 (Fig. 5); palpal claw five-pronged 7 Sternal setae 2:4 or 2:6; palpal claw trifurcate (Fig. 6) 8 7. Palpal laterotibial seta nude (Fig. 6) soiiciniis, page 52 Palpal laterotibial seta branched orcgonensis, page 53 8. Sternal setae 2:4 cordiiemus, page 53 Sternal setae 2:6 ; cynomyicola. page 53 9. Two genualae I (Fig. 2) .- 11 One genuala I 10 10. Palpal claw trifurcate (Fig. 6); laterotibial seta nude or with 1 or 2 branches (Fig. 6) kind, page 53 Palpal claw five-pronged; laterotibial seta branched decipiens. page 53 11. Genuala III present ^^ 13 Genuala III absent 12 12. Palpal claw trifurcate (Fig. 6); galeal seta nude or forked (Fig. 6) ^ luteodcma, page 54 Palpal claw five-pronged; galeal seta branched (Fig. 4) ... pomerontzi, page 54 13. Subtenninaia and parasubterminala present (Fig. 2) 14 Subterminala and parasubterminala absent fassola. page 54 14. One pair of humeral setae (Fig. 2) 15 Two pairs of humeral setae rotunda, page 54 15. Palpal claw trifurcate (Fig. 6) 16 Palpal claw with four to seven prongs 19 52 BiiiGHANt ^OiNc; Univekmtv Science Bulletin 16. Scutum and cheliceral bases punctate (Fig. 408) 17 Scutum and cheliceral bases impunctate (Fig. 417) 18 17. Sensilla capitate; AL etjual to PL (Fig. 408) radfordi, page .54 Sensilla pyriform; AL shorter than PL (Fig. 413) ohesa, page 55 18. Dorsal setae of two form.s — majority narrow lanceolate, rest of usual form lanccolata, page 55 Dorsal setae of usual form criceticola, page 5.5 19. Sensilla capitate; AL equal to or greater than PL (Fig. 408) . ladfonli. page .54 Sensilla cordate; AL shorter than PL; scutum and cheliceral bases impunctate (Fig. 422) sciiiricola, page 55 Euschoengustia laceiia Brennan, 1948 Fig. 430 Distribution. CALIFORNIA: Citellus heechetji, Neotoma fuacipes, Perogtuithus californicus, Peromyscus boylii, Peromyscus maniculatus, Reithrodontomys sp., Sceloporus occidentalis, Syhilagus audulxmi ( Brennan, 1948; Brennan ;ind Jones, 1954; Gould, 1956; Jiuneson and Brennan, 1957). COLORADO: Neotoma albigulu, Neotoma cinerea, Neotoma mexicana; KANSAS: Cyno- mys ludovicianus, Neotoma micropus, Sylvilagus florid- anus ( Loomi.s, 1956). NEVADA: Dipodomijs microps (Goates, 1963). TEXAS: Liomys sp. (Eads, Trevino and Campos, 1965). UTAH: Garfield Co.: Neotoma lepida; Tooele Co.: N. lepida (Brennan and Beck, 1955). Other Utah records. Duch&sne Co.: Citellus lateralis. Seasonal occurrence. Five larvae were col- lected in Julv and August. Comments. Chiggers of E. lacerta are known from both the Great and Upper Colorado River basins. They have been found in onlv three counties. This species has been taken from a variety of hosts, but more commonly from Neotoma spp. than from others. Euschoengustia Itoffiiiannue Could, 19.56 Fig. 433 Distribution. CALIFORNIA: Citellus lateralis (Gould, 19.56). COLORADO: Neotoma mexieana (Finley, 1958), UTAH: Sevier Co,: C. lateralis (Bren- nan and Beck, 1955). Other Utah records. Beaver Co.: C. lateralis. Sanpete Co.: C. lateralis. Summit Co.: Oehotoiui prin- ce ps. Seasonal occurrence. A total of 105 larval E. hoffmanriae was collected in June and August. Greatest numbers were found in June. Comments. Specimens were found most commonly in southwestern Utah in the Great Basin. This species is known from only four counties. Most collections and greatest numbers of E. hoffmanae were taken from the squirrel, C lateralis. In tliree of its five collections, E. hoffmannae was associated with chiggers of other species. In one collection it was the only mite on its host. Eu.^c)ioengastia fiirmani Gould, 19.56 Fig. 432 Distribution. CALIFORNIA: Citellus leueurus, Neotoma lepida, Perognathus californicus, Reithrodon- tomys megalotis (Gould, 1956). UTAH: Washington Co.: N. lepida (Brennan and Beck, 1955). Other Utah records. None. Euschoengastia utahensis Brennan cUid Beck, 19.55 Fig 426 Distribution. NEVADA: Neotoma lepida (AUred and Goates, 1964b). UTAH; Juab Co.: N. lejnda (Brennan and Beck. 19.55). Other Utah records. Wasliington Co.: Pero- gnathus formosus. Seasonal occurrence. A single chigger was collected in April. Euschoengastia soricintis Gould, 19.56 Fig. 425 Distribution. CALIFORNIA: Sorex palustris, Sorex trowbridgii ((knild, 19.56). Mites of Utah Mammals 53 Utah records. One chigger was taken from Ocho- toiui pnnccps in llt;ih County in August. C^oniments. Looinis (personal correspond- ence) indicated that the Utah specimen is simi- lar to .wricimis from California, hut the sensilla is a different shape. Tliis mav represent an undeserihed species, but is here tentatively rele- gated to E. soricinus. Etiichocngustia orco^onensis (Ewing), 1929 Kig. 428 Distribution. CALIFORNIA: CitL'Uui, heldiugi. Citellus lateralis, Eutamias townsendii, Microtiis califor- nicus (CIoulcl, 1956); Ncurotrichus sp. (Jameson and Brenniui. 1957 ) ; Sorex pacificus, Sorex palustrui, Sorex towhridgii, Tumuisciurus douglasii; MONTANA: PiJca; OREGON: Mole (Gould. 1956). UTAH: Cache Co.: Peromt/scus maniciilutus; Iron Co.: Ocliotonu princeps, Sorex ohscurus; Sevier Co.; Microtus sp.; Ut;ih Co.: O. princeps; Wayne Co.: O. princips; WASHINGTON: ? host (Brennan and Beck, 1955). Other Utah records. Beaver Co.: Microtus longi- caudtis. O. princeps. Bo.\ Elder Co.: M. longicattdtis, Perogtuithus parvus. Duchesne Co.: P. numieulatus. Se- vier Co.: Eutamias quadrivittatus. Seasonal occurrence. A total of 226 larval E. orpgonen,si.$ was collected in June, July, August and November. Greatest numbers were taken in July. Comments. Chiggers of this species are com- mon in the Great Basin. Tliev are known from eight counties. This species is most closelv associated with the pika, O. princeps, and other hosts such as slirews and voles living in a similar habitat. In seven of 14 collections E. oregonen.?i.? was the only mite on its host. In five collections it was associated with chiggers of other species. Euschoengastia cordiremus Brennan, 1948 Fig. 429 Distribution. CALIFORNIA: Citellus heldingi, Marmota sp. (Gould, 1956). MONTANA: Citellus lateralis, Peromijscus maniculatus (Brennan, 1948) NEN'ADA: ? liost, P. maniculatus (Brennan ;uid Beck, 1955; AUred and Goates, 1964a). UTAH: Cache Co.: P. maniculatus (Brennan and Beck, 1955). Utah Co.: P. maniculatus (Ash, 1963). Other Utah records. Box Elder Co.: C. lateralis, Perogtuithus parvus. Seasonal occurrence. Si.xteen larval E. cor- diremus were taken in June, July and October. Comments. Mites of this sj>ecies are known only from three counties in northwestern Utah in the Great Basin. In its tliree collections, E. cordiremus was associated each time with mites of a different species of Euschoengastia and with other meso- stigmatids. Euschoengastia cijnomt/icola Crossley and Lipovsky, 19.54 Fig. 423 Distribution. NEBRASKA: Ct/nomi/s ludovician. us; KANSAS: Citellus tridecemlineatus. C. ludovician- us, Perognathus hispidus (Crossley and Lipovsky, 1954). Utah records. ^a>i Juan Co.; Citellus spilosoT7ui. Seasonal occurrence. Nine mites were col- lected in May. Euschoengastia land Brennan and Beck, 195.5 Fig. 419 Distribution. NEVADA: Peronn/scus maniculatus (Allred and Goates, 1964a). UTAH:' Box Elder Co.: P. maniculatus, Reithrodontomt/s megalotis; Uintah Co.: P. maniculatus (Brennan and Beck, 1955). Other Utah records. Utah Co.: Ochotona prin- ceps. Seasonal occurrence. Two specimens were taken in July. Comments. This species has been taken from both the Great and Upper Colorado River basins. Euschoengastia decipiens Gould, 1956 Fig. 431 This species is very similar to Euschoengastia radfordi and may prove to be synonymous with it. The two are differentiated by the presence of two genualae I on radfordi and one genuala I on decipiens. In a series of chiggers taken from Leptis californiciis in southern Utah, four mites possess two genualae I and two have one genuala I. Distribution. CALIFORNIA: Perognathus par- vus (Gould, 1956). NEVADA: ? host, Dipodomys merriami, Dipodomys microps, Perognathus formo.ius, Perogtuithus longimemhris, P. parvus ( Brennan and Beck, 1955; Allred, 1963; Goates, 1963; Allred and Goates. 1964a). OREGON: ? host (Brennan and Beck, 1955). UTAH: Box Elder Co.: Lcpus calif orni- cus; Cache Co.: Peronn/scus tnaniculatus; Duchesne Co.: Citellus lateralis; Garfield Co.: C. lateralis; Tooele Co.: Neotowa lepida; Utah Co.: Dipodomys ordii, P. parvus; Washington Co.; D. merriami, P. formosus, P. longimem- hris (Brennan and Beck, 1955). 54 3bigham Young Univehsitv Science Bulletin Other Utah records. Box Ekltr Co.: P. numku- liitus. Sijlvilagus idahoensh. Daggett Co.: Pcrornyscus truei. Juab Co.: L. calijornicus, Sijlvilagus auiluhonii. Kane Co.: D. ordii, N. lepida, P. parvus. S;in JuiUi Co.: P. nianicuUitus. Sanpete Co.: C. lateralis, Erethizon dor- mtiim. Utah Co.: D. microps, P. maniculatus, Sylviligus mitUillii. \Va.shington Co.: L. calif amicus, Ferojmjsctts crinitus, Perorni/scus ereiiiicus, S. audulionii. Seasonal occurrence. A total of 1,129 larval £. decipiens was collected. Mites were found during February through June, and in August and November. Greatest numbers were taken in February, March and April. Comments. Mites of E. decipiens are known from the Great and Upper Colorado Riv^r Basins in 12 counties in Utah. ThLs species has been taken from a variety of rodents, but the majority of the collections were as follows: 28% from Dipodomtjs spp., '25% from rabbits, 242 from Pcrognathus spp., and 11% from Pcromi/scus spp. Twenty percent of the collections were from D. ordii. and 15f from L. californictis. In 32 of 55 collections, £. dccipiem was the only mite on its host. In only seven collections was it associated with other species of chigger mites. Eitschocngd.'itia pomeiantzi Brennan and Jones, 19.54 Fig. 416 Distribution. CALIFORNIA: Citetlus heecheyi, Citcltus lateralis, Microtus califomicus, Mus niusculus, Neotoma fu.scipes, Perognathus califomicus, Peromyscus bot/lii, Peromyscus califomicus, Peromyscus maniculatus, Peromyscus truei. Tamiasciurus douglasii ( Brennan and Jones, 1954; Gould, 1956). NEVADA; ? ho.st; UTAH: Garfield Co.: Eutamias umhrinus (Brennan ;uid Beck, 19.55). Other Utah records. Garfield Co.: P. manicu- latus, P. truei. Seasonal occurrence. The 49 mites were taken in June. Comments. This species is known in Utah from only one county. Euschocnga.^tia fa.'ioUa Brennan and Beck, 1955 Fig. 414 Distribution. NEVADA: Eutamias dorsalis, Neo- toma lepida, Perognathus parvus (Allred, 1963; Allred and Goates, 1964a, 1964b). UTAH: Washington Co.: N. lepida (Brennan and Beck, 1955). Other Utah records. None. Eit.ichocito(istkt hiteodema Brennan, 1948 Fig. 420 Distribution. CALIFORNIA: Marmota fUnivcn- tris (Gould, 1956). IDAHO: M. flaviventer (Brennan. 1948). MONTANA: Citcllus cotumhianus, M. flaviven- ter, Tamiasciurtis Inuhonicus ( Brenmui. 1948). UTAH: Beaver Co.; M. flaviventris (Allred, 1961). Cache Co.: Peromyscus maniculatus (Brennan and Beck, 1955). Utah Co.: P. nuwiculatus (Ash, 196.3). Other Utah records. Duchesne Co,: Citellu.-, lat- eralis. Seasonal occurrence. A total of 114 larval E. hiteodema was collected during January, June, and from September through November. Great- est numbers were found in June. Comments. Mites of this species have been found in both the Great and Upper Colorado River basins, but are more common at the northerly latitudes. They are knowm from only four counties. In seven of its ten collections, E. hiteodema was associated with chiggers of other species. In two instances it was apparently the only mite on its host. Etischoengaslia rotunda Bremian and Beck, 195.5 Fig. 411 Distribution. UTAH: San Juan Co.; Ochotonu princeps (Brennan and Beck, 1955). Utah Co.: Pero- myscus nianiculatu.ris Perognathus parvus 6% Peromyscus crinitus Peromyscus eremicus Peromyscus maniculatus Pcronu/scus truei 'Sylvikigus audubonii 1% 'Sylvilagus idahoensis \'2% 'Sylvilagus nuttallii 5% Euschoengastia fasolla Neotoma lepida Euschoengastia furmani Neotonui lepida Euschoengastia hoffmanrvje Citellus lateralis Ochotona princeps Euschoengastia lacerta Citellus lateralis Neotoma lepida Euschoengastia lanei Ochotona princeps Peromyscus maniculatus Reithrodontomys megalotis Euschoengastia lanceolata Neotoma lepida Peromyscus eremicus Euschoengastia luteodema Citellus lateralis Marmota flaviventris Peromyscus maniculatus Euschoengastia obesa Peromyscus maniculatus Euschoengastia oregonensis Eutamias quadrivittatus Microtus longicaudus 'Ochotona princeps 5% Perognathus parvus Peromyscus maniculatus Sorex obscurus Euschoengastia pomeruntzi Eutamias umhrinus Peromyscus maniculatus Peromyscus truei Euschoengastia radfordi Duma hcmionus Dipodnmtis microjis Dipodomys ordii Erethizon dorsatum Lepus culifornicus Neotoma lepida Perognathus formosus Peromyscus crinitus Peromyscus eremicus Peromyscus maniculatus Reithrodontomys megalotis 'Sylvilagus idahoensis 6% Euschoengastia rotunda Peromyscus maniculatus Euschoengastia sciuricohi Citellus armatus 'Citellus lateralis 5% Citellus variegatus Eutamias minimus Eutamias umhrinus Marmota flaviventris 'Ochotona princeps 5% Peronu/scus maniculatus Reitljrodontomys megalotis 'Tamiasciurus hudsonicus 6% Euschoengastia soricinus Ochotona princeps Bricham Young University Science Bulletin EtischoengastUi utahcusis Neotoma lepida Perogiuitluis formosus Gahrliepia amcricuna Eutainkis iimbrinus Haemogarnasus alaskensis Blurina hrcvicauda (probably Sorex sp.) Microtus longicaudus Ochotona princepi Pcromyscus maniculatus Xapus princeps Haemogarnasus amhulans Citclhts armatus Citelhis lateralis Clethrionoviys gapperi Dipodomys ordii Eutamias iimbrinus Glaucomys sabrinus Microtus longicaudus Microtus montanus Mus musculus "Neotoma cinereu 12% Neotoma lepida Oclwtona princeps Omjchomys leucogaster Perognathus parvus Pcromyscus maniculatus Peromyscus truei Tamiasciurus hudsonicus Thomomys hottac 'Thomomys talpoides 5% °Zapus princeps 6% Haemogarnasus Uponyssoides Bhrina brevicauda (probably Sorex sp. Microtus longicaudus Haenwgamasus longitarsus Microtus longicaudus Peromyscus maniculatus Thomomys talpoides Haemogarnasus pontiger Eutamias quadrivittatus Neotoma lepida "Peromyscus boylii 7^ Peromyscus eremicus Peromyscus maniculatus Haemolaelaps casalis Citellus lateralis Dipodomys merriami "Mus musculus 5X Neotoma lepida "Peromyscus boylii 7% Peromyscus crinitus Peromyscus maniculatus Peromyscus truci Rattus norvegicus "Thomomys bottae 10% Thorrwmys talpoides Haemolaelaps g^■(>llllJs Omjchomys leucogaster Haemolaelaps glasgowi "Citellus armatus 10^ "Citellus lateralis 15% "Citellus leucurw 10% Citellus richardsonii Citellus townscrulii Citellus trideccrn tincatns "Citellus variegalus 24% "Cynomys leucurus 35% "Dipodomys merriami 10% Dipodomys microps Dipodomys ordii Eutamias minimus Eutamias quadrivittatus Lcpus californicus Mcirmota flat ii ciitrh "Microtus longicaudus Microtus mexicanus 'Microtus montanus Microtus permsylvanicus "Mus musculus Neotoma cinerea "Neotoma lepida Ochototui princeps "Onychomys leucogaster Onychomys torridus Perognathus formosus Perognathus longimemhris Perognathus parvus Peromyscus boylii Peromyscus crinitus Peromyscus eremicus Peromyscus maniculatus "Peromyscus truei Rattus norvegicus Reithrodontomys megalotis Siilvilagus auduhonii Tamiasciurus hudsonicus Thomomys hottac Thomomys talpoides TAifnts princeps H irstionyssus afjinui Eutamias minimus Perognathus formosus Peromyscus truei H irstionyssus hisetosus Neotoma lepida Hirstionyssus femu talis Citellus armatus "Mustela frenata Neotoma lepida Peromyscus eremicus "Thomomys bottae Thomomys talpoides Hirstiorujssus geornydis Neotoma lepido H irstiorujsstis hilli Dipodomys ordii Neotoma lepida Onychomys leucogaster Perognathus flavus Perognathus longimembris Perogruithus parvus Perorn i/scus eremicus Peronryscus mi longUdiiilus 4% 'Microtus moiitaiiii.s 6% Microtua pcnnsi/lvaniciis Ochotoua prince/is Om/chornt/n torridus Pt'romiiscii.s iinmiculutus Phciutcomijs inlirmedius Reithrodoutomy.i megalotis Thomomi/.t talpoidcs H irstionyssiis ncotonuie Dipodomi/s ordii 'Neotomo cincrcu 6% Neotoma lepidu PerognatJius jmrvus Peromt/scus maniculatus- 'Peroniyscus truci 4% Thomomijs tulpoides Hintionyssus slajjordi Spilog^idc gracilis 10% Hirstiomjss-us triucinithiis Citellus leucunis Dipodomys merriumi 'Dipodonu/s microps 10% "Dipodomys ordii 6% Lcpus C(difornicus Neotoma lepidu Perogruithus formosus Perognatlnis longirnemhris Peromyscus crinHus Peromyscus eremicus Peromyscus miiniculatiis Rutttis nonegicuf- Reithrodontoinijs megalotis Hirstionyssus thomomys Thomomys talpoides 3% Hirstionyssus longiehelae "Thomomys talpoides 3% Ochotoua i)riiiceps Hirstiouysstis punctatus Eutamius minimus Eutamius quadrivittutus Olaucoimjs sahrinus Peromyscus maniculutus Hirstionyssus tarsalis Peromyscus maniculutus H irstionyssiis torus Seiurus aherti Hirstionyssus puhistris Sorex palustris Hirstionyssus angustus Citellus armatus Hirsiionyssus eutamiue "Eutamius ipwdriiittatus 3% Reitlirodontomys megalotis Hirstiimyss-us utahcnsU: Citellus armatus Citellus lateralis Citellus tridecendiiiealus Dipodomys ordii "Eutamiai minimus 5% "Eutamius (puidrii ittatus 122 Neotoma lepid^i Perognathus longimend>ris Peromyscus crinitus Peromyscus municidatiis Peromyscus truei Reithrodontomys megalotis Thomomys talpoides Hirstionyssus invaginatus "Citellus armatus "Mus musculus Thomomys talpoides Hypoaspis gurahensis Peromyscus maniculutus Thomomys talpoides Hijpoaspis luhrica Citellus lateralis Citellus spilosoma Peromyscus manicidutus Rattus norvegicus Thomomys hottae Ichoronyssus rohustipes Tadurida mexicana Ischyropoda armatus Citellus varicgatus Dipodomys merriami "Dipodomys microps Dipodomys ordii Erethizon epixanthum Lepus californicus Myotis californicus "Onychomys leucoguster Onychomys torridus Perogruithus flatus "Perognathus formosus Perognathus longimemhris Perognathus parvus Peromyscus manicukitus Peromyscus truei Ischyropoda furmani Dipodomys ordii Microdipodops niegacephalus Neotoma lepida Onychomys leucogaster Perognathus longimemhris Peromyscus inaniculatus Kleenwnia sp. Citellus lateralis Dipodomys microps Dipodomys ordii Eutamius minimus "Onychomys leucogaster Perognathus formosus Perognathus longimemhris Perognathus parvus Peromyscus crinitus Peromyscus maniculutus Luelaps kochi Dipodomys ordii Eutamius minimus Eutamius quadrivittutus "Mierotus longicaudus ° Aticrotus montanus Mierotus pennsylvanicus Phenacomys intermedius Laelaps multispinosus Ondatra zibethicus Peromi/scus manicutalus Laelaps niiltallii Peromi/scus maniculalus Laelaps iiicitis Eutamius quadrivittutus Mierotus longicaudus Mierotus montanus 21% Bkigiiam Vounc University Science Bulletin Pcromy.scus niuiticuliitii), LeewenJwekia umericana BUirina sp. (probably So/ex sp. Peromyscus maniculatus Scupanus- sp. ( probably Sorex s Listrophonis sp. Dipodomys ordii Perognathns formoxu^ Perognathus hngimembris Peromi/scus maniciihituti Macrochcles sp. Citcttus armutus Dipodomys ordii Marmota flaviventris °Mus musculus Onychomys torridus Perognathus hngimembris Peromyscus maniculatus Reithrodontomys megalotis Thomomys hottae Thomomys talpoides Zapus princeps Myobia .sp. Neotoma Icpida Myocoptes sp. Peromyscus maniculatus Reithrodontomys megalotis Myonyssus montanus Ochotona princeps Odontacarus hirsutus °Mus musculus "Neotoma cinerea Neotoma lepida Odontacarus linsdalei Dipodomys microps Dipodomys ordii Neotoma lepida Perognathus formosus Perognathus parvus Peromyscus maniculatus Odontacarus micheneri Neotoma cinerea 'Neotoma lepida Onychomys torridus Perognathus parvus Sylvilagus sp. Omithonysstds aridus Citellus leucurus Ornithonyssus hacoti Citellus tounscndii Dipodomys ordii Eutaniias minimus Neotoma lepida Perognathus parvus Peromyscus boylii Peromyscus crinitus Peromyscus eremicus Peromyscus maniculatus 'Peromyscus truci Rattus norvegicus OrnitJtonyssus sylviarum Marmota flaviventris 'Sylvikigus nultallii Paraxpinturnix globosus \l If Otis sp. Radfordiu bachai Dipodomijs ordii Radfordia lemnina Peromyscus maniculatus 5% Radfordia subuliger Peromyscus maniculatus Reithrodontomt/s 7ncgalotis Spinlurnix orri Antrozous pallidas Steatonyssus antrozoi Corynorhinus rafinesquii 6% Trombicula allredi Neotoma lepida Trombicula arenicola Citellus leucurus Citellus spilosoma Cipiomys parvidens Dipodo/nys merriami 'Dipodomys microps 5'Jf Dipodomys ordii Neotoma lepida 'Perognathus formosus 20i? Perognathus hngimembris Perognathus parvus Peromyscus maniculatus Peromt/scus truei Reithrodontomijs megalotis Tromhicuhi bakeri Citellus armatus Dipodomys ordii Peromyscus maniculatus Trombicula belkini Citellus hteralis Citellus leucurus Clethrionomys gapperi Dipodomys ordii Peromyscus truei Trombicula californiea Citellus armatus Clethrionomys gapperi Microtus hngicaudus Peromtiscus maniculatus Zapus princeps Trombicula doremi Perogiuithus longimembris Dipodomys merriami Dipodomys ordii Trombicula esoensis Peromyscus nuinicukitus Trombicula harperi Citelhts armatus Citellus lateralis 'Clethrionomys gupperi 6? Eutamias mininuis Human Marmota flaiivciilris 'Microtus longieaudus 5'i Microtus montanus Ncotonui lepida 'Ochotona princeps 26% Onychomys leucogustcr Perognathus panus Peromt/scus inanieulatus 'Zapus princeps- 6% Trombicula hoplai CitcUus leucurus Neotonui lepida Perognathus apache Perognathus fxirvus Peromyscus truei Trombicula jewetti Peromyscus manicidatus Trombicula kardosi Mites ok Utah Mammals EutuniUis iimltrinus Troiiihicuhi iiiotitanensis Citcllti.i hitcralis Ct/nomys leucurus Peromyscus crinitus Peromyscus truci Peromyscus niajiicukitus Trotnhicula myotis Bat Mtjotis culifomicus Peromyscus iiutnicuUitus TrombicuUi juinamcnsis Neotonia lepida Peromyscus maniculatus Tromhiculii potosina Perogiuithtis parvus Neotoma Icpidu Microtus loiif^icuudus Tromhiculu sargenti Neotoma lepida Tromliicuhi sid>signatii Reithrodontomys mcgulotis Troinhicula univari PipistreUus liesperus Wluirtoma perplexa Autrozous pallidus Myotis calif nrnicus ZumpticUu bukcri Citellus armatus Eutamias quadrivittatus TABLE 3. HOST-MITE RELATIONSHIPS IN UTAH. Mites indicated with an asterisk are those which were found most frequently on the host. Where applicable, the order of those so marked is designated, number one being most common. Species not marked with an asterisk occurred about equally or with insufficient frequency on the host to warrant frequency designation. Names of hosts enclosed within parentheses are as designated by Hall and Kelson (1959). Antrozous pallidus Spiiiturnix orri Whcirtonia perplexa Bat Troinhicula ini/oth Wliartonia perplexa Citellus armatus ( = Spermojthilus armatus) Amlrolaelaps leviculus Eulaelaps stahulark Euschoengastia criceticola Euschoengastia sciuricola Haemogamasus amhulans 'Haemolaelaps glasgowi 1 Ilirstioni/ssus angustus llirstiout/ssus femuralis "Hirstionyssus invagmatus 2 Hirstiomjss-us isabellinus Hirstionyssus utuhensis Macrochcles sp. Trom]>icula bukcri Trombicula califoniica Tromhiculu bar peri ZumpticUa bakiri Citellus lateralis ( —Spermophilus lateralis) Androlaelaps leviculus Euschoengastia cordiremus Eusclioengustia criceticola Euschoengastia deci/iicns Euschoengastia hoffnninnae Euschoengastia lacerta Euschoengastia luteodema Euschoengastia sciuricola Haemogamasus amhulans Hacmolaelaps casalis ° Hacmolaelaps glasgowi Hirstioni/ssus utahensis Hypoaspis lubrica Kleemania sp. Trombicula helkini Trombicula harperi Trombicula montanensis Citellus leucurus ( ^Ammospermophilus leucurus) 'Hacmolaelaps glasgowi Hirstionyssus incumptus Hirstionyssus triacanthus Ornithonyssus aridus Trombicula arenicola Trombicula belkini Trombicula hoplai Citellus richardsonii { =Spermophilus richardsonii) Hacmolaelaps glasgowi Citellus spilosoma ( =-- Spermophilus spilosoma) Euschoengastia cynomyicola Hypoaspis lubrica Trombicula arenicola Citellus townsendii (—Spermophilus townsendii) Euschoengasiia criceticola Hacmolaelaps glasgowi Ornithonyssus bacoti Citellus tridecemlineatus ( ^Spcrmophdus tridecem- lineatus) ° Hacmolaelaps glasgowi Hirstionyssus utuhensis Citellus variegatus ( = Spermophilus variegatus) Brevisterna utahensis Euschoengastia sciuricola 'Hacmolaelaps glasgowi Hirstionyssus incomptus Ischyropoda armatus Clethrionomt/s gupperl Haemogamasus und)tdans Trombicula belkini Trombicula californica Trombicula harperi Corynorhimts rafinesquii ( =Corynorhinus toivn- sendii ) Eubruchyluelaps dcbilis Steatonyssus antrozoi Cynomys parvidens Tromb icu la aren icola Cynonu/s leucurus 'Hacmolaelaps ghisgowi T rombicula montanensis Dama hemionus Euschoengastia radfordi Dipodomi/s merriami Brigham Young Univehsity Science Bulletin Brevisterna utuhensis Euschoengastia decipiens HaemolaeUips ctisalis 'HacnwUieUips g/«,sgoai Hirxtiontjsaus incoiuptiis Hirstioni/ss)is triiiciintluis Ischijropoda armutus Trombicuhi arenicola Tmmhicuhi doremi Dipodormjs microps Eit.'ichocnfiii.',1ui criceticola Eiisclwcngditia decipiens Euxcliocitf^ii.stia nidfordi Huemohehips glasgowi Hirst iomjstius incomptuti 'Hirstinmjssus triacanthus Ischijropoda armatus Kleem-ania sp. Odontacarus linsdalei Tromhiculti arenicola Dipodomijs ordii Androhielaps (cii'ru/u.v Euschocngastiu criceticuhi Euschofiigdstia decipiens Euschociigiislia radfordi Hiiciiuigiiiiuisus amhulans 'Htuimiluihii's gh.sgowi 2 Hirstionys.ms hilli 'Hirstionysstts incoinptus 1 Hirstiomjssus neotoinae "Hirstionyssus triacanthus 3 Hirstionyssus utahensh: Ischyropoda armatus Ischyropoda furtnani Kleemania sp. Laehips kochi Listrophorus sp. Macrocheles sp. Odontacarus li)tsdalei Ornithonyssus hacoti Radfordiu bachui Trond>icuhi arenicola Tromhicula hakeri Trondncula helkini Tromhicula doremi Erethizon epixanthum ( ^Ercthizon dorsatum] Euschoengustia decipiens Euschocngastia radfordi Ischyropoda arjnutus Eutamias dorsalis Hirstiomjssus sp. Eutamias minimus Euschocngastia sciuricola Haemolaehips glasgowi Hirstionyssus uf finis Hirstionysstis punetatus Hirstionyssus utahensis Kleemania sp. Laelaps kochi Ornithonyssu\ hacoti Tromhicula harju-ri Eutamias ipiadrivitt-titus Brevisterna moiUunus Dernuinyssus hecki Euschocngastia oregoncnsis Hacmogamaxus pontiger Haemolaelaps glasgowi Hirstionyssus eutamiae Hirstionyssus isahellinus Hirstionyssus punetatus "Hirstionyssus utahensis Laelaps kochi Laehips incilis Zumptietla hakeri Eutamias uiidirinus Euschocngastia pomerantzi Euschocngastia sciuricola Gahrliepia americana Hcwmogamasus amhulans Tromhicula kardosi Claucomys sabrinus Haemogamasus amhulans Hirstionyssus jmnctatus Human Tromhicula harperi Lepus califomicus Brevisterna utahensis Dermanyssits hecki Etdjrachylaelaps circularis ° Euschocngastia decipiens Etischoengastia radfordi Haemolaelaps glasgowi Hirstionyssus triacanthus Ischyropoda armatus Marmota flaviventris Dernwnyssus hecki Euschoengastiti luteodcma Euschocngastia sciuricola Haemolaelaps glasgowi Macrocheles sp. Ornithonyssus sylviurum Tromhicula harperi Microdipodops mcgucephalus hiiuiropoda furmani Micriitus leri Ondatra zihethicus Laelaps multispinosus Onychomys Icucogastcr Androliirlaps Icvicuhis Euhrachi/lavlups crincii Haemogamasus amhulans Haemolaelaps geomys " Haemolaelaps glasgowi Hirstionyssus hilli 'Ischyropoda arntatus Ischyropoda furmani Kleemania sp. Tromhicula harperi Onychomys torridus Haemolaelaps glasgowi Hirstionyssus isahellinus Ischyropoda armatus Macrocheles sp. Odontacarus micheneri Perognathus apache Androlaelaps leviculus Tromhicuhi hoplai Perognathus jlavus Hirstionyssus hilli Ischyropoda armatus Perognathus formosus Androlaelaps leviculus Brevisterrw utahensis Euschoengastia decipiens Euschoengastia criceticola Euschoengastia utahensis Haemolaelaps glasgowi Hirstionyssus affinis Hirstionyssus triacanthus Ischyropoda armatus Kleemania sp. Listrophorus sp. Odontacarus Unsdalei 'Tromhicula arenicola Perognathus longimemhris Bernia marita Euhrachylaelaps circularis Euschoengastia decipiens Haemolaelaps glasgowi Hirstionyssus hilli Hirstionyssus incomptus Hirstionyssus triacatxthus Hirstionyssus utahensis Iscfiyropoda armatus Ischyropoda furmani Kleemania sp. Listrophorus sp. Macrocheles sp. 64 Bricham Young University Science Bulletin Tromhicula arenicola Troml)icula doremi Perognathus parvus Androhielaps leviculus Euhrachtjlaclups circtilaris Eubrachylaelups hollisteri Euschoenga.ttia cordiremus Euschoengdiiia decipiens Euschoengastia oregonensis Haemogantaxus innbulans 'Haemolaelaps glasgowi Hirstionyssus hilli H irstionyssus incomptus Hirstionyssus neotomae Ischyropodii armatus Kleenwniii sp. Odontacarus linsdulei Odontacarus micheneri Ornithoni/ssus bacoti Trombicula arenicola Trombicuhi harperi Trombicula hoplai Trombicula potosina Perogimthus sp. Eubrachylaelups crowei Ischyropoda furmani Trombicula doremi Peromyscus boylii Brevisterna utahensis Dermanyssus becki Eubrachylaelups circularis Euschoengastia criceticola Haemogamasus pontiger Haemolaelaps casalis Haemolaelaps glasgowi Ornithonyssus bacoti Peromyscus crinitus Androlaelaps leviculus Brevisterna utahensis Dermanyssus hechi Euhrachyhiclaps circularis Euhrachylaelaps debilis 'Eubrachylaelups hollisteri Euschoengastia decipiens Euschoetigastia radfordi Haemolaelaps casalis Haemolaelaps glusgowi Hirstionyssus incomptus H irstio n i/ssus triacunth us Hirstionyssus utahensis Kleemaniu sp. Ornithonysstis bacoti Trombicula monlanensis Peromyscus eremicus Brcvistrena utahensu Dcrmanysstis bccki Eubrachyluelaps circuluris Eubrachyluclaps debilis Eubrachylaelups hollisteri Euschoengastia criceticolu Etischoengavtia decipiens Euschoengastia lanccolata Euschoengastia radfordi Haemogamasus pontiger Haemolaelaps glasgowi Hirstionyssus femuralis Hirstionysstts hilli Hirstionyssus incomptus Hirstionyssus triacanthus Ornithont/ssus bacoti Peromi/scus maniculutus Androlaelaps leviculus Brevisterna utahensis Chatia ochotorui Cliatiu setosa Dermanyssus becki ° Eubrachylaelups circularis E\d>rachylaelups debilis Eubrach i/laelaps hollisteri Eulaclaps stabularis Euschoengastia cordiremus "Euschoengastia criceticola Euschoengastiu decipiens Euschoengastia lunei Euschoetigastia luteodema Euschoengastia obesa Euschoengastia oregonensis Euschoengastia pomerantzi Euschoengastia radfordi Etischoetigastia rotunda Euschoengastia sciuricola Haemogamasus alaskensis Haemogamasus ambulans Haemogamasus longitarsus Haemogamasus pontiger Haemolaelaps casalis "Huemolaelaps glasgowi Hirstionyssus hilli Hirstionyssus incomptus H irstii^nyssus isabellinus Hirstionyssus neotomae Hirstionyssus punctatus Hirstiomjssiis tarsalis Hirstioni/ssus triacanthus ° Hirst ioni/sstis utahensis Hypoaspis gurabensit Hypoasf)is lubrica Ischyropoda armatus Ischyropoda furmani Kleemaniu sp. Laelaps multispinosus Laclaps nuttallii Laelaps incilis Lecuwcnhockia americanu Listrophorus sp. Macrochehs sp. Mt/ocoptes sp. Odo ntacarus linsdalei "Ornithonyssus bacoti Radfordia lemnina Rudfordia sidyuliger Trombicula arenicola Trombicula bakeri Trombicula calif ornica Trombicula esoensis Trombicula harperi Trombicula jewetti Trombicula montancnsis Trombicula myotis Trombicula panamensis Peromyscus truei Brevisterna utahensis Dcrnuinyssus becki ° Eubracht/hielaps circularis Eubrachyluelaps debilis Eubrachylaelups hollisteri Euschoengastia criceticola Euschoengastia decipiens Mites of Utah Mammals 65 Eufichocnf^u.stui i>oiiicr(iiitzi Hacmogurnaiiis amhulans Udciiwlut'UijK castdi.s ° HacmoUichips glus'^oiLi Hirstionyssus uffiiiis Hirstionysstts ncotoniuc Hirstionyssus tttahensis Ischt/ropoild unnatus 'Omithonyssus hacoti Trotnhiculu intniicohi TrornliicuUi hclkiiii Tronihiculu hoplui Tromhiculu iiuintuncn.six Pcromyscus sp, Hacmofji^unuisus ulaskensis Pheriaconu/s intcniicilius Hirstionysstts i.siihellinus Luelaps hoclii Pipistrclliis licsj>cnis Tromhiculu univuri Rtrttus norvegicus Euhielups stahularis HucriiohicUips casalus ° Hucnwlaelups glasgowi H irstio nyssti s incom))tus Hirstiomjssus triacanthus Hypoaspis luhrica 'Onithoiiysstis Imcoti Reithrodontomtjs megalotis Eubruchtjluclaps dehilis Euschocngasiia criceticolu Eusclwengastiii lanei Euschoengastia radfordi Euschoengustia sciurivola 'Haemolaelaps glasgowi Hirstionyssus etitamiae Hirstionyssus incomptus Hirstionyssus isuhellinus Hirstionyssus triacanthus Hirstiomjsstis utahensis Mticritcheles sp. Myi^coptcs sp. Radfordia stdndiger Tromhicula arenicola Tromhiculu suhsignata Sciurus aberti Hirstionyssus torus Sorex obscurus ( = So rex vagruns) Euschoengastia oregonensis Sorcx pultistris Hirstiont/sstis palustris Spiloguk' grucili.',- Hirstionysstts stafjordi Stjhilagus uudubonii Etthrucltylaclaps circuluris Euschoengastia decipiens Hacmolaelups glasgotvi Syliilagus idahoensis Euschoengastia decipiens Euschoengastia radfordi Sylvilagus nuttallii Euschoengastia decipiens Omithonyssus sylviarum Sylvilagus sp. Odontacartis iiiicheneri Tamiasciurtts hudsonicus Euschoengastia sciuricoUi Haemogamastts ambtiUins Haemolaelaps glasgotvi Thomomys bottae { ^^Thomomys iinibrinus) Haemogamasus ambtdans 'Haemolaelaps casalis Haemolaelaps glasgowi 'Hirstionyssus femuralis Hypoaspis luhrica Macrocheles sp. Tho mo m ys talpoides 'Haemogamasus ambtilans Haevtogamasus longitarstts Haemolaelaps casalis Haemolaelaps glasgowi Hirstionyssus femuralis Hirstionysstts invagittatus Hirstioiiyssus isabellinus 'Hirstiontjssus longichelae Hirstionyssus neotomae Hirstionyssus thomomys Hirstionyssus utahensis Hypoaspis gurahensli- Macrocheles sp. Zapus princeps Haemogamastts ulaskensis 'Haemogamasus ambtilans Haemolaelaps glasgowi Macrocheles sp. Trombicula californica 'Trombicula harperi DISCUSSION More than 9,(KK) small mammal.s repre.sent- ing 94 species of 51 genera were e.xamined for ectoparasites. Approximately 13,(K)0 mites were found on 4,015 of those examined. These, plus other previously listed records for Utah, repre- sent 111 species of 34 genera of mites which in- clude 12 new species in thre^ genera, one genus being new. During the si.xteen-vear period covered by this report, numerous individuals were involved with the collection of specimens. Even though stan- dardized technicjues were used, there likely was considerable variation in the efficiency of ecto- parasite retrieval. Consequently, the incidence of ectoparasites as shown in Table I is relative only to tlie efficiency of our techniques. Hosts were collected by different methods, e.g., live-trap- ping, shooting, etc. and in almost all cases were examined in the field under varying conditions of weather, pressure of time, and lack of ade- (juate facilities. It is likely that some ectopara- sites were lost or overlooked, particularly those 66 Bhigham Young Univekshv Science Bulletin requiring special techniques for their recovery. Nevertheless, over a period of years involving large numbers of specimens, we expect that each of the mite taxa would have been taken at least once. The validity of host identification may be questionable in some instances, although for the most part identifications were made by a quali- fied mammalogist in the field, or representative specimens were returned to the reference mu- seum at Brigham Young University for com- parison and verification. As shown in the listing of mite-host relation- ships ( Table 2 ) , the variety of hosts from wliich mites of a particidar species were recovered may be correlated with the numbers of animals ex- amined. For example, we examined more mice belonging to Peromijscus maniculatus than of any other species, and mites of more species were found on tliis mouse than on any other mammal studied. We assume that mites of almost any species occasionally are accidental symbionts of host animals living in the same habitat. Examination of large numbers of hosts increases the potential of finding such infestations. There is a tendency for host preference, and perhaps even for host specificity in some cases, whereas mites of some species are widely distributed geographically and occur on a variety of animals. For example, Haemolaclaps glasgowi is a widely distributed species and occurs on the greatest variety of mammals in Utah. This distribution and host relationship apparently is typical for this most cosmopolitan species. Other sui-veys may reveal a frequency of occurrence of mites of a particular species dif- ferent from that indicated by us (Table 3). We assume that mites of all species normally as- sociated with a particular host have an equal opportunity of infesting that host, but the season- al activity of different species may vary. Mites of one species may be more common on a host in summer, whereas those of another species may be more common in winter. In our field activities we collected the year round, but most mammals were taken in late spring, summer and early autumn. Even though our data on mite-host relation- ships, relative abundance, frequency of occur- rence and seasonal and geographic distribution may vary from other surveys, the information presented here should serve as a basis for fur- ther studies directed at specific localities or species. More work needs to be done on mam- mals that have been taken only in few numbers or only at certain times of the year. REFERENCES AUred, D. M. 1954a. Mites touiul on mict- of the genus Peromijscus in Utah. Unpub. Ph.D. disserta- tion, Dep. Inv. Zool. and Entomol., Univ. Utiih, Salt Lake City. AUred, D. M. 1954b. Observations on the stylosome (feeding tube) of some Utah chiggers. Proc. Ut;di Acad. Sci., Arts, and Letters, 3:61-63. Allred, D. M. 1954c. Mites found on mice of the genus Peromijscus in Utah. I. General infestation. Great Ba.sin Nat. 16( 1-4) :23-31. Allred, D. M. 1957a. A new species of mite, Hirs- tionyssus hisetosus, from the nests of the desert wood rat, Neotoma lepidu lepida Thomas (Acarina: Denriimyssidae ) . Proc. Entomol. Soc. \Va.shington, 59(2):83-89. Alb-ed, D. M. 1957b. Mites found on mice of the genus Peromijscus in Ut;Ui. II. Family Haemogama- sidae. Proc. Entomol. Soc. Washington. 59(1):3I- 39. Allred, D. M. 1957c. Mites foiuid on mice of the genus Peromi/scus in Utah, III. Familv Dermanvs- .sidae. Amer.' Midland Nat.. 57( 2) :45()-460. Allred, D. M. 1957d. Mites fomid on mice of the genas Peromyscus in Utah. V. Trombiculidae and miscellaneous families. Great Basin Nat., 17(3-4): 95-102. Allred, U, M. 19.57c. Notes on the life history and bionomics of the wood rat mite. Brevislertui utahen- sis ( Acarina ) , Trarts. Amer, Microscop. Soc., 76 (l):72-78, Allred, D, M, 19.57f, The male, deuton\mph and proton\mph of the mite Euhrachijlaehips circularis (Ewing) (Acarina: Laelaptidae) witli notes on morphological variations, Ann, Entomol, S(K. Amer., .5()(2):2()6-209. Allred, D. M. 1957g. Setal variations on mites of the species Brevisterna uiahensis (Ewing) (Acarina). Proc. Utah Acad. Sci,, Arts and Letters, 34:51-54. Allred. D. M. 19.58. Mites found on the genu.s Pero- myscus in Utah, I\'. Families Laelaptidae and Ph\-toseiidae. Pan-Pacific Entomol., 34(1): 17-32. .Mired, D, M, 1961, Parasitic mites on miumots in Utixli, J. Parasitol., 47(1):124. Allred, D. M. 1962. Mites on gniishopper mic-e at the Nevada Atomic Test Site. Great Basin Nat.. 22(4);101-104. Allred, D. M. 1963. Mites from pocket mice at the Nevada Test Site (Acarina). Proc. Entomol. Soc. Wa,shington, 65( 3) :231-232. Allred, D, M. 1965, Clarification of type data for Isclii/ropoda furmani Keegan. J. Parasitol., 51(4): 604.' Mites of Utah Mammals 67 AUred, D. M., ;uid D E. Beck. 1953a. Mite fauna of woodrat nesLs in Utali. Proc . Utah Acad. Sei., Arts, and Letters, 30:53-56. AUred. D. M., and D E. Beck. 19631). A new speeies of Acomatacanis ( Acarina, Trombiciilidae ) from Utali. Cireat Ba.sin Nat. 13 ( .3-4 ): 87-90. AUred, D. M.. and D E. Beck. 1962. Eeological dis- tribution of mites on lizards at the Nevada Atomic Test Site. Herpetologiea, 18(1):47-51. AUred. D. M., and D E. Beck. 1964. Mite.s on rep- tiles at the Nevada Atomic Test Site. Trans. Amer. Microscop. Soc, 83( 2 ) :266-268. AUred, D. M., and M. A. Goates. 1964a. Mites from Mammals at the Nevada Test Site. Great Basin Nat., 24(2):71-73. AUred, D. M., and M. A Goates. 1964a. Mites from wood rats at the Nevada Test Site. |. Parasitol., 50(1):171. AUred, D. M., and N. J. Marcliette. 1957. E.xperiment- al feeding of tlie mite Brevisterrui utahensis ( Acar- ina: Haemogamasidae ) . J. Parasitol., 43(l):51-54. AUred, D. M., and E. J. Roscoe. 1957. Parasitic mites in desert wix)d rat nests with notes on free-living forms. Trans. Amer. Microscop. Soc. 76(4):389- 403. Asanuma. K. 1952. Two new species of blood- sucking mites parasitic on the striped mouse Apodemus agrarius, from Manchuria. Res. Inst. Nat. Resources, Tokyo, Misc. Rep., 25:86-92. Ash, D. B. 1963. Seasonal study of mites on deer mice {Perotnyscus inaniculatus) in a chaparral community. Unpub. master's thesis, Brighimi Young Univ.. Provo, Utah. Baker, E. W., T. M. Evans, D. J. a)uld, W. B. Hull, and H. L. Keegan. 1956. A manual of parasitic mites of medical or economic importance. Nat. Pest Control Assoc, Inc., New York. Beck, D E. 1955. Distributional studies of panisitic arthropods in Utah, determined as actual and po- tential vectors of Rocky Mountain spotted fever ;md plague. Brigham Young Univ. Sci. Bull., Biol. Ser.. l(l):l-64. Bradsliaw, G. V. R., and A. Ross. 1961. Ectoparasites of Arizona bats. J. Arizona Acad. Sci., 1(4) : 109- 112. Brennan, J. M. 1946a. Two new species of Trom- hiculo: T. moutaneiisis and T. aplodontitic ( Ac;u- ina, Trombiculidae ) from northwestern United States. J. Para.sitol., .32(51:441-444. Brennan. ]. M. 1946b. A new genus and species of chigger, Cliatia seto.ia ( Trombiculidae. Acarina ) , from northwestern United States. J. Parasitol., 32(2): 132-1.3.5. Brennan, J. M. 1947. New species of chiggers (Acar- ina, Trombiculidae) from bats of the Nearctic Re- gion. J. Para.sitol., 33(3) :245-252. Brenn;m, J. M. 1948. New North American chiggers (Acarina, Trombiculidae). J. Parasitol., 34(6) :465- 478. Brennan, J. M. 1952. Tromhicula cynos Ewing, 1937, and three related new species (Acarina: Trombi- culidae). Wasmiuin J. Biol.. 10(l):55-65. Brennim. J. M. 1965. Five new chiggers from soutli- westem United States (Acarina: Trombiculidae). J. Parasitol., 51( 1 ): 108-113. ♦See footnote on p. 3. Brennan, J. M., and D E. Beck. 1955°. The cliiggers of Utah (Acarina: Trombiculidae). Great Basin Nat., 15(1-4): 1-26. Brennan, J. M., and E. K. Jones. 1954. A report on the chiggers (Acarina: Trombiculidae) of the Frances Simes Hastings Natural History Reservation. Monterev County, California. Wasmann J. Biol., 12(2): 155-194. Brennan, J. M., and E. K. Jones. 1959. Keys to the chiggers of North America with synonymic notes and descriptions of two new genera (Acarina: Trombiculidae). Ann. Entomol. Soc. Amer., .52(1): 7-16. Brennan, j. M.. and G. VV. Wliarton. 19,50. Studies on Nortli American chiggers. No. 3. The subgenus Neotromlyicidti. Amer. Midland Nat.. 44(1):153- 197. Burgess, G. D. 1955. Arthropod ectoparasites of Richardson's ground squirrel. J. Parasitol., 41:639- 640. Crossley. D. A., and L. J. Lipov.sky. 1954. Two new chiggers from the central states. (Acarina, Trom- biculidae). Proc. Entomol. Soc. Washington, 56(5): 240-246. Drummond, R. O. 1957. Ectoparasitic acarina from small mammals of the Patuxent Refuge, Bowie, Maryland. J. Parasitol., 43(1):.50. Durrant, S. D. 1952. Mammals of Utah. Univ. Kansas Pub.. Mus. Nat. Hist., 6:l-.549. Eads, R B. 1951. New mites of the genus Androlae- laps Berlese. J. Parasitol., 37( 2) :212-216. Eads, R. B., and B. G. Hightower. 1951. A new N eoichoromjssus from the pack rat, Neotoma mi- cropus (Acarina, Lelaptidae). Proc. Entomol. Soc. Washington, 53 ( 5 ) : 295-300. Eads, R. B., and B. G. Highwater. 1952. A new mite of the genus N eoichoromjssus (Acarina, Laelap- tidae). Proc. Entomol. Soc. Washington, 54(1)32- 35. Eads, R. B., G. C. Menzies, and V. 1. Miles. 1952. Acarina taken during west Texas plague stxidies. Proc. Entomol. Soc. Wa.shington, 54( 5) :250-253. Eads, R. B., H. A. Trevino, and E. G. Campos. 1965. Ectoparasites of the spiny pocket mouse Liomys irroratus texensis Merriam. Southwestern Nat., 10: 17-21. Eddie, B., D. F. Meyer, F. L. Lambrecht, and D. P. Funman. 1962. Isolation of Ornithosis bedsoniae from mites collected in turkey quarters imd from chicken lice. J. Infectious Dis., 110:231-237. EUis, L. L., Jr. 1960. A survey of the ectoparasites of certain mammals in Oklahoma. Ecology, 36:12- 18. Elzinga R. J. 1960. Biotic associations of rodent and ectoparasite populations. Unpub. Ph.D. dissertation. Dept. Zool. and Entomol.. Univ. Utah, Salt Lake City. Elzinga, R. J., and D, M. Rees. 1964. Comparative rates of ectoparasite infestation of deer and harvest mice. Proc. Utah Acad. Sci., Arts, and Letters, 41(2);217-220. Ewing, H. E. 1922. The dermanvssid mites of North America. Proc U.S. Nat. Mus.', 62(13): 1-26. Ewing, H. E. 1933. New genera and species of para- sitic mites of the superfamily Parasitoidea. Proc. U.S. Nat. Mus., 82(30): 1-14. Bnit;HANr VouNc; IFninehsitv Science Bulletin Farrell, C. E. 1956. Chiggers of tlie genus Euschoen- gastia (Acarina: Trombiculidae ) in North America. Proc. U.S. Nat. Mils., 106:85-235. Finley, R. B., Jr. 1958. The wood rats of Colorado: distribution and ecology. Univ. Kansas Publ. Mus. Nat. Hi.st., 10(6):213-552. Florschutz, O., Jr., and R. F. Darsie, Jr. 1960. Addi- tional records of ectoparasites on Delaware mam- mals. Entomol. News. 71(2):45-52. Fonesca, F. da. 1948. A monograph of the genera and species of Macronyssidae Oudemans, 1936 (synom.: Liponissidae Vitzthum, 1931) (Acari). Proc. Z(X)1. Soc. London, 1 18( 11 ) :249-334. Francis, E., and G. C. Lake. 1922. Transmi.ssion of Tularemia by the bedbug, Cimex lectularius. U.S. Public Health Rep., 37:83-95. Furman, D. P. 1954a. A revision of the genus Pnciiinonyssus (Acarina: Halarachnidae). J. Para- sitol., 40(l);31-42. Furman, D. P. 1954b. A new species of Androlaelaps from Perognaihus in southern California. Pan- Pacific Entomol., 3()( 2) :119-124. Furman, D. P. 19.5.5. Revision of the genus Euhraclnj- laelaps (Acarina: Laelaptidae) with the description of two new species from Mexico. Ann. Entomol. Soc. Amer.. 48( l-2):51-.59. Furman, D. P. 1957a. Revision of the genus Sterno- stotna Berlese and Trouessart (Acarina: Rhinonys- sidae). Hilgardia, 26( 10) :473-495. Furman. D. P. 19.59a. Observations on the biology and morphologv of Haeinogcinuisu.s ambukitis (Thor- ell) (Acarina: Haemogamasidae). J. Parasitol.. 45(3):274-280. Furman, D. P. 1966. Biological studies on Haemolue- lapti centrocarpus Berlese with observations on its cla.ssification. J. Med. Entomol., 2(4) :331-335. Furman, D. P., and V. J. Tipton. 1955. The genus Stiionijs.ms Tiralxischi (Acarina: Dermanyssidae ) including a new species from Pika. J. Parasitol., 41(2):179-184. Furman, D. P.. and F. J. Radovsky. 1963. A new speci&s of Ortiitliom/ssus from the \\ hitc-tailetl antelope squirrel, with a rediagnosis of the genus Oniitlwnijssus. Pan-Pacific Entomol., 39(2):75-79. George, J. E., and R. W. Strandtmann. 1960. New recx>rds of ectoparasites on bats in west Texas. Southwestern Nat., 5(4) :228-229. Goates, M. A. 1963. Mites on kangaroo rats at the Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol. Ser., 3(4): 1-12. Gould. D. ). 1950. A new species of chigger from California (Acarina; Trombiculidae). Wasmann J. Biol., 8(3):367-370. Gould. D. J. 1956. The larval Trombiculid mites of California (Acarina: Trombiculidae). Univ. Cali- fornia Pub. Entomol., 2(1):1-116. Greenberg, B. 1952. A review of the New World Acotiiatac-anis (Acarina. Trombiculidae). Ann. Entomol. S(K-. Amer., 45(3 ) :473-491. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. Ronald Press, New York. V. 1 and 2. Hansen, C. G. 1964. Ectoparasites of mammals from Oregon. Credit Basin Nat., 24( 2) :7.5-81. Hays, K. L., and F. E. Guyton. 1958. Parasitic mites (Acarina: Mesostigmata ) from Alabama mammals. J. Econ. Entomol, 51:259-260. Hayward, C. L., D E. Beck, and W. \V. Tanner. 1958. Zoolog)' of the Upper Colorado Ri\er Basin. I. The biotic communities. Brigham Young Univ. Sci. Bull, Biol. Ser.. 1(3): 1-74. Hightower. B. G., V. W. Lehman, and R. B. Eads. 1953. Ectoparasites from mammals and birds on a quail preserve. J. MammiJ., 34:268-271. Ho. B. C. 1962. Ectoparasite-host associations in three areas in Utah and Wyoming. Unpub. PhD dissertation, Dept. ZckjI. and Entomol, Univ. Utah, Salt Lake City. Howell, J. F., and R. J. Elzinga. 1962. A new Rad- fordia (Acarina: Myobiidae) from the Kangaroo Rat and a kc) to the known species. Ann. Entomol. Soc. Amer., '55( 5) :547-555. Howell J- F-, D- M- AUred. and D E. Beck. 1957. Seasonal population fluctuations of mites in desert wood rat nests in central Utah. Ecology. 38(1):82- 88. Huebner, R. J., W. L. Jelli.son, and C. Pomerantz. 1946. Rickettsialpox — a newly recognized rickett- sial disease. IV. Isolation of a rickettsia apparently identical with the causative agent of rickettsialpox from Alhdermantjssus sanguineus, a rodent mite. U.S. Public Hedth Rep., 61:1677-1682. Jameson, E. W^. Jr. 1947. A new mite, Euhrachylae- laps crowd, from the Grasshopper Mouse, Onycho- mys leucogaster arcticeps (Rhoads). J. Parasitol, 33(5):391-392. Jameson, E. W., Jr. 1950a. H irstionyssus ohsolctus, a new mesostigmatic mite from small mammals of the western United States (Acarina). Proc. Biol. Soc. Washington, 63:31-34. Jameson, E. W., Jr. 1950b. Notes on mites of the genus Neoichoronyssus. v\'ith the description of a new subgenus and three new species of tlie sub- genus Hirst iont/ssus (Acarina, Laelaptidae, Macro- nyssinae). Proc. Entomol. Soc. Washington, 52(4): 161-172. [ameson, E. W., Jr. 1950c. EubrachyhieUips debilis, a new Laelaptid mite (Acarina: Laelaptidae) para- sitic on the Deer Mouse. Peromyscus maniculatus (Mammalia: Cricetidae). J. Parasitol. .36( 1 ) :62-64. [ameson, E. W.. Jr. 1950d. External parasites of the short-tailed shrew, Blarina hrcvicauda (Say). J- Mammal. 31(2) : 138-145. Jameson, E. W., Jr. 1955. A summary of the genera of Myobiidae (Acarina). |. Parasitol, 41(4):407- 446. Jameson, E. W.. Jr., and J. M. Brennan. 1957. An environmental analysis of some ectoparasites of small forest mammals in the Sierra Nevada, Cali- fornia. Ecol Monog. 27:45-54. Jenkins, E. 1965. The parasitology of the ground squirrels of western Utah. Unpub. master's tliesis, Dep. Zool and Entomol, Univ. Utah, Salt Lake City. Johnston, D. E. 1959. Some new sviionymy in the Haemogamasidae, Laelaptidae and Diplogyniidae indicated by an examination of Bank's types of Mesostigmata (Acarina). P.syche, 66(4):60-62. Kardos. E. H. 19.54. Biological and systematic studies of the subgenus Neotrondncula (Genus Tromhicula) in the central United States. Univ. Kansas Sci. Bull. 36(4):69-123. Keegan, H. L. 1944. On a new genus and species of 'parasitid mite. J. Parasitol, ,30; 181-183. Keegan, H. L. 1946. Six new mites of tlie super- family Parasitoidea. Trans. Amer. Microscop. Soc., 65(r);69-77. Mites ok Utah Mammal> 69 Keegan, H. L. 1949. Euhufmogutnaxtjs utalictiii-s- Ewing, 1933, redescribed as type species of new- genus Brevitterna (Acarina; Laelaptidae ) . Trans. Amer. Microseop. Soc.. 68(3) :222-227. Keegiui, H. L. 1951. The mites of the subfamily Haemogamasinae (Acari: Laelaptidae). Proc. U. S. Nat. Mus., 101 (3275): 203-268. Keegan. H. L. 1953. Collections of parasitic mites from Ut;ili. Great Basin Nat., 13( 1-2) :35-42. Keegan, H. L. 19.56b. Original illustrations of Hae- molaelaps rnarsupialis Berlese, 1910, and of five additional Haeiyiolaclaps species described, but not figured bv Berlese. Trans. Amer. Microseop. Soc., 75(3):314-319. Keegan, H. L. 1956b. Ischyropoda furnuini n. sp., a new ectopara-sitic mite from Utah. J. Parasitol., 42:311-315. Krantz, G. W. 1959. New s)Tionymy in the Der- manyssinae Kolenati, 1859, with a description of a new species of Deniianyssus ( Acarina, Demiiinys- sidae). Entomol. Soc. Washington, 61(4) : 174-178. Krantz, G. W. 1962. A review of the genera of the family MacrocheUdae Vitzthum (Acarina: Meso- stigmata). Acarologia, 4( 2) : 143-173. Lipovsky, L. J., D. A. Crossley, Jr.. and R. B. Loomis. 1955. A new genus of chigger mites. J. Kansas Entomol. Soc, 28(4) : 136-143. Loomis, R. B. 1954. A new subgenus and six new species of chigger mites (Genus Trombicuh) from tlie central United States. Univ. Kansas Sci. Bull., .36(13):919-939. Loomis, R. B. 1956. The chigger mites of Kansas (Acarina, Trombiculidae ) . Univ. Kansas Sci. Bull., 37(19): 1195-1443. Loomis, R. B. 1963. The discovery of cliiggers ( Acar ina: Trombiculidae) in the nasal passages of cri- cetid rodents from California, with descriptions of two new species. J. Parasitol.. 49:330-333. Loomis, R. B., and M. Bunnell. 1962. A new species of chigger, genus Euschoengastia ( Acarina, Trom- biculidae), with notes on other species of chiggers from the Santa Ana Mountains, California. Bull. Southern California Acad. Sci., 61 ( 3) : 177-184. Macfie. J. W. S., and J. G. Thomp.son. 1929. A trvpanosome of the canarv. Trans. Roy. Soc. Trop. Med. Hyg., 23:185-191. Manwell. R. D., and C. M. Johnson. 1931. A natural trypanosome of the canary. Amer. |. Hvg., 14:231- 2.34. McKinley, I). J. 1963. The morphologN and biologv of Hacmolaehps camlis Berlese (Acarina: Meso- stigmata). Ann. and Mag. Nat. Hist.. 6(62) :6,5-76. Miller, R. S., and R. A. Ward. 1960. Ectoparasites of pocket gophers from Colorado. Amer. Midland Nat.. 64(2):382-391. Myklebust, R. J. 1951. Some studies on tlie consortes of the Norway rat in Utah County. Utah. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. Pratt, H. D., and N. E. Good. 19.54. Distribution of some common domestic rat ectoparasites in tlic United States. J. Para.sitol., 40(2) : 1 13-129. Pratt, H. D., J. E. Lane, and F. C. Harmston. 1949. New locality records for Allodermanyssus siin- guineus, vector of Rickettsialpox. J. Econ. Entomol., 42(3):414-415. Radovsky. F. J. 196<)b. Uacinoguiiuixus li potty ssokles hesperus, n. ssp., with a discussion of the H. liponyssoides complex (Acarina; Haemogamasidae ) . J. Para.sitol.. 46(4) :401-409. Radovsky, F. J., and D. P. Furman. 1963. The Nortli American species of Steatonyssus (Acarina: Der- m;mvssidae). Ann. Entomol. Soc. Amer., 56(3): 268-276. Randolph. N. M.. and R. B. Eads. 1946. Ectoparasitic survey of mammals from Lavaca County, Texas. Ann. Entomol. Soc. Amer., 39:597-601. Rapp, W. F. 1962. Distributional notes on parasitic mites. Acarologia, 4:31-33. Rudnick, A. 1960. A revision of the mites of the family Spintumicidae (Acarina). Univ. California Pub. Entomol., 17(2) : 157-284. Seddon. H. R. 1951. Disea.ses of domestic animals in Australia. Part 3. Tick ;uid mite infestations. Div. Veterinary Hyg., Dep. Public Health, Common- wealth of Australia. Strandtmann, R. W. 1946. Atricholuctups sigmodoni, a new species of mite parasitic on the cotton rat, and notes on the genera Atricholaetaps and Ischno- laelaps (Acarina: Laelaptidae). J. Parasitol., 32(2): 164-169. Strandtmann, R. W. 1949. The blood-sucking mites of the genus Haemolaelups (Acarina: Laelaptidae) in the United States. J. Parasitol., .35(3) :325-352. Strandtmann, R. W., and D. M. Allred. 1956. Mites of the genus Brevisterna Keegan. 1949 (Aciuina - Haemogamasidae). J. Kansas Entomol. Soc., 29(3): 113-132. Strandtmann, R. W., and O. E. Hunt. 1951. Two new species of Macronyssidae, with notes on some estab- lished genera (Acarina). J. Parasitol., 37(5) :460- 470. Strandtmann, R. W., and H. B. Morlan. 1953. A new species of Hirstionyssus and a key to the known species of the world ( Dermanyssidae). Tex;Ls Rep. Biol, and Med., 11(4 ) :627-637. Strandtmann. R. W., and G. W. Wharton. 1958. Manual of mesostigmatid mites. Univ. Maryland, College Park. Contrib. no. 4. Talley, G. M. 1957. The incidence of na,sal mites in over-wintering Red-wing Blackbirds in the vicinity of Utiili Liike, Utah. Unpub. master's thesLs. Brig- ham Young Univ., Provo, Utah. Thurman, D. C, Jr., J. A. Mulrennan, and N. Bnuich. 1949. Description of the male of Cosmohielaps guraheiviis Fox (Acarina: Laelaptidae). J. P;u-a- sitol., .35(5):496-499. Till, W. M. 1963. Ethiopian mites of the genus AndroheUips. Bull. British Mus. Nat. Hist., 10(1): 1-104. Tipton, V. J. 1960. The genus Lcielups with a review of the Laelaptinae and a new Subfamily Alpha- laelaptinae (Ac;irina: Laelaptidae). Univ. California Pub. Entomol., 16( 6) :233-356. Traub, R., and M. Nadchatram. 1966. A revision of the genus Chatiii Brennan, with svnonymic notes and descriptions of two new species from Pakistan. J. Med. Entomol.. 2( 4) :.37.3-383. Wharton, G. W., J. M. Brennan. D. W. Jenkins, H. S. Fuller, G. M. Kohls, and C. B. PhiUp. 1951. The terminology and classification of Trombiculid mites (Acarina: Trombiculidae). |. Parasitol.. 37(1):13- 31. 70 Brigham Young IInivebmtv Science Bulletin Wharton, G. W., and H. S. Fuller. 1952. A Manual of the Chiggers. Entomol. Soc. Wa.shington Mem- oir No. 4. Whitaker. J. O., Jr. 1963. Food, habitat and para- .sites of the Woodland Jumping Mouse in central New York. J. Mammal., 44:316-321. Woodbury, A. M. 1956a. Ecological cheek lists. The Great Salt Lake Desert series, Uni\'. Utah. Dugway. Woodbury, A. M. 1956b. Host-piu-asite correlation charts. The Great Salt Lake Desert series, Univ. I'tah. Dugwav ( mimeographed series, unpulilished data ) . Yunker. C. E. 19.58. The parasitic mites of A/i/ofi.s lucifugus ( Le Gonte). Proc. Helminthol. Soc. Wash- ington, 25(l):31-34. Mites of Utah Mammals 71 Figs. 7-11. 7. Pach)laflaptidac ventral; 8, Purmpinturnix globus ventral; 9, Radfordia subliger dorsal; 10, Radfordm Icmnina dorsal; 11, Sjiinturnix orri, vuntral. 72 Brigham Young University Science Bulletin 24, thomomijs; 25, forus; 26, neofomnc. Mites of Utah NlANtMALS 73 Figs. 27-41. Dorsal plates. 27, Dermanyssus heckii 28, D. sanguinetis; 29, D. gallitiae; 30, Omithomjssus sijl- vuirum: 31, O. sylviarum variant; 32, Steatomjs.'ius antrozoi anterior plate; 33, Hirstiomjssus hilli; 34, H. isabel. linus; .3.5. S. antrozoi posterior plate; .36, H. hisetosus; 37, O. aridus; 38, H. triacanthus; 39, H. incomptus; 40, IchoTonyssus rohu.stipes; 41, H. hilli variant. Bhigham Young University Science Bulletin Fitrs 4" 53 Dorsal plates 42 hch/ropoda annatus; 43, Brcvi,tcnu. monlanus; 44. B^ utahcu.ps: f ■""<'"">- Figs. 4„-5J. uorsal plates^ 7 ' „„, J/,, ;,.li„r;. 47 / furmuni.- 48, H. lipomjssoides; 49, H. flmb.Jflfw form B; 50, pormsus alaskensis; 46, LumpUcUa oaken, •*(, j. /i""' > / j „„),„;„„. fnmi n H. ,;orvf>ger; 51, H. amhulans form A; 52, 11. amhulans form C; 53, H. am/;.iian. form U. MiTF.s OF Utah NTamm u s 75 65, Hiicmolachps ciisalis:; 66, //. gimgoui. ' . ■ • crouct; 63, £, cirruliins: 64 Brigham Young University Science Bulletin KiPs 67-77 Dorsal plates 67 Dcnmnupsus hccki; 68, Hir.v(io^,/.v.s.^s Imctosus- 69, Uchyropoda armatus; 70, H. g//i.sgoiii; 76, llirstiomjssus torus; 77. H. neotomac. \lni:s OF Utah Mwimaia 77 Figs. 78-92. Dorsal platfs. 78. Brcviaterna utahcnsis; 79, H aemogamams amhuhms form B; 80. Hirstionyssus tlwiiwjuijs: 81. H. nrotonmc; 82, H. ncotomr(tchiilaclaps circiilriris; 98. Haemolaelaps glasgowi. 99, RadfordUi hacJiai dorsal. Anal plate,s: 100, Ischtjropoda armtitus; 101, D. hccki; 1G2. Haemogamastis (daskensia; 103, Haemolaelaps casalis; 104, H. glasgoui: 105, E. circidari.'i. Mites of Utah Mammals 79 Figs. 106-121. Anal plates. 106, Uchyropoda armatus; 107, Haemogamasus Hpomjssoidc.s; 108, hchyropoda furmani; 109, Brrvi.sternu utahensis; 110, H. alaskemis; 111, Myomjssiis muntanus; 112. H. amhulans form B; 113, Macrochclcs sp.; 114, H. amhulans form C; 115, H. pontiger: 116. H. ambtilam form A; 117. H. amhulans form D: 118, B. mnntanus; 119, H. hngitarsus^ 120. Zumptiella hakeri; 121. Eulaclaps stahularis. 80 Brigham Young University Science Bulletin Figs. 122-135. Anal plates of Hlrstionussus. 122, stafiordi; 123, tar^alj,; 124. a/fini.,^ 125 punc^af^^^^^ chelae; 127, invofiumtus variant; 128, eutamiae; 129, utahensis; 130, ,nvagtnatus; 131, pa/u^t™, 132, angustus, 133, f/iomomi/,?; 134, toru.s; 135, femuralis. Mites of Utah Mammals 81 Figs. 136-148. Anal plate.s. 136. Hirstiomjssus neotomae; 137, //. hisctosus; 138, H. isaheUinus; 139, H. hilli; 140, //. hilli variant; 141. //. triacanthus; 142, H. incomptua; 143, Ichorontjssus rohustipes: 144. Dermanysstts sanguineus; 145. D. gallinae; 146, Ornithonyssus hacoti; 147, Spinturnix orri; 148, Paraspinturnix gloho.^is. 82 Brigham Young Univehsity Science Bulletin 159 160 161 Figs 150 Steatont/ssuf,- uutrozoi; 151, Oniithomjssu!,- syliiarutn; iaz u anau.s- .o^, ^«..«p, ....^.......o, ..... L. kochi; 155, L. wcills; 156. Kleenuwic sp.; 157. EuWc/iy- k/kpVcroL; 158;AnJroL./«,«/«,c./..,.- 159, L. n„»„/;,i,- 160. £. hollistcri; 161. Hj/,,oa.p« g«r«/,rn....- 162. Hiwmoladaps geoniys: 163. E. dcbitis. 149-16,3. Anal plates. 149. Dcrmtinyssus hcchi. 152. O. aridus; 153. Laelups multispinnsus; 154. Mites of Utah Mammals 83 Figs. 164-178. Anal platos. 164, 165, Euliracht/laehips circulars; 166. Hijpoaspis hihrica; 167, Haemolaehps glas- goui; 168, H. cdsatU-. 169, Ischijropnda furniani; 170, I. armatus; 171. Ichoroni/ssus rohtistipes; 172, Laelaps kochi: 173, H. g/a.vr;ou(,- 174, Dcrmany.s\m.s I>cchi; 175. L. mulfispinosus; 176, Omithonyssus hacoti; ni , E. circularis: 178, L. koclii. 84 Bricham Young Univebsity Science Bulletin Fi« 179 191 Anal plates 179 Uch,,ropoda furmani; 180, Laelups muhispiuosu.: WLHtrsUonyssus "eoW'mae; ^ mo B ;..i,t / ^^u...;.,- 183. H.iMinus; 184, H. bisetosus; 185. H. for,,.; 186, H. ,ncomptus: 187 . H. Mites of Utah Mamnials 85 ^'-'' ri!f;f^f^'Qr'lrP/'"T,'^r Parasiticlae; 193, Phytoseiidae; 194, Gamasolaelaptidae. Ventral setae ar- rangement: 195, Eubwchijlaelaps crowei; 196, E. dehilis; 197. £. hoUhtcri: 198, Ecirculark. 86 Bhigham ■^oung University Science Bulletin 199 Fies 199-207 Sternal plates 199 Macrochch-s sp.; 200. ZumptUlla hakcri: 201, Brrvhtcrrui montanus; 202, Haemogamasus amhulans form A; 203, Euhelaps stahulam; 204, H. pontijier; 205. B. utahemis; 206, H. lipotit/fimides; 207, H. longHarsus. Mites of Utah Mammals 87 igs. 208-215. Stemal plates: 208, Ischyropoda furmani; 210, Haemogamasus amhulans form C; 211, H. alasken- sis; 212. H. amhulans form B; 213, Hirstkmyssiis staffordi; 215, H. tar.ialU: Gcnitoventral plates-' 209 Zumn- ticlhi hakrri: 214 1 f„rnu,r,s ^ ' ' Brigham Young University Science Bulletin 216 217 221 219 220 222 FiBs 216-224 Sternal plates: 217, hchi/wpoda armntus; 218, Haemonamusus amhulans form D; 2\9, Myonyssus montanus; 221, Hirstiomissus affinh; 222, H. palustm; 224, //. punclatus. Genitoventral plate-s: 216. Brev,- sterna xitahensis; 220. Uaemogamasus liponyssoides, 223. Macrochelcs sp. NtiTEs OF Utah Mammals 225 Figs. 225-234. Sternal plates: 225, Hirstiom/ssus invaginatus; 227, H. invaginatiis variant; 228, H. eutamiae; 229, H. utahensis- 230, //. thomomys; 2.32. H. femuralis; 233, H. angustits; 234, H. longichelae. Genitoventral plates: 226, Hcicmogiimasii.s alaskensis; 231, Brevistcrna montanu.':. 90 Bhic.ham Young University Science Bulletin 245 Fi,s, 235-247. Sternal plates: 235. ^ ^rsHo,n>s.us torus 2^^^ Hjeolo^^^ Euhelaps stabtilaris. Mites of Utah Mammals 91 252 253 256 Figs. 248-256. Sternal plates: 248 Icltoroni/ssus robusfipes: 2.50, Dernianyssus sanguineus; 251. D. hecki; 253, Steatonyssus antrozoi; 2.54, OrnitJuiniissiis bacoti: 2.56. O. aridus. Genito\eiitraI plates: 249. Haenipgamasiis ambulans form A; 252, H. ambulans form B; 255, Mi/oni/ssus montanus. 92 Brigham Young UNn-EBsixY Science Bulletin 259 263 Figs. 257-265. Sternal plates: 257, Ornithonyssus sylviarum; 2.59, O. sijlviarum variant; 260 AndwUielaps levi- culus; 262, Kleemanin sp.; 263, Laelaps multispinostts; 265, L. kochi. Genitoventral plates: 258 Hirsfionussus punctatus; 261, H. palustris; 264, H. utahemis. ' Mites of Utah Mammals 93 272 Figs. 266-274. Sternal plates: 266, Laelaps inciluv 268, L. nuttallii; 269, Euhrachylaekps crowei; 271, E. hol- listeri; 111, E. circulars; 21 A, Hijpoaspvi lubrica. Genitoventnil plates: 267, Hirstkmysstis imaginatus variant; 270, H. tarsalis; 273, Haemogamasus ambulans form C. Bricham Young University Science Bulletin 275 276 278 277 282 281 283 Figs. 275-283. Sternal plates: 275, EtthrachylaeUips dehilis; 277, Ihjpoaspls gumbcmh; 278, H aemoladaps cas- alis; 280, H. ghsgowi; 281, Paraspinturnix globosus; 283, Spintumix nrri. Gt-nitoventral plates- 276 Hirst- ionyssus invagiiuitus; 279, H. eutamiac; 282, H. staffordi. Mites of Utah Mammals 95 284 285 288 289 290 291 292 295 Figs. 284-295. Genitoventral plates; 284. Haeviogamastis amhuJans foim D; 286. Hirstiont/ssus affinis; 287. Haemogiimasus longitarsus; 288, Hirstionyssus fcmurcilis; 289, H. thomomijs; 290, H. longichclac; 291, H. torus; 292, H. neotomae; 293, H. incomptus; 294. H. angu.sfi/.v; 295, //. i.mhellirius. 285, sternal plate of Haemohelaps geomijs. 96 Brigham Young University Science Bulletin 301 302 305 298 306 307 308 Figs. 296-308. Genitoventral plates. 296, Hirstionyssus triacanthus: 297, H. hilli; 298, H. hilli vari;uit; 299. H. bisetosus; 300, Ichoronyssus robustipes; 301, Dermanyssiis sanguineus; 302, D. gallinae; 303, OrniOwnyssus bacoti; 304, O. sylviarum; 305, Androlaelaps leviculus; 306, Eubrachylaelaps crowd; 307, Laelaps ineilis; 308, D. becki. Mites of Utah Mammals 309 313 317 318 320 314 316 322 323 Fies 309-323 Genitoventral plates. 309, Laehps kochi; 310. L. multispinosttsi 311. Ornithonyssus syhwrum; 312, Steatonussus antrozoi; 313, Hypoa.y,U gurabensis; 314, O. aridus; 315, L. nuttdli; 3\&.Haemoyelaps geomys; 317, Hypoaspis lubrica; 318, Haemolaehps casalis; 319, Kleemania sp.; 320, Eubrachylaekps hoUtsten; 321, E. debilis; 322, E. circularis; 323, Haemolaelaps glasgowi. Brigham Young Unhersity Science Bulletin . r. ■ > 1 1 . lOA 'ininturnix OTTV 325 Paraspinturnix ghhostis. Stemogenital plates: 326. FiBs 324-334. Genitoventral plates: 324, bpmturntx orn, o-j, / f„,J^R. -xoa H alaskensis- 330. Brevi- tionyssus utahensis. Mites of Utah Mammals 344 Figs. 335-346. Stemogenital plates. 335, Hirstionyssus torus; 336, H. thomomys; 337, H incomptus; 338, H. hisetosi,s- 339 H isahellinns; 340, Laelaps kochi; 341, Euhmchyhelaps circulark; 342 Dennanyssus beck:; 343 L multispinosus: 344, Haemolaelaps glmgowi; 345, D. ^lecfci; 346, Ornif/ionyssus bacofi. Brigham Young UNrvEHsiTV Science Bulletin 355 Figs. 347-358. Sternogenital plates: 347, Laelaps kochi; 348, L. multlipinos^is; 349, Euhrachylaelaps circularis: 350, Haemohelaps glasgowi. Stemo-genitoventral plates: 351, Ischyropoda armatus; 353, /. furmani. Holo- ventral plates: 352, Haemogamasus ambulans form B; 354, H. pontiger; 355, Hirstionyssus utahemis; 356, H. torus; 357, H. femttralis; 358, H. neotomae variant B. Mites of Utah Mammals 360 364 365 Figs. 359-367^ Holovcntral plates: 359. Brevu^terrm utahemh; 360, Hirstionysms thomomys; 361 H neotomae variant C; 363, H. neotomae variant A; 364, H. mcomptus; 365. Eubmchylaelaps circularis; 366, H. bisetosus- 3b7. Dermanysstis becki. 362, stemogenital plate of Ichomnyssus rohustipes. 102 Brigham Young Univehsity Science Bulletin 369 Fi.s 368-372. Holoventral plates: 368, Laelups .,ultispino... 369, Ornitho.yssus oridus; 371. Haemolaelap. ' glasgowi; 372, O. hacoH. 370, stemo-genitoventral plate of L. kocln. Mites of Utah Mammals 103 375 378 379 381 Figs. 373-381. Scuta: 373, Leeuwenhoekia americana: 375, Odontacarus micheneri, 376, O. Unsdalei; 378, O. hirsutus; 379, Whartonia perplcxa; 381. Tromhkula hclkini. Chelicerae: 374, Ischyropoda arm'atus; 'sil, Haemogamasus Uponyssoides; 380, H. amhulans form B. 104 Brigham Young University Science Bulletin variant; 389, Dermanyssus hecki. Mites of Utah Mammals 105 392 393 J >\'I : 395 394 397 Figs. 391-397. Scuta: 391, Tromhicula potosina; 392, T. suhsignata; 393, T. urihari^ 394, 7'. harperi; 39.5. T. jewetti; 397. T. esoeusis. 396, chelicera of H aemo gamasus ambttlans form D. 106 Bhigham Vounc University Science Bulletin 404 Figs. 398-405. Scuta: 398, TwmhKula sarcentu 399, T. /,a/.tri, 40-, /. '""'"""'""Z,' . ,,^„ „, „„-^„,, doremi; 405, T. allredi. CheUcerae; 400, Haemogamcu^us amhuhm form C; 401 Ormtiwnyssus anm Mites of Utah Mammals 107 F.gs. 406-416. Scuta: AO^ Tromhicua arenicoh; 408, Euscfwenga.,tui radfordi; 409, £, criceticola^ 411 £ rotun- 4l'h F?; t "''Z\V: ^'"""^'■416. E. pomerantzi. 407, Macrocheles sp. specialized seta of palpal tarsus; 410, Eulaelops mbuhr^ metapodal plate; 412, Ichoronyssus whustipes femur IV, 415, Omithonys^J syhiarum 108 Bbigham Young University Science Bulletin 417 420 423 424 Figs. 417-425. Scuta: 417, Euschocngastia lanceolata; 419, E. land; 420, E. luteodema; 422, E. sciuricola; 423, E. cynomijicola; 425, E. soricinus. Palpal trochanters: 418, Ichoronyssus robustipes; 421, Steatonyssus antro- zoi. 424. Ht/ponspis gurabensis body seta. Mites of Utah Mammals 109 432 Figs. 426-434 Scuta: 426, Euschoengastia utahensis; 428, E. oregonensis; 429. E. cordiremus; 430 E. lacerta- 431, Edecipiert,; 432, E. furmarU; 433, E. hoffmannae. 427. Ornithomjssus aridus palpal trochanter- 434' Hacmolaetaps glasgowi specialized seta on cheliceral digit. 110 Bricham Young University Science Bulletin Figs. 435-449. Coxae II of Hirstionyssus: 438, incomptus; 439, hilli variant; 440, hilli; 441, isabelUnus; 442, tria- canthus; 443, bisetosus; AAA, neotomae; 445, torus; 446, thomomys; 447, longichelae; 448, jemuralis; 449, angustus. Scuta: 435, hernia marita; 437, Cahrlepia americana. 436, H. staff ordi coxa I. Mites of Utah Mammals 111 Figs. 450-467. Co.vae II of Hirstionyssus: 450, utahensis; 451, eutamiae; 452, invagimtus variant- 453 invae tus; 454, punctatus; 455, palustris; 456, affinis; 457, staff ordi; 459, tarsalis. 458, hchyropoda armatus c^.. II. Coxae III of Hirstionyssus: 460, staffordi; 461, tarsalis; 462, a//mi.?,- 463, palustris; 464 moao,no o o 37 ARIZONA Fig, 615. Map of Utah showing counties and division into Great Basin and Colorado River Basin. ^( Brigham Young University Science Bulletin RHYNCHOPHORA BEETLES OF THE NEVADA TEST SITE by VASCO M. TANNER s^tC 3 0 1956 UNIVERSITY BIOLOGICAL SERIES — VOLUME VIII, NUMBER 2 ^ NOVEMBER, 1966 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Dorald M. Aixked, Department of Zoology and Entomology, Brigham Young University, Provo, Utah Associate Editor: Eabl M. Chbistensen, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: J. V. Beck, Bacteriology C. Lynn Haywabd, Zoology W. Derby Laws, Agronomy How^AED C. Stutz, Botany Wblmer W. Tanner, Zoology, Chairman of the Board Stanley Welsh, Botany Ex officio Members: RUDGER H. ^ Science Ernest L. Olson, Chauman, University Pubhcations RuDGER H. Walker, Dean, College of Biological and Agricultural Sciences The Brigham Youtng University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Pubhcations, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah. Brigham Young University Science Bulletin RHYNCHOPHORA BEETLES OF THE NEVADA TEST SITE by VASCO M. TANNER BIOLOGICAL SERIES — VOLUME VIII, NUMBER 2 NOVEMBER, 1966 LIBRARY DEC 30 1966 UNIVERSITY TABLE OF CONTENTS Page INTRODUCTION 1 ACKNOWLEDGMENTS 2 LIST OF THE SPECIES 2 SYSTEMATIC AND ECOLOGICAL DISCUSSION 7 Trigonorhinus irregularis 7 Thinoxenus nevadensis 7 Thricolepis inortuita 8 Eucyllus vagans 9 E. unicolor 9 Aragnomus hispidulus 10 Cryptolepidus leechi 11 C. nevadicus 11 C. aridus 11 C. cazieri 12 Ophrtjastes varius 13 O. gemirmtus 14 Orimodema protracta 14 Paracimbocera atra 15 P. artemisiae 15 Miloderes mercury ensis 16 Dirotognathus sordidus 17 Cleonus denticollis 17 C. lobigerinus 17 Smicromjx imbricatus '. 18 Smicrouyx sp 18 Promecotarsus densus 19 Apion varicome . 19 Auletobius humeralis 19 Auletobius sp 20 Myrmex lineata lineata 20 Magdalis Iccontei tenebrosa , 21 Macrorhoptus hispidus 22 Anthonomus peninsularis 23 A. haematO])us confusus 23 A. hirtus 23 A. omatulus 24 A. sphaeralceae 24 A. cycliferus 24 A. tenuis 24 Epimechus gracilis 25 Brachyogmus ortuttus 25 Tychius prolixus 26 Zascelis irrorata 26 Ceutorhynclius adjunctus 27 C. tescorum 27 Onychobaris mystica 27 O. near depressa 28 Yuccaborus frontalis 29 NOTES ON THE BIOLOGY OF SOME NEVADA TEST SITE WEEVILS 29 REFERENCES 31 LIST OF ILLUSTRATIONS Figure Page L Cidciulru ochrctis, general morphology 3 2. Trigonorhinus irregularis, dorsal view 7 3. Thinoxenus nevadensis, dorsal view 8 4. EiicijUus Vdgans, dorsal view 9 5. Cnjptolcpidus aridus, dorsal view 12 6. Cnjptolepidus aridus, female genitalia 12 7. Cnjptolepidus aridus, spermatheca 12 S. Mitodcres mercuryensis, dorsal view 16 9. Miloderes mercuryensis, aedeagus 16 10. Magdalis lecontei tenebrosa 21 11. Map of the areas of the Nevada Test Site 33 12. Extent of the major plant communities of die Nevada Test Site 34 LIST OF TAXONOMIC KEYS Page Families of Rhynchophora 4 Subfamilies of Anthribidae 4 Subfamilies of Curculionidae 4 Species of Eucyllus 8 Species of Cryptolepidus 10 Species of Ophryastes 13 Species of Paracimbocera 15 Species of Miloderes 15 Species of Promecotarsus 19 Genera of Myrmecinae 20 Genera of Anthonomini 21 Species of Macrorhoptus 22 Subgenera of Anthonomus 23 Species of Zascelis 26 Species of Yuccahorus 28 RHYNCHOPHORA BEETLES OF THE NEVADA TEST SITE' by Vasco M. Tanner" INTRODUCTION It is interesting to contemplate the progress that is taking place in the accumulation of in- formation and imderstanding of the large and interesting group of snout beetles, the Rhyncho- phora. Following the major contributions made by LeConte, Horn, Say, Casey, Blatchley, Leng, Fall, Schaeffer, and Dietz, there was a decided lull and lack of interest in the study of the wee- vils. Within the past 30 vears, however, many new workers have been attracted to the study of the Curculionidae of this country. Attention has been turned to careful curating and amassing in permanent collections of the old and newly col- lected specimens of Nearctic species. Major con- tributions by recent workers which must be con- sulted when studying this insect complex have been made bv such authorities, to name a few, as Pierce, Van Dyke, Buchannan, Ting, Sleeper, Warner (Mrs. Spilman), Kissinger, Burk, Gilbert, and Anderson. Two timely and useful recent contributions are Amett's "The Beetles of the United States, A Manual for Identification," and Kissinger's "Cur- culionidae of America North of Mexico, A Key to the Genera 1964." These bring together and make available information on the weevils that will greatly facilitate research work on this difficult group. There is great need for careful internal as well as external morphological studies of the weevil genera. Without doubt, the more than 3,600 genera can be reduced in number and a better understanding of their phvlogenv brought to light. Too little is known about the immature stages and food habits of the United States weevils. It is the aim of this study to classify those collected at the Nevada Test Site (Fig. 11) according to latest available studies and to report their seasonal abundance and plant hosts. No special effort was made to collect Rhynchophora; only specimens that were trapped along with other insects or were occasionally taken from plants came into this collection. The collecting extended from autumn, 1959 to and including the summer of 1965. A total of 310 specimens representing 28 genera and 44 species was col- lected. The fauna of this southern portion of the Great Basin is tvpicalh Sonoran in its insect ele- ments. The summers are iiot and long; whereas the winters are mild and short. Allred, Beck, and Jorgensen (1963a) reported temperatures of 112° F in July. Van Dyke (1939) pointed out that the Sonoran fauna is a peculiar one, well adapted to its environment. The dominant gi'oups of in- sects in this fauna are such coleopterous families as the Tenebrionidae or darkling beetles (Tan- ner and Packham, 1965), the Meloidae or blister beetles, and somewhat related Alleculidae and Oedemeridae, and a certain large element of broad-nosed weevils like the Brachyrhinae (Otorhvnchinae) as well as numerous genera of Buprestidae or jewel beetles. Other dominant insects than the beetles which favor these areas are the Bombyliidae or bee flies (Allred, Johnson, and Beck, 1965) and the Asilidae or robber flies among the Diptera. The great portion of the adult beetles are nocturnal, burving themselves in the soil during the day, and most of their larvae are subterranean in habit. Many of the nocturnal forms are somber in color or with very little color, the diurnal forms often gray and many of them very hairy, as is true of many desert plants. Though so very distinct and highly modified, I am convinced that most of this fauna has been derived from the Neotropical or South American fauna. Species of the tribe Anthono- mini which live mainlv in the flowers, fruits, and seeds of plants are well represented. The plant communities of the test site were deteiTnined and discussed by Allred, Beck, and Jorgensen (1963a, 1963b). They used t^^'o major tvpes of vegetation at the test site which they divided into plant communities (Fig. 12). The Desert Scrub tvpe was divided into the Larrea- Franseria, Gravia-Lycium, Coleogyne, Atriplex- Kochia, and Salsola communities. The Desert Woodland t\^:>e was designated as one commun- ity, the Pinyon-Juniper. Some areas were not typical of the above plant communities, and iReport B,Y,U-A,EC. COO-1 355-U. Field work 2Departnient of Zoology and Entomology, Brigha npleted under AEC Young University, Pr ATi III )--.Sl. and .Vrdl-ll-l U Brigham Young University Science Bulletin Allred et al. proposed that mountain areas, natur- al springs, reservoirs, and playas be grouped to- gether as mixed communities. In this study the weevil species are discussed under plant relationships, and the number of specimens collected, month and year of collec- tion, and plant host or plant communities where collected are given. Specimens taken in can traps are related only to the plant community where trapped. This may not be entirelv accurate since it does not account for the flight or movement of the specimens from contiguous areas. Very little information, if any, on the life history of species of this study was obtained. ACKNOWLEDGMENTS I am indebted to many individuals who were associated with the collecting, preparing, and fur- nishing of ecological information on the weevils from the Nevada Test Site. Tins study was made possible by a grant to Brigham Young University from the Atomic Energy Commission, contract AT(ll-l)-786. Drs. Dorald M. Allred and D Elden Beck, project directors. Department of Zoology and Entomology of Brigham Young Ll^ni- versity, have been most active in gathering data as outlined by the grant and very successful in getting the findings published. I acknowledge and thank the Atomic Energy Commission and Drs. Allred and Beck for financial support, facili- ties, and many kindnesses during this study. Appreciation is also extended to Drs. H. R. Burk, Department of Entomology, Te.xas A & M University, who kindly furnished liis opinion on some species of Antlionomini, and Harold J. Grant, Academy of Natural Science of Pliiladel- phia, who made it possible for the author to com- pare and study many species of the Mason weevil collection at the academy. Mrs. Rose Ella Spilman, U.S. National Museum, and Mr. Hugh B. Leech, California Academy of Sciences, were helpful in providing loans and comments on specimens submitted to them. I also thank the preparators who, under the supervision of Drs. Allred and Beck, were most courteous and help- ful in providing collection data and sifting out ecological information, and Mr. Douglas Hill of Brigham Young Uni\'ersity who drew some of the illustrations used in this study. LIST OF THE SPECIES Two families of the Rhynchophora are repre- sented. In the main I have followed Kissinger's (1964) taxonomic proposals as to subfamilies, tribes, and genera of the Curculionidae. He made a number of major clianges in the subfamily and tribal sequence. Tliis departure from the Leng Catalogue and Bradley Manual arrangement of higher categories will no doubt, if followed, be reflected in a new catalogue of the North Ameri- can Coleoptera. I agree with the way Kissinger has dealt with the Otiorhynchinae of Leng. Tlie new arrangement of the genera is far more natur- al and morphologically sound. I have made use of the keys and of the genus and species characterizations of previous contri- butors to our knowledge of the species included in this study. Tlie labeled drawing of Calendra ochrens Lee. ( Fig. 1 ) may be useful in inter- preting the moqjhojogical tenns used in the keys and descriptions. Family Platystomid.\e ( Anthhibid.ae) Subfamily Anthribinae Tribe Anthribini Trigonurhiniis hrcg^ularis (Tanner) Family Curculionidae Subfamily Brachyrhininae Tribe Peritelini Thinoxcnus nevadensis Casey Thricolepis inoinata Horn EucijUits D(inomii.s hispidiihts Casey Subfamily Tlivlacitinae Tribe Banaotini Cn/pfolcpidn.s Ircclii Ting Weevils of the Nevada Test Site Occiput EpimeralAbea Epipleura T PROSTERNUM I i..^^Lr,.^r T EpisternalArea Int^erspace Apex Proncttum Vertex Front Compound Eye Gena Groove Club Beak- mandibles Epimehum -Claws Fig. 1. Calendra ochreus, general morphology Tarsus Claws Crijptolcpidiis nevadicus (Van Dyke) Crtjptolcpidus aridtis, new species Cnjptolepidus cazieri (Van Dyke) Subfamily Leptopiinae Tribe Ophryastini Ophryastes variiis (LeConte) Ophryastes geminatus Horn Tribe Leptopiini Orimodema protracta Horn Paracimhoccra atra Van Dyke Paracimhocera artcnmiae Ting Miloderes mercuryensis, new species Dirotognathus sordidus Horn Subfamily Cleoninae Cleonus denticollis Casey Cleomis lohigerinus Casey Subfamily Erirhininae Tribe Smicronychini Smicronyx inibricatus (Casey) Smicronyx sp. Proinecotarsus densus Casey Subfamily Apioninae Apion vdricorne Smith Subfamily Rhynchitinae Auletobius htimercdi^ Boheman Auletobius sp. Subfamily Myrmecinae Myrmex lineata ( Pascoe ) Subfamily Magdalinae Tribe Magdalini Magdalis lecontei subsp. tenehrosa Fall Subfamliy Anthonominae Tribe Anthonomini Macrorhoptus hispidus Dietz Anthonomus peninsularis Dietz Anthonomus haematopus si.bsp. confiisus Dietz Anthonomus ornatulus Dietz Anthonomus hirtiis LeConte Anthonomus spluicralccae Fall Anthonomus cyclifcrus Fall Anthonomus tenuis Fall Epimechus gracilis Fall Brachi/ogmus ornatus Linell Subfamily Tvchiinae Tribe Tychiini Tychius jnolixus Casey Siibfamilv Cryptorhyncliinae Tribe Cryptorhynchini Zascelis irrorata LeConte Subfamily Ceutorhyiichinae Tribe Ceutorhynchini Ceutorhijnchus acljunctiis Dietz Ceutorhynchiis tcscorttm Fall Brigham Young University Science Bulletin Subfamilv B;u-idinae Tribe Madarini Ontjchoharis mystica Casey Omjchoharis near deprcs.sa Casey Sublamilv Rhvnchophorinae Tribe Sipalini Yi4ccahoni.s frontalis (LeConte) KEY TO THE FAMILIES OF RHYNCHOPHORA OF THE NE\ ADA TEST SITE Rostrum broad, flat, short, never separated from the front bv a transverse im- pression. Labrum present. Antennae usually inserted under the sides of the front, slender, straight; segments 9 to 11 broader, more or less compressed, rarely forming a compact club. Labial palpi of three segments, slender, long- er than in other Rhynchophora, the last segment of all palpi elongate. Base of pronotum always with a transverse elevated ridge. Pygidium exposed; notched to receive sutural ape.x of elytra. Middle coxal cavities broadh' closed by sterna. Tarsi brushlike beneath, of four segments; the second emarginate, tri- angular; the third small, bilobed Anthribidae Rostrum variable in length, often long and curved downwards. Palpi short, rigid, nearly always concealed within the mouth. Head usually spherical. Antennae generally geniculate with a three-segmented club, usuallv ringed. Front coxal cavities closed behind. Elytra usually with a strong fold on the underside, limiting a deep groove into which the upper edge of the abdomen fits. Epi- pleura wanting. Five stemites, the first two closely united. Front and middle co.xae rounded, the hind pair oval Curculionidae KEY' TO THE SUBFAMILIES OF ANTHRIBIDAE OF THE NEVADA TEST SITE (From Bradley, 1930, and Arnett, 1962) Antennae inserted on the sides of the rostrum (Brachytarsinae) Anthribinae Antennae inserted on the front of the head, or dorsally at the base of the rostrum. Pronotal ridge basal Choraginae Tribe Anthribini Since only one species of this family is included in this study, the tribe Anthribini to which it is assigned may be briefly characterized as follows: Antennae inserted on the sides of the rostrum; transverse ridge situated at the base, the surface behind it perpendicular. KEY TO THE SUBFAMILIES OF CURCULIONIDAE OF THE .\'E\'ADA TEST SITE (After Kissinger, 1964) 1. Trochanter long and somewhat cylindrical. Femur attached to apex of trochanter; base of femur distant from coxa. Antenna straight; funiculus with seven seg- ments, segments 6 and 7 distinctly narrower than club Apioninae Trochanter short and triangular. Femur attached to side of trochanter; base of femur closely adjacent to coxa 2 2. Antenna straight; rostrum generally lacking conspicuous scrobe to receive anten- na. Prothorax not margined; elytra concealing tergites 1-5 .. 3 Antenna distinctly elbowed between scape ;ind funicular segment 1; rostrum gen- erally uith apparent scrobe to receive scape of antenna. Tarsus with four dis- tinct segments. Funiculus consisting of five to eight segments 6 Weevils of the Nevada Test Site 5 3. Antenna with eleven segments, including distinct club composed of tliree seg- ments. Prothorax not pedunculate at base. Mandible flattened, toothed on outer margin. Claw with basal process Rhynchitinae Rostrum not received into prosternal emargination, or, prosternum not a triangular plate in front of procoxae and rostrum usually slender and longer than head or tarsal claws toothed at base. Tarsal segment 3 usually strongly bilobed 4 4. Mandible lacking deciduous cusp, not scarred, either glabrous on lateral aspect or with few minute setae; mandible usually small in size. Rostrum slender, gen- erally longer than prothorax; if rostrum shorter than prothorax then frons and rostrum lacking deep, median longitudinal groove, and scape not obviously extending bevond anterior margin of eye. Prothorax with margin more or less truncated, margin not produced into rounded lobe behind the eye; eye usual- ly distant from anterior margin of prothorax, slightlv round in outline. Pro- thorax lacking long setae projecting anteriorly from front margin adjacent to eye .-. ■ 5 Mandible with prominent scar indicating point of attachment of deciduous cusp, or else clothed laterally with many fine scales or setae; mandible large in size. Rostrum stout, quadrate in form, shorter than prothorax, rarely longer, often expanded laterally toward apex. Prothorax with anterior margin pro- duced into prominent, rounded lobe adjacent to eye; eye usually partly cov- ered by anterior margin of prothorax, mainly transverse in shape. Vlandible with four or more large setae Leptopiinae 5. Antennal scrobe vaguely defined posteriorly; scape usually passes over middle of eye when retracted ne.xt to head Brachyrhininae Antennal scrobe lateral, dorsal margin sharply defined, ventral and dorsal bound- aries strongly bent ventrally so that scape usually rests below eye when re- tracted next to head Thylacitinae 6. Scape not extending beyond hind margin of eye, generally fitting into scrobe; an- tenna generally inserted toward apex of rostrum; club uniformly clothed with erect, fine, short setae, first segment with distinct setae; if firsf segment is bare then funiculus with seven segments or prosternum with apical channel 7 Scape stout and long, projecting some distance past posterior margin of eye, not fitting into short antennal scrobe; inserted close to eye (exception to preced- ing statement: Yuccahonts); funiculus six segments, first segment of club large, virtually glabrous and polished. Large uncus on tibia 3 projects from anterior margin. Prosternum not channeled. Hind tarsal segment not bilobed (except Yticcahonis) . Pygidium of male resembles that of female Rhynchophorinae 7. Rostrum free, not received into deep, median sternal channel 8 Rostrum received into deep, median sternal channel in repose. Funiculus with six or seven segments. Eyes partially covered by prothoracic postocular lobes when rostrum is in repose. Pygidium covered by elytra. Tibia 1-3 armed with an uncus. Antennal club rather evenly clothed with pubescence. Tibia 3 with distinct apical comb of setae. Body lacking dense vamishlike coating. Tarsal segment .3 bilobed, usually distinctly wider than segments 1 and 2. Front coxae obviously separated by prosternum or claws toothed or connate Cryptorhynchinae 6 Brigham Young Univebsity Science Bulletin 8. Mesepimeron not ascending and not visible in dorsal view 10 Mesepimeron strongly ascending, truncated by elytral humeri and visible in dorsal view between prothorax and elytra; elytra not produced anteriorly over base of prothorax 9 9. Funiculus with seven segments. Claws simple, free, connate or with a single claw Baridinae Funiculus with six segments. Claw simple or toothed Ceutorhynchinae 10. Front coxae contiguous, not separated by process of prosternum 11 Front coxae separated by complete process of prosternum; separation may be slight. Sterna unequal in length, 2 about as long or longer than 3 and 4. Claws simple. Tibia 3 definitely uncinate, praemucro if present small, less than one-half as long as uncus. Eyes narrowly separated by frons which is not wider than maximum diameter of club. Femur 1 toothed, sometimes strongly so. Elytra broadly exposing pygidium; elytra black, usually with a reddish area, lacking obvious vestiture. Tooth on femiu- 1 small and acute. Rostrum nearly subcylindrical. Metepimeron visible Magdalinae 11. Suture between sterna 2 and 3 nearly straight, at most slightly produced back- wards laterally, not reaching suture between sterna 3 and 4 12 Suture between sterna 2 and 3 strongly produced backwards laterally, reaching or passing suture between sterna 3 and 4. Claw with basal process. Hind coxa distant from margin of elytra. Rostrum definitely tapered apically Tychiinae 12. Claws connate at base. Funiculus with seven segments. Elytron lacking acute subhumeral tubercle. Tibia lacking angulation on inner surface near middle 13 Claws free at base, simple, lacking basal process; sterna unequal in length. Front tibia with apex not produced into elongate, flat paddle; tibia 3 unarmed at apex or mucronate, mucro not more than one-half as long as claw; eyes well developed 14 13. Eye distant from anterior margin of prothorax; anterior margin of prothorax with long postocular vibrissae and not produced into postocular lobe. Vestiture fine, hairlike. Metepimeron visible, punctured and scaly at metepisternum Cleoninae Eye partially concealed by postocular lobe of anterior margin of prothorax; long postocular vibrissae absent. Rostrum obviously more slender than maximum width of femur 2, usually separated from frons by erect group of scales and transverse impression. Elytra usually with round or narrow scales. Eyes usual- ly not contiguous ventrally Erirhininae 14. Rostrum long and slender, more or less glabrous; width at insertion of antenna (in lateral view) much less than width of eye; rostrum longer than pro- thorax; eyes nearly round (subfamily in part) Anthonominae Rostrum stout, front coxae much closer to hind margin of prosternum than front margin, distance to front margin not less than two times as great as dis- tance to hind margin; prothorax longer than wide, globose in middle, base much narrower than middle; tibia 3 uncinate; frons narrower than diameter of antennal club; scrobe directed toward ventral aspect of rostrum; body nearly glabrous (subfamily in part) Myrmecinae Weevils of the Nevada Test Site SYSTEMATIC AND ECOLOGICAL DISCUSSION Fainilv Anthribidae Subfamily Anthribinae Tribe Anthribini Genus Trigonorhintis Wollaston Reference. Wollaston, 1861:102. Morphological characteristics. Genus Trigo- norhintis. Valentine (1960), a specialist on this family, pointed out that the generic character- istics of this genus may be recognized in all the species assigned to the genus ". . . by their ros- trum, the dorsal surface of which is progressive- ly narrowed from base to apex, and by the shape of the apex itself which has the central portion distinctly produced beyond the comers and slightly emarginate. . . . The lobes of the third tarsal segment are separate, thus this segment appears profoundly emarginate. . . . Species of tliis genus are plant feeders like the majority of the family." Trigonorhintis irregularis (Tanner) Fig. 2 References. Tanner, 1934:285-286. Valentine, 1960:41-85. Morphological characteristics. Form robust oval. Covered with loose whitish pubescence, except for brown checkering of prothorax and elytra. Head and rostrum blackish; mandible rufescent with whitish pubescence extending down over base of mandibles; labium distinct, brown and glabrous; eyes but slightly emargin- ate; scrobes reaching eye, antennal carina promi- ment, extending from upper margin of eye to base of mandible. Antennae rufescent, except segments of club which are brownish black; first two segments about equal; segments tliree to eight equal, but only about one-third as wide as Fig. 2. Trigoriorhinus irregularis, dorsal view first two; segments of club as long as segments tliree to eight. Prothorax wider than long; lateral carina extending forward only about one-third length of prothorax; decumbent coarse pubes- cence covers surface; distinct brownish spots on disc. Elvtra \\'ider at base than prothorax, with conspicuous bro%vn patches of pubescence sur- rounded bv white elongate scales. Legs rufescent and evenlv covered with white pubescence; claws widelv divergent with small tooth, about etjuidistant on claw. Under surface of the body covered with white scales. Length 1.9 mm, width 1.3 mm. Plant relationship. One specimen was col- lected in July, 1965, on Franscrio acanthicarpa in Area 17. Comments. This species was described orig- inally in the genus Brachi/tarsoidcs (Pierce, 1930) which was recently made a synonvm of Trigonorhintis (Wollaston, 1961) by Valentine (1960); hence, the new combination. In 19.54, Dethlefsen described several species of Trigonorhintis from California and Nevada. I have been unable to relate Trigonorhintis ir- regularis to any of the species discussed in his paper. Trigonorhintis nevailensis agrees to some e.xtent in color and pubescence, but it is much larger than T. irregtilaris. The species of this genus will no doubt be critically dealt with in the near future by Valentine, who is well pre- pared to do so. Family CuRCULiONmAE Subfamily Brachyrhininae Tribe Peritelini Genus Thinoxentis Horn Reference. Horn, 1876:74. Morphological characteristics. Genus Thino- xentis with mesopleural sclerites unequal, epis- temum larger and attaining elytra; first ventral suture arcuate; cotyloid surface of the hind tibiae squamose; scrobes lateral, not convergent, very shallow, not attaining the eyes; scape feebly ar- cuate. Thinoxentis nevadensis Casey Fig. 3 Reference. Casey, 1888:267. Morphological characteristics. Casey de- scribed Thinoxeinis iievadensis as an oblong- elongate convex species, clothed with smaU, rounded, dark brown scales; abdomen covered Bricham Young University Science Bulletin Fig. 3. Thinoxenus itevadensis, dorsal view with crust of similar scales, intermixed with squamiform hairs. Head and rostrum conical, flattened; rostnim with short median stria in basal half; eyes small, very distant on sides; scrobes widely distant, narrow and deep near apex; antennae inserted nearly at apices of scrobes; first joint of funicle one-half longer than second. Prothorax one-half wider than long, wider than head; disc not densely punctate. Elytra emarginate at base; sides nearly straight and parallel; feebly striate. First ventral suture strongly arcuate; second segment as long as ne.xt two together. Legs densely squamose, with short semi-erect hairs. Length 5 mm, width 3.0 mm. Plant relationship. One specimen was taken in a Pinyon-Juniper community, Area 12, in August, 1964. Genus Thricolepis Horn Reference. Horn, 1876:68). Morphological characteristics. This genus is common throughout the Great Basin region. It has the following characteristics according to Pierce (1909): side pieces of mesostemum un- equal, epistemum large and attaining elytral margin, epimeron usually small; first ventral sutin-e arcuate; cotyloid surface of hind tibiae glabrous; integument squamose; scrobes super- ior, convergent above; rostnim narrowed to tip, longer than head; scrobes very short, terminal. Thricolepis inornata Horn Reference. Horn, 1976:68-69. Morphological characteristics. Thricolepis in- ornata, one of the three species of this genus found on the oaks, is common along the Wasatch Front and in the Great Basin. It has the follow- ing distinguishing characteristics: form oval and robust, head and rostrum larger than thorax, surface sparsely punctured and not densely scaly. Antennae prominent, rufotestaceous. Pro- thorax cylindrical, slightly wider than long, mth deep punctures, sparsely scaly and with erect hairs. Elytra about three times as long as thorax, with closely placed punctures, intervals flat, short black erect setae, scales of pearly luster, and body surface black and shining. Length 3.6 mm, width 2 mm. Plant relationship. One specimen was taken from Quercus gamhelii. Area 12, in August, 1965. Genus Euci/llus Horn Reference. Horn, 1876:74. Morphological characteristics. Genus Eucyl- hm with first ventral suture arcuate, cotyloid sur- face of hind tibiae squamose; scape arcuate, slightly twisted; scrobes lateral, not convergent, deep, attaining the eyes. Specimens representing two species of this genus were collected at the test site. Van Dyke (1936:31-32) prepared the following key to separate the species of EtictjUus: Larger species, 5.5 to 7.5 mm in length; bicolored; second funicular segment from tliree to four times as long as broad; setae of entire upper surface hispid, many times as long as broad and acute at apices vagans Horn Smaller species, 5 mm or less in length; second funicular segment but little more than twice as long as broad; setae of upper surface very short and robust, squamose or club-like and blunt at apices Weevils of the Nevada Test Site 2. Bicolored, three brown stripes on pronotiim and numerous browm patches on elytra; erect setae of upper surface club-Hke, several times as long as broad echinus Van Dyke Unicolored, cinereous; erect setae of upper surface peg-like or tubercular; but little longer than broad tinicolor Van Dyke Eucylhts vagans Horn Fig. 4 Reference. Horn, 1876:75. Morphological characteristics. Eucijllus va- gans is an elongate oval species, densely clothed with scales and grayish setifonn hairs. Head and rostrum not as long as thora.x, covered with cinereous and pale-brownish scales intermi.xed, and short erect hairs; prothorax cylindrical, slightly wider than long, disc sparsely punctured, densely scaly and with some hairs. Elytra twice as long as wide, finely striate, fine punctures on the striae, intervals flat; long erect hairs on each interval. Venter of body scaly, with some short setae. Legs with cinereous scales; femora with dark ring near tip. Length 7 mm, width 3 mm. Plant relationships. A total of 42 specimens was collected, mostly during 1961 (Table 1). Table 1. Collection records of EucyUus vagans at the Nevada Test Site. Area Plant Host° or Community Month and Number of Specimens 1 5 6 Grayia-Lycium Larrea Lycium ML\ed AprU I. May 2 May 1. June 1, Sept. 2, Oct. 3, Nov. 3 Sept. 2, Nov. 1 Nov. 1 10 Coleogyne April 1, Aug. 1, Oct. 1, Nov. 1 12 16 Grayia-Lycium Pinyon-Juniper 'Eurotia lanata June 1 Aug. 1 June 1 23 Mixed Oct. 6, Nov. 6 28 Mixed Feb. 2, Oct. 1, Nov. 1 Cane Springs Mixed May 1, Oct. 1 Fig. 4. Eucijllus vagans, dorsal view EitcijUus tinicolor Van Dyke Reference. Van Dyke, 1936:32. Morphological characteristics. Euci/Uiis tini- color is a small species, unicolored, lacking browTi stripes common to other two species, cinereous with erect setae which are peg-like or tubercular. Prothorax with sides less broadly rounded, elytra narrower with striae deeper and with closer punctures. Length 4.9 to 5.1 mm, width 2 to 2.1 mm. Plant relationships. Four specimens were taken from a mixed community. Area 28, and two from Cane Springs (a mi.xed community), all in September and October, 196L Genus Aragnomtis Horn Reference. Horn, 1876:72. Morphological characteristics. Aragnomtis, a genus related to the genera Thricolepis, Etictjl- Itis, and Thinoxeniis. is distinctive because of an arcuate ventral suture. Cotyloid surface of liind tibiae glabrous; integument squamose; scrobes more lateral, not convergent above, and scape much shorter than flagellum. Bhigham Young University Science Bulletin Aragnomiis IiKpiclidus Casey Reference. Casey, 1888:266. Morphological characteristics. Aragnomus hispidulus is robust, covered with rounded, pale brown scales. Head wider than rostrum; rostrum longer than wide and dilated at tip. Antennal scrobes lateral, deep, nearly straight, but not reaching eyes; scape extending beyond anterior margin of prothorax, shorter than flagellum. Pro- thorax one-half wider than long, sides parallel, finely and sparsely punctate, each puncture with erect scale. Elytra slightly inflated, declivous at apex; one-third longer than wide, middle much wider than prothorax. Legs short and well cov- ered with short erect scales. Length 5.5 mm, width 2.7 mm. Plant relationships. Twenty-five specimens were collected, mostly during 1962 (Table 2). Subfamily Tliylacitinae Tribe Barynotini Genus Cryptolepidtis Van Dyke Reference. Van Dyke, 1936:91. Morphological characteristics. Kissinger (1964) placed Cnjptulepidiis in the subfamily Tliylacitinae which he characterized as follows: claws free; eyes in part encroaching upon dor- sal area of head, frons narrower than rostrum; Table 2. Collection records of Aragnomus hispidulus at the Nevada Test Site. Plant Host" or Community Montli and Number of Specimens 2 'Coleogyne ramosissima Jan. 1, April 1 4 "Grayia spinosa Nov. 1 5 "Larrea divaricata May 1 12 Pinyon-Juniper July 14, Aug. 7 frons and dorsal surface of rostrum lacking deep median sulcus, with at most a broad, shallow, longitudinal depression; rostrum separated from frons by distinct transverse furrow or depression; dorsal aspect of tarsal segments with scales, tarsi clothed ventrallv with coarse setae. Mentum concealing maxillae; frons lacking distinct tu- bercle above eye; eye visible in dorsal view; elytra with suberect, fine, acute setae. The genus Crijptolepidiis includes six species confined to southern California, Nevada, and Arizona. The main generic characters are no pubescence on ventral surface of tarsal segments, only bristle-like setae present. Third tarsal seg- ment only slightly, if at all, larger than second. Postocular prothoracic lobes and vibrissae ab- sent. Antennal scrobes deep and only slightly widened at posterior end. Four species are known for Nevada and the test site. The following key modified from Ting's (1940)) study will separate them. KEY TO SPECIES OK CRYPTOLEPIDUS 1. Elytral setae short and reclinate, not longer than one-fourth to one-half \\ddth of intervals; head slightly convex between eyes; pronotal sides faintly convex -- leechi Ting Elytral setae long and suberect, as long or longer than width of intervals; head between eyes flat or concave; pronotal sides strongly convex 2 2. Pronotum not tuberculate or rugose 3 Pronotum distinctly tuberculate and rugose 4 3. Base of rostrum with broad, somewhat arcuate, transverse constriction; pronotum as broad as long; elytral intervals flat; fore tibiae witli six spinules and pale colored mucro at apex, exclusive of those in ventral comb -"- nevadicus (Van Dyke) Base of rostrum with well-defined basal constriction, \\'ith median longitudinal groove free from scales; head one-third wider than rostrum at base; corbel spinules amber, widely spaced, six in anterior, eight in mesothoracic tibial comb; setae white to golden, long, three times width of elytral intervals, sparsely placed, none on medial discal area, short on head and rostrum; an- tennal club reddish-brown; scales silvery; size 4 to 4.5 mm aridiis new species Wkevils of the Nevada Test Site Base of rostruin flat; pronotiim one-fourth wider than long; elytral intervals slightly convex; fore tibiae with eleven spinules and black mucro at apex, exclusive of those in ventral comb planifions Ting 4. Dorsum of rostrum longitudinally convex between base and subapical area; med- ian sulcus narrow, line-like; color brown cazieri (Van Dyke) Dorsimi of rostrum longitudinally flat; median sulcus broad; color grey rugicoJUs Ting Cryptolefndtis Icechi Ting Reference. Ting, 1940:147. Morphologioal characteristics. Nloderately elongate, clothed with white, imbricate, circular and ovate scales; rostnmi with faint basal trans- verse constriction; narrow median sulcus; setae sparse and same length as those on head; sub- apical area with V-shaped glabrous region back of nasal plate. Head with area between eyes slightly convex. Prothorax with broad apical con- striction; broader than long; disc pimctate, but covered by crust of white scales; setae about same length as head setae. Elytral sides nearly parallel; intervals flat, striae fine, setae short. Spinules black, hind tibia with six teeth in distal comb and five teeth in anterior comb. Length 6.5 mm, width 3.0 mm. Plant relationships. Two specimens were taken, one in a Larrea community. Area 5, in March, 1961, and the other in a Lycium com- munity. Area 5, in February, 1961. Comments. Ting points out that the salient characters of C. Jeechi are its extremely short setal vestiture whose length on the elytra is one- fourth to one-third the inter\'al width; the shghtly convex front between the eyes; and the nearly black tarsal bristles and corbel spinules. Crtjptolepidus nevadicus (Van Dyke) Reference. \'an Dyke, 1936:77. Morphological characteristics. Cn/ptolepidtis nevadicus is reddish-brown to black with grav and irridescent scales. Tarsal bristles and tibial spinules pale yellow in main, but light brown in some specimens. Rostrum broad, with transverse constriction at base; median sulcus narrow, ex- tending from antennal articulation to posterior margin of basal transverse constriction. Pronot- Tim smooth, as broad as long, sides convex; long- er setae four times longer than head setae; no median sulcus. Elytral intervals flat; setae two and one-half times longer than interval width. Metepistemal suture visible at posterior end. Hind tibia with seven spinules in distal comb; anterior comb with five to seven spinules; how- ever, number of spinules variable. Length 3.4 to 4.6 mm, width 2 mm. Plant relationships. A total of 20 specimens was taken, mostly during 1961 and 1962 (Table 3). Table 3. Collection records of Crtjptolepidus nevadicus at the Nevada Test Site. Area Plant Host" or Community Month and Number of Specimens 1 5 Grayia-Lycium Lycium Jan. 1, Feb. 3, Mar. 8, April 2, May 2 Feb. 1, Mar. 1 6 Atriplex- Kochia Feb. 1, Mar. 2 17 'Oenothera californica June 1 28 Mixed Mar. 1 Cane Springs Mixed Mar. 3, April 2, Dec. 1 Cnjptolepidits aridity, new species Figs. 5, 6, 7 Morjihological characteristics. Black. Anten- nal club reddish-brown, compact. Vibrissae ab- sent. Scales on elytra imbricate, silvery with few brown to black intermi.xed in some specimens, with sheen which gives all specimens whitish color. Head and rostrum as long as prothorax; rostrum with median longitudinal groove free from scales; head one-third wider than rostrum at base, separated by well-defined basal constric- tion. Scrobes deep, terminating below eyes wliich are ovulate black and flattened; scape reaches posterior margin of eye; first joint of funiculus longer than second and third com- bined; club received in cupped seventh funicular segment. Prothorax wider than long, greatest width near apical region; postocular lobes and vibrissae absent. Dorsal surface covered with round, silvery white scales. Lateral surface with long setae. No median or lateral vittae. Elytra parallel, surface flat, striae fairly distinct, but punctation obscured by imbricate scales; setae Brigham Young University Science Bulletw similar in color and length to those on prothorax, but tliree times as long as ones on head and rostrum. Legs whitish, except for corbel spinules which are amber colored, and black ventral tar- sal spines. Spinules of tibial comb widely spaced and short. Fourth segment of tarsus as long as other three combined. Claws widely separated and long. Aedeagus (Fig. 8) characteristic of C. aridus, but differs in a number of respects from that of other species of this genus. (See Ting, 1940). Type locality. Nevada Test Site near Mer- cury, Nye County, Nevada. Collected by mem- bers of the Brigham Young University A. E.G. Project, 1960 tol962. Holotype in the U.S. Na- tional Museum. Four paratypes in the ento- mological collection at Brigham Young Univer- sity. Plant relationships. Five specimens were taken as follows: two in a Grayia-Lycium com- munity. Area 1, in May, 1960, and April, 1962; two in a Coleogyne community, Area 2, in Janu- ary and March, 1961; and one in an Atriplex- Kochia community, .Area 6, in February, 1961. Comments. Crijptolepidus aridus is a small species. The corbels are open with widely spaced amber colored spinules. The small size and silvery-colored scales are distinctive. Fig. 5. Cryptolepidus aridus, dorsal view sty Fig. 6. Cryptolepidus aridus, female genitalia, ventral view, .sty — .stylus; c — co.\ite; vf — valvifer; 9ths — ninth stemite; 8ths — eighth stemite Fig. 7. Cryptolepidus aridus, spemiathex;a ( receptacu- lum seminis). cii — eornii, distal portion of sperma- the.ca; rm — ramus, portion of spermatheca receiv- ing the seminal fluid; nd — nodulus, that part of the spermatheca attaching to the spermathecal gland Cn/ptolepidus cazieri (Van Dyke) References. Van Dvke, 1936b:73; Ting, 1940: 148. Morphological characteristic. Oblong, robust, black, appendages rufous, densely clothed with imbricated, gray and light brown scales, latter forming median and lateral vittae on pronotum, and irregular patches on elytra. Rostrum with pronounced basal, transverse constriction; dor- sum convex from base to nasal plate; narrow median sulcus; setae on rostrum and head same length; first funicular segment of antennae more elongate than in other species of genus; scrobes Weevils or the Nevada Test Site 13 nearer posterior end than in other species; nasal plate broadly and shallowly emarginate. Pro- notum tuberculate, broader than long. Elytra one-fourth longer than broad, rounded at base and apex, flattened, setae arranged in two or three irregular rows per interval. Hind tibia with distal comb with 10 spinules; anterior comb with five spinules. Scales dense beneath; first and second ventral segments slightly conca\e. Length 5.1 mm, width 2.8 mm. Plant relationship. One beetle was taken in a Lycium community. Area 5, in March, 1961. Comments. Cryptnlepiclus cazieri, according to Ting ( 1940 ) , may be easily distinguished from the other described species of this genus as fol- lows: pronotum tuberculate; dorsum of rostrum conve.v from base to nasal plate; only species pre- dominantly tan to brown. Subfamily Leptopiinae Tribe Ophr)'astini Genus Ophn/astes Schoenherr References. Schoenherr, 1833:508; LeConte and Horn, 1876:30; Fall, 1907:260, 1910:189; Davis, 1947:483-551; Kissinger, 1964:31. Morphological characteristics. The subfamily Leptopiinae, according to Kissinger, has elbowed antennae with compact club, rostrum grooved for their reception; rostrum not received into prostemum; rostrum never long and slender; tarsi usually dilated, third segment bilobed, brush-like beneath; mandibles with deciduous cusp, leaving scar; prothorax with anterior mar- gin produced into prominent, rounded lobe ad- jacent to eve; eye partly covered b%- anterior margin of prothorax; mandible with four or more large setae. The tribe Ophn/astini is characterized by well-developed ocular lobes, eyes elongated, transverse, accuminate beneath, and in part covered; mentum large, covering maxillae; scrobes directed beneath; rostnim large, tjuad- rangular; eyes narrow and acute below, partially concealed by ocular lobes. If the species listed in the Leng Catalogue under the genera Eupagoderes and Ophn/astes are now to be considered in the genus Opltn/as- tes, the characteristics as presented by Kissinger (1964) should be followed; antennal funiculus and dorsal portion of tarsal segments 1 to 3 clothed with decumbent, usually broad scales; rostrum trisulcate — if sulci are obliterated then tarsal segments in basal half not dotted ventrally with coarse setae; antennal scrobe usually well defined, dorsal margin rather straight; hind coxae separated by distance not greater than width of coxa; suture between sterna 1 and 2 straight; elytra regularly 9 to 10 striate, not nod- ose in apical fourth; lateral rostral sulcus not abruptly turned laterally towards antennal scrobe; prothorax wider at base than apex; sternum 2 shorter than 3 and 4 combined; tarsal segment 3 pubescent beneath and broadly bi- lobed, at least in male; prothorax more or less tuberculate laterally; apex of tibia 3 with two rows of setae, corbel enclosed. The two species of Ophrijastes collected at the test site may be separated from other species as follows: Rostrum with well-developed rnedian sulcus; thorax finely and sparsely punctate, tibiae not denticulate within; lateral sulcus of rostrum short, linear; elytral striae fine, inteivals flat, punctation fine; length 6 to 12 mm , geminuius Horn Rostrum without median sulcus; thorax more coarsely and closelv punctate; tibiae, at least anterior, denticulate within; lateral sulci of rostnim longer, arcuate, convergent basally; elytral striae closer together, deeper punctures; length 7 to 14mm varitis (LeConte) Ophrijastes vaihis (LeConte) Reference. LeConte. 1853:439-448. Morphological characteristics. Dorsum of rostrum and front continuous, without interrup- tion by transverse impression at base of rostrum; rostrum without median sulcus, convex and broadened at apical portion of rostrum. Lateral sulci of rostrum long, arcuate and convergent basally. Surface sparsely punctured, covered with white scales. Prothorax coarsely and close- ly punctate. Surface covered with silvery white scales, with median and lateral plumbeous stripe. EKtra broadest behind middle, finely striate with coarse punctures \videly separated, surface co\'ered with white scales interspersed vvdth plumbeous ones which in some species form spots or vittae. Legs and body beneath clothed with white scales. Length 5.5 to 10.5 mm. 14 Brigham YouNt; University Science Bulletin Plant relationships. A total of 63 beetles was taken. Most specimens were taken during 1960 and 1961, with others collected in other years from 1959 to 1965 (Table 4). Tabic 4. Collection records of Ophryastes vuritis at the Nevada Test Site. Plant Host" Month and Number Area or Community of Specimens 1 Grayia-Lycium Mar. 2, April 7, May 6, June 3 4 Grayia-Lycium Mar. 1, April 2. May 3, Jime I, Nov. 1 5 "Larrea divaricata May 1, June 1, July 10, Aug. 1 Lycium Mar. 1, April 1, May 3 6 Atriplex-Kochia April 1, May 7, June 1 10 ° Larrea divaricata July 1 Mixed Aug. 2 12 'Jitniperus osteospernm June 1 Pinyon-Juniper July 1 16 "Atriplcx canescens Aug. 1 26 Larrea July 1 28 Mixed May 1, June 1 Cane Springs Mixed AprU 1 Ophryastes geminatus Horn Reference. Horn, 1876:35. Morphological characteristics. Rostnnn with slight interruption by transverse impression at base of rostrum; median sulcus variable in im- pression; lateral sulci short and varying in depth; covered with wliite scales, except for small plumbeous spot in front of eyes and at base of scrobes. Prothora.x wider than long, median line rather impressed, disc coarsely punctate, covered with white scales except for plumbeous stripes on each side. Elytra one-third longer than wide, striae fine with small pimctures covered with plumbeous scales which gives vittate appearance in most specimens. Legs and body beneath covered with white scales intermi.xed with plumbeous ones. Length 6.5 to 12 mm. Plant relationships. A total of seven speci- mens was taken. Two were from a Grayia- Lycium community. Area 10, in June, 1964; one on Craijia sp.. Area 4, in June, 1960; one on Atriplcx canescens. Area 26, in June, 196.5; one on Chrijsothamnus sp., Midvalley, in June, 1965; and two on Larrea divaricata, .\rea 5, in June and September, 1961. Tribe Leptopiini Genus Orimodema Horn Reference. Horn, 1876:43. Morphological characteristics. Genus Orimo- dema with seventh segment of flagellum distinct from club; third tarsal segment broader than second, tarsi densely pubescent beneath; scrobes deep, well defined, strongly arcuate, passing be- neath at distance from eyes; first ventral suture arcuate; second sternite usually as long or long- er than third plus fourth; vestiture squamose but not intermixed with setae or pubescence; an- terior tibiae denticulate within; nasal plate de- pressed, but not sharply defined. Orimodema protracta Horn Reference. Horn, 1876:44. Morphological characteristics. The species O. protracta is elongate, densely covered with brownish scales intermixed with grayish small ones; head and rostrum as long as thorax. Pro- thorax cylindrical, broadest near apex; elytra elongate, base not wider than prothorax; disc feebly convex; feebly striate, intervals flat, densely covered with brownish-gray scales. Un- der surface densely clothed with brown and gray scales. Tibiae sparsely fimbriate. Length 10 mm, width 3 mm. Plant relationship. One specimen was taken in a Pinyon-Juniper community. Area 12, in August, 1962. Genus Paracimbocera Van Dyke References. \'an Dvke, 1938:1; Ting, 1940: 136. Morphological characteristics. Species of the Leptopiini tribe have tarsal segments on ventral surface with bristle-like setae only or with great- ly reduced pubescent tufts at apices; last seg- ment of funiculus closely applied to base of an- tennal club. The genera of this tribe are fairly common and found mainly in the western United States. The genus Paracindwcera proposed by Van Dyke (1938) has been characterized by Ting (1940) as follows: small tufts of pubescence present at apices of tarsal segments. Third tar- sal segment generally distinctly larger than sec- ond ( both characters much more pronounced on male specimens ) ; antennal scrobes shallow and greatly widened at posterior end. Postocu- Weevils of the Nevada Test Site 15 lar prothoracic lobes prominent; pubescent tar- sal tiifts in both sexes present only on third seg- ment of all tarsi. The tu'o species of this genus known for the test site may be separated, by the aid of Ting's (1940) kev, from other species as follows: Elytral setae length on declivity four times width of intervals; three to four ir- regular rows of setae per interval on declivity; tarsal bristles and corbel spin- ules dark brown, nearly black; metaepisternal suture distinct .. atra Van Dyke Elytral setae length on declivity two times width of intervals; two or tliree irregu- lar rows of setae per interval on declivity; tarsal bristles and corbel spinules reddish brown; metaepisternal suture obscure . . ... artemisiae Ting Paracimhocera atra Van Dkye Reference. \^an Dyke, 1938:2. Morphological characteristics. Black and elongate; head, prothorax and elytra densely clothed with black scales, those of head and pro- thorax granular white; scales of elytra flat and closely set; elytra with long pile, denser on de- clivity, whitish to fulvous, underside of body with mixture of whitish and pliunbeous scales and setae. Length 5 to 7.5 mm. Plant relationships. Eight specimens were taken from 1961 through 1965 as follows: four in a Grayia-Lycium community. Area 10, in June; one from Ephedra nevadensis. Area 6, in June; and three in a Pinyon-Juniper communitv. Area 12, in July and August. Paracimhocera artemisiae Ting Reference. Ting, 1940:139. Morphological characteristics. Body color gray and black mottled; with black irregular vitta along second elytral interval. Tarsal bristles and tibial spinules reddish brown. Rostrum transversely constricted at base. Prothorax wider than long; surface with scales raised, giving slight tuberculate appearance; setae shorter than elytral setae. Elytral striae fine; intervals flat, setae about length of interval widths; setae on declivity twice length of intervals width. Length 5 to 7 mm. Plant relationships. Nine beetles were taken from 1960 to 1962 as follows: two in a Larrea community. Area 5, in January and February; one in a Lycium community, Area 5, in March; two in a Coleogyne community, Area 10, in February and May; one in a Grayia-Lycium community. Area 10, in May; one in a Pinyon- Juniper community, Area 12, in August; one in an Artemisia community, Area 14, in March; and one in a mixed community. Cane Springs, in Vlarch. Genus Miioderes Casey Reference. Casey, 1888:252. Morphological characteristics. The genus Mi- ioderes now placed in the tribe Leptopiini has small tufts of pubescence present at the apices of tarsal segments. In male specimens third tar- sal segment generallv distinctly larger than sec- ond. Antennal scrobes deep and only slightly widened at posterior end. Postocular prothoracic lobes absent or only slightly evident. Pubescent tufts present in males at apices of basal tliree tarsal segments on fore legs. Rostral length slightly less than distance between eyes. An- tennal funicular segments one to six not cupped at apices. Postocular lobes slightly evident. KEY TO SPECIKS OF MILODERliS (After Ting, 1940:151) Color brown and silvery gray. Fore tibiae with outer apical portion evenly round- ed. Scales of pronotum and elytra with central punctures setosus Casey Color brown to sienna witli white scales in irregular pattern on head and elytra, scales without central punctures. Prothoracic tibae rounded apically; 12 to 13 spinules on middle tibia; black vestiture, sparse setae mercurt/ensis, new species Color uniform green. Fore tibae with outer apical portion elongated. Scales of pro- notum and elytra without central punctures viridis Pierce 16 BuiGHAM ^'olIN^. University Science Bulletin Milodeies mercun/ensis. new species Kigs. 8. 9 Morphological characteristics. M ilo d e r e s mcrciin/cnsi.s, one ot the tiiree species ;i,scribed to this genus, may be separated from the other two as follows: size 4 to 4.3 mm; brown to sienna witii white scales in irregular pattern on head and elvtra. Retldisli-broun scales on mar- gins of prothorax, on legs, especially t;irsi and antennae. Rostrum wider than long with trans- verse impression lust slightly posterior to an- tennal luticulation. with median fo\ea on trans- verse impression, r'ronotum broader than long, greatest width near apex; postocular lobes pres- ent, rounded opposite eye, with long, golden colored \ibrissae which reach eve; setae on head imd between closeh" arranged round punctures. Elvtra with closeh' placed, det^p stiial punctiu-es; setae length three to four times width of inter- vals. W'lvite scales irregularb' arranged, thus making mottled white and brownish pattern; sides parallel with rounded lumieral angle. Pro- thoracic tibiae rounded apicallv; third segment ml — Fig. S. Milodcres mercurticnsis. dorsal N-icw Fig. 9. Milodcres iiiercunjensis, aedeagus. ventral view, mo — mediiui orifice; op — orificial plate; b — ^bacu- liun: ml — median lobe; tg — tegmen; ts — tegminal stmt; ms — median strut of tarsus broad, deeply emarginate; under-sur- face covered with white setae. Middle tibiae with 12 to 13 spinules. Posterior tibia with nine spinules, corbels open. The small size, color, scales without central punctures, short rostrum, 12 to 13 spinules on middle tibia, black \esHture, sparse setae, and shorter length than M. sctostis and M. tiridus characterizes M. merciin/ensis. The male geni- talia (Fig. 9) is distincti\e. Type locality. Nevada Test Site, near Mer- cury, Nye Coiuitv, Ne\ada. Collected by mem- bers of the Brigham Yoimg Uni\ersitv A. E.G. Project, 1961 to 1965. Holotyj^e male in' the U.S. National Museum. Two parat\'pes in the ento- mological collection at Brigham Yoimg Univer- sity. ' Plant relationships. One specimen was taken from Sphacralcca amhigua. .\rea 5, in June, 1964; one from Grai/ui spinosa. .\rea 26, in June, 1965; and one from a pit-can trap on the Jackass Flats approach road in a mixed plant association with- in the Larrea-Franseria communit\-. Comments. Figure S show s the cusp in place on the tip of the mandible of one of the speci- mens. This is a most interesting species. Genus Diiotognuthii.'i Horn Reference. Horn, 1876:79. Morphological ch;vracteristics. This mono- t\'Y>ic genus has ociihu" lobes, eyes elongate, trans\erse. acuminate beneath, and at least in part co\ered; mentum small, not retracted; Weevils of the Nevada Test Site 17 maxillae free; scrobcs feebly angled, rapidly evanescent. Dirata^nathu.s sordidus Horn Reference. Horn, 1876:80. Morphological characteristics. Dirotognathn.s sordidu-i i.s oval, robust in form; covered with dark cinerous scales with short, decumbent, dark brown setae. Head and rostnim as long as thora.x which is oval, broader than long. Elytra broad- ly oval, broader at base than thorax, finely striate; intervals flat. Body beneath and legs less densely scaly and very sparsely hairy. Length 5.2 mm, width 3 mm. Plant relationship. Six specimens were col- lected in a Pinyon-Juniper community, Area 12, during June, July, and August, 1961. Subfamily Cleoninae Tribe Cleonini Genus Cleoiius Schoenherr Reference. Schoenherr, 1826:14.5. Morphological characteristics. The genus Cleonus may be characterized as follows: beak not dilated at tip; gular peduncle shorter than in Lixu-s; front and middle tarsi broad, third seg- ment at least spong)' and bilobed; hind tarsi hairy beneath first division, but broad and spongy in second; pubescence in longitudinal stripes, no transverse or oblique bands. Body elongate. Blatchley and Leng (1916:.329) ob.served that it is difficult to set forth in a key any defin- ite characters which sharply define the two genera, Cleonus and Lixus. Cleonus denticollin Casey Reference. Casey, 1891:180. Morphological characteristics. Cleonus den- ticollis is narrow to sUghtly robust, somewhat de- pressed above, elongate-suboval; black, densely clothed with short, recumbent, squamiform pu- bescence, cinereous to ferruginous in color; some sparse, short erect hairs; head sparsely punctate; beak shorter than prothorax, broadly bisulcate, densely pubescent above, median carina narrow but not acute, moderately ele- vated. Prothorax one-fourth wider than long, widest at anterior lateral tubercles which are very pronounced; sides behind them parallel to base, latter broadly cusped in middle, wider than apex; disc courselv foveolate, sides covered with white pubescence; middle sparsely clad. Sciitel- luin inconspicuous. Elvtra t^vo-thirds longer than wide; humeri not j^rominent; disc with ele\'ated altcniatc iutcrxals; pubescence denuded on spots at basal j)ortion of each elytron and along suture near apex in some specimens. Abdomen densely pubescent. Legs moderately stout, femora feebly annulate at apical third. Length 10.0 to 10.2 niiu., width 4.0 to 4.2 mm. Plant relationships. A total of 1.52 specimens was taken, mostly in 1960 and 1961 (Table 5). Table 5. Collection records of Cleonus denticollis at the Nevada Test Site. Plant Host" Month and Number Area or Community of Specimens 1 Grayia-Lycium Salsola Jan. 1, Mar. 48, April 1.5, May 3, June 5, July 22, Aug. .5, Sept. 3, Oct. 2 Mar. 1, July 6, Aug. 2, Sept. 1, Nov. 2 4 Grayia-Lycium April 1, Oct. 1 5 Larrea June 8, July 4, Aug. 8, Oct. 1 6 Atriple.v-Kochia June 1 10 Coleogyne July 1 ° Hymenoclea fasciculata July 7 'Larrea divaricata July 1 28 Mixed Feb. 1, July 1, Aug. 1 Cleonus lohigerinus Casey Reference. Casey, 1891:191. Morphological characteristics. Casey charac- terized C. lobigerinus in this manner: body slen- der and convex, integument feebly shining, pu- bescence cinereous, a broad discordal anteriorly constricted spot denuded along middle, margin- al vitta on pronotum, also denuded on second, sixth and eighth elytral intervals. Head trans- versely impressed between eyes, rostrum rather long, feebly carinate, subequal in length to pro- thorax. Prothorax nearly as long as wide, apex broadly bisinuate, narrower than base, latter angularly lobed in middle, sides convergent from base to apex; disc coarsely punctate, deeply im- pressed in basal third, slight evidence of being carinate anteriorly. Elytra more than two times as long as wide; sides parallel and nearly straight, disc with deep approximate punctures. Abdomen and legs densely pubescent with few sub-denuded punctures. Length 8.9 mm, width 4 mm. Plant relationships. Three specimens were taken as follows: one in a Grayia-Lycium com- 18 Brigham Young University Science Bulletin munity. Area 10, in June, 1964; one in a Salsola community. Area 7, in July, 1962; and one from Atriplex canesccns. Area 16, in August, 1964. Subfamily Erirliininae Tribe Smicronvchini Genus Smicroni/x Scboenherr Reference. Scboenherr, 1843:313. Morphological characteristics. The genus Smicromjx has tibia 3 unarmed or with spine less than one-half as long as tarsal claw; body lack- ing waterproof covering. Prostemum with coxae situated much closer to hind margin than to front margin; rostrum distinctly longer than head (in lateral view ) ; scape not reaching anterior mar- gin of eye; tarsal claws connate at base; femora unarmed; prothora.x with postocular lobes; tarsal segment 4 shorter than segments 1 to 3 com- bined. Smicrotnjx inihricattis ( Casey ) References. Casey, 1892:391-392; Anderson, 1962:203-305. Morphological characteristics. Rostra of both sexes black to piceous, slender and curved; squamose, punctate from base to near apex in male and female. Head black, squamose; an- tennae piceous, with pale brown scales. Pro- thorax black, sides subparallel; evenly covered with deep rounded punctures; scales ovate, white at sides and along midline, medium to light yellowish-brown elsewhere; prosternum deeply emarginate, not concave. Elytra black, intervals covered with imbricate elliptical scales, mostly white in humeral regions, light or medium brown with scattering of white posterior to humeral regions. Underside of thorax and ab- domen black covered with ovate, white scales having violet to bluish irridescence. Femora red- dish, covered with elongate and ovate, pale brown to white scales. Tibiae black. Tliird seg- ment of tarsi broader than first two; fourth seg- ment extending beyond third; claws coimate near base. Length 2 to 2.4 mm, width 0.70 to 1.00 mm. Plant relationships. A total of 54 specimens was taken, mostly during 1962, 1964 and 1965 (Table 6). Smicronyx sp. Specimens of Smicrom/x not referred to a species were taken at the test site. They are very Table 6. Collection records of Smicronyx imhricatus at the Nevada Test Site. Plant Ho,st° Montli and Number Area or Community of Specimens 5 "Lijcium pallidum July 1 Larrea July 1 12 ° } uniperus osteospcrma June 4, JiJy 5 °Pinus monophijlhi June 8, July 5, Aug. 7 " Chnjsothumnus vixcidif torus Aug. 1 ° Chriisothamnus luiuseosus Aug. 1 'Ribes sp. Aug. 2 16 'Artemisia tridentata July 1 ° Argemone corymlyosa June 1 'EuTotiu lanata June 1 'Mentzclia albicaulis (une 1 'Sallugia paradoxa June 2 'Piiius monophylhi June 2 17 Unknown Aug. 1 18 'Grciyia spinosii |une 1 °Pinus monophylhi June 2 °]uniperus osteospcrma June 3 'Guticrrezia micrncephula Aug. 1 23 Mixed July 1 26 'Atriplex canescens June 2 °Pinus monophylla June 1 similar to S. cognatus Dietz, now considered as a synonym of mucidus Dietz. Plant relationships. Three specimens were collected in Julv, 1962, as follows: one from Artemisia tridentata. Area 14; one in a Larrea community. Area 5; and one in a mi.xed com- munity. Area 23. Genus Promecotarsiis Casey Reference. Casey, 1892:408. Morphological characteristics. Body cylindri- cal, convex, longer and with more glabrous tarsi having small third segment, fourth segment very long and subequal in length to entire remainder. Vestiture dense over entire body, consisting of small imbricated scales, with recurved, subre- cumbent setae on each strial interval. Rostnim constricted at base, head spherical, eyes oblong. Prothorax constricted at apex, ocular lobes more or less distinct. Scutellum small. Abdomen flat, second segment as long as the ne.xt two and not cjuite as long as fifth. The test site species may be separated from others as follows: Wee\ils of the Nevada Test Site 19 1. Tarsal claws siibparallel; prothorax much wider than long, with ocular lobes prom- inent ftimatus Casey Tarsal claws widely divergent; prothorax very nearly as long as wide; ocular lobes not prominent 2 2. Prothorax abruptly, deeply constricted near the apex, latter but slightly narrower than base maritimus Casey Prothorax gradually more strongly narrowed and broadly, feebly constricted to- ward apex, latter scarcely more than two-thirds as wide as base .. densus Casey Prumecotaniis densus Casey Reference. Casey, 1892:410. Morphological characteristics. Promecotar- sus densiis is robust, subcylindrical, convex, black, densely clothed witli oval, whitish over- lapping scales, with distinct recurved setae. Head glabrous, constriction deep; beak slender, polished, nigosely punctate near base, almost tliree-fifths as long as elytra; antennae inserted beyond basal third, second funicular segment about as long as next two; club compact and shining towiu-d base. Prothorax wider than long, sides parallel, convergent and constricted at apex; disc convex and very densely punctate tlirougliout. Elytra at base about one-fourth wider than prothorax, one-half longer than wide; striae indicated by fine but shaqily defined part- ings of dense crust of scales. Legs stout, tarsi long, divergent, connate at base. Length 2.4 mm, width 1.1 mm. Plant relationships. Three specimens were collected in 1964 and 1965 as follows: two from Artemisia tridentafa in August, and one from Jtinipertis osteospenna in July, all in Area 12. Subfamily Apioninae Genus Apion Herbst Reference. Herbst, 97:100. Morphological characteristics. Weevils of the subfamily Apioninae are characterized by having straight antennae, scrobes wanting; segments of antennae club compactlv united; pygidium con- cealed; trochanters large, femora attached to apex; form pear-shaped. Small, not over 4.5 mm. Two genera constitute this subfamily in the United States. The genus Apion may be distin- guLshed from Podapion as follows: front femora not stouter than others; antennal club large, last segment larger than preceding. Apion vctriconie Smith Reference. Smith, 1884:60. Morphological characteristics. The only species of the genus Apion collected on the pro- ject at the test site is varicorne. It is black in color, covered with dense white pubescence. Antennae black except basal segment which is yellow; rostrum of female longer than that of male; intervals of elytra flat. Length 1 to 1.3 mm. Fall (1898:160) discussed several varieties of this species. Kissinger (1964:32) placed vari- corne in a group with three other species which occur throughout the western and southern United States, Mexico and Guatemala. Plant relationships. Fourteen specimens were collected. Four were taken from Dalea pohja- denia near Buckboard Mesa in July, 1965; five from D. pohjadenia in Jackass Flats in August, 1965; four from D. pohjadenia and one from Eriogonum nodosum near Tippipah Spring in August, 1965. Subfamily Rhynchitinae Auletobius hiimeralis Boheman Reference. Boheman 1859:117. Morphological characteristics. Three-fourths longer than wide; black, with elytra reddish- brown, except for nan-ow black line along suture; punctation fine; pubescence sparse, short, semi- erect, whitish. Head with occiput wider than long, feebly convex between eyes; punctation between eyes shallow, polished, eyes large and prominent. Rostrum as long as thorax and occi- put, slightly arcuate behind base of antennae; antennae inserted within basal third of beak, two basal segments stout, others small; club large, very loosely jointed. Prothorax widest at basal third, one-third wider than long; base broadly and feebly arcuate; disc feebly convex, scutellum large. Elytra broadly and separately rounded at apex, one-half longer than wide, two- Brigham Young University Science Bulletin thirds wider than prothorax; disc convex, im- pressed along suture; claws armed with tooth. Length 2.1 mm. Plant relationship. Two specimens were col- lected in June, 1965, on Ephedra nevadensis in the environs of Cane Springs. Auletohius sp. Represented by three specimens. Color of head and body bluish black with sheen; rostrum and legs nifus. Size 2 mm. I am unable to determine this species. It does not agree with any described species. I have before me specimens of all the species listed in Leng's catalogues except mariposae Zimmerman and those described by Voss. One specimen was taken from Ephedra nev- adensis near Cane Springs in June, 1965, and two specimens from Chrysothamnus viscidiflorus on Rainier Mesa in August, 196.5. Subfamily Myrmecinae Genus Mijrmex Sturm Reference. Stium, 1826:172. Morphological characteristics. Members of the subfamily Myrmecinae to which the genus Mtfrmcx belongs have tibiae that are not fos- sorial; rostnmi free, not received by prosternum; humeri not truncated by protruding mesopleura; lateral angles of first sternite covered by elytra; beak usually at least as long as prothorax; gular peduncle usually long; sternites nearly equal, or first longer; claws usually more or less dentate; prothorax pedimculate; form ant-like. Four genera in this tribe may be separated by the following key (Kissinger, 1964:50-51): KEY TO THE GENERA OF MYRMECINAE 1. Front coxae inserted near middle of prosternum, co.xae more or less etjuidistant from anterior and posterior margins of prosternum; inner margin of mandible toothed - 2 Front coxae much closer to hind margin of prosternum than to front margin, dis- tance to front margins more than three timej as great as distance to hind mar- gin; mandible slender, triangular in outline, inner margin with minute basal tooth; rostrum slender, very long; femora clavate, with large triangular tooth Erodisctis Schoenherr 2. Elytra elongate oval, humeri well developed 3 Elytra ovate, rounded, humeri absent Oopterinus Casey 3. Femora toothed; male lacking dorsal excavation on rostrum; length more than 3.0 mm Mijrmex Sturm Femora not toothed; male with dorsal excavation on rostum; length less than 3.0 mm Micromt/rmex Sleeper Mtjrmcx lineata lineata (Pascoe) Reference. Pascoe, 1872:454. Morphological characteristics. Mijrmex I. lineata is a fairly common sjjecies in the south- ern portions of the Great Basin. Tlie single speci- men contained in the collections made at the test site is elongate, subcylindrical, color black with dense gray pubescence. Rostrum three- fourths as long as pronotum, with median, im- punctate polished area. Antennae black, scape reaching eye, first segment of fimicle longer than second; club elongate, oval, feebly pubescent Head clothed with white pubescence, sparsely intermixed with dark brown setae; punctate. Eyes with many facets, round. Pronotum slight- ly arcuate at middle, with pronounced median carina; densely punctate; from each puncture issues many branched setae (palmate); scutel- lum prominent, densely covered with very fine, white setae. Elytra with glabrous prominent in- tervals; dense pubescence on intervening inter- vals, surface slightly punctate, sides practically parallel. Legs and venter densely clothed with white setae; each femora with small acute tooth; claws toothed. Length 12.3 mm, width 3 mm. Plant relationship. One specimen was collect- ed in a Larrea community. Area 5, in October, 1961. Weenils of the Nevada Test Site Subfamily Magdalinae Tribe Magdalini References. Germar, 1817:340; Horn 1873- 407-469; Fall, 1913:27. Morphological characteristics. Species of this tribe and genus have anterior ccvae contiguous; elytra unicolorous, usually black, glabrous and widened posteriorly; claws simple or toothed. Magdalis Iccontei tenebrosa Fall Fig. 10 Reference. Fall, 1913:28. Morphological characteristics. Magdalis Je- contei tenebrosa Fall is black throughout; fe- mora toothed, not impressed at base; hind angles of thorax more or less produced and divergent. Antennal club normal, second funicular segment twice as long as wide, and as long as two fol- lowing. Length 2.5 to 5 mm, width 2 to 3 mm 21 Fig. 10. Magdalis lecontei tenebrosa Plant relationships. Eleven specimens were collected in 1964 and 1965 as follows: nine in June and one in July from Piiuis monopht/lla. Area 12; one from a Pinyon-Juniper community Area 12, in July. Subfamily Anthonominae This is a large subfamily according to Kis- singer (1964), consisting of two tribes, Endaeini with eleven genera and AntJwnomini with four- teen. The following characters will aid in placing species in this subfamily collected at the Nevada Test Site: trochanter short and triangular, femur attached to side of trochanter, base of femur closely adjacent to co.xa; antennae distinctly el- bowed, funiculus consisting of five to eight seg- ments; scape fitting into scrobe; usually inserted toward apex of rostrum; club elongate, oval, en- tirely pubescent, distinctly annulate; rostrum not received into prosternal emargination, slen- der, usually longer than prothora.x, mandibles small, lacking deciduous cusp; tarsus with four distinct segments, segment 3 usually strongly bi- bbed; mesepimeron not ascending and not visi- ble in dorsal view; front co.xae contiguous; claws free at base and with basal tooth; hind tibiae mucronate at tip; mesepimeron not ascending and not visible in dorsal view; suture between sterna 2 and 3 deep and straight. Tribe Anthonomini The characteristics for the tribe Anthonomini are, in the main, the same as for the subfamily Anthonominae. Thus, if a specimen agrees with the above assemblage of characters, it may well be considered as belonging to the tribe Anthono- mini. The species from the test site which belong to this tribe represent four genera which may be separated by the following key: 1. Claws toothed Claws simple Hind tibiae mucronate; prosternum short in front of coxae. Form convex; tibial armature very evident Posterior tibiae unguiculate;pygidium covered; claws armed with a long tooth- mat Sn^l' ' ''''''*' '^'' ^^''''"^ latero-inferiorly, somewhat !pproxi: ^"^'^ ^ Macrorhoptus LeConte ^°''^enn/^fi'' 7 ""°"^;*"\^^''°b"- l°"g' directed against the eye; funicle of an- tennae 6 or 7 jomted; claws elongate, ovoidal J. Anthonomus Germar 3. Scrobes directed against the eye £„ imecbus Dietz 22 Bhigham Young University Science Bulletin Scrobes broad and deep for about one-half the distance toward the eyes, then gradually evanescent Biachiiogmiis Linell Genus Macrorhoptus LeConte Reference. LeConte, 1876:208. Morphological characteristics. Specimens of the genus Macrorhoptus have the following char- acteristics: claws toothed; middle coxae separ- ated; posterior tibiae unguiculate; pygidium covered; rostnmi rather short and stout; eyes latero-inferior, somewhat approximate beneath. With the aid of the key prepared by Sleeper (1957:71) the species of this genus may be separated. KEY TO THE KNOWN SPECIES OF MllcrorhoptUS 1. Scaly vestiture on elytra without intermixed erect or semierect clavate bristles Scaly vestiture on elytra with intennixed erect or semierect clavate bristles 2. Scales on rostrum more or less prostrate, broadly oval, those on "ocular lobes" broadly oval; Texas and California estriatus LeConte Scales on rostrum erect, elongate and narrowly clavate, those of "ocular lobes" elongate, narrowly clavate; British Columbia sidalceae Sleeper 3. Prostrate scales on disc of elytra longitudinally striate, for the most part Prostrate scales of disc of elytra not striate at all; smaller and paler species; Texas sphaeralciae Pierce 4. Scales of elytra prostrate, having a smoothly, evenly placed appearance; rostrum of female short and stout; Arizona hispichts Dietz Scales of elytra very grizzled, being roughened in appearance; rostrum of female elongate and cylindrical; California griseus Sleeper Macrorhoptus hispid us Dietz Reference. Dietz, 1891:185. Morphological characteristics. Macrorhoptus hispidus has pale grav or brownish scales which are longer and broader than in estriatus. Pro- thorax densely and finely punctured, with broad stripe of pale brown scales along middle; sides paler. Legs moderately long; femora slightly clavate, anterior armed with triangular tooth, middle and posterior mutic. Tarsi moderately stout, [X)sterior more slender, third segment bi- lobed. Claws armed with tooth. Length 2.4 to 3.3 mm. Plant relationships. Ten specimens were col- lected during 1965 as follows: one from Sphaer- alcea sp., Cane Springs, in June; two from Sphaeralcea sp., Area 17, in June; three from Sphaeralcea sp., Area 26, in June; one from Gratjia spinosa, Area 18, in July; and three from Artemisia sp.. Area 18, in July. Comments. It is difficult to separate M. his- pidus and M. sphaeralciae. The specimens be- fore me representing the three species as above probably would be considered as two valid species with sphaeralciae as a subspecies of his- pidus. Genus Anthonomus Germar Reference. Gennar, 1821:320. Morphological characteristics. Rostrum usual- ly long and slender; scrobes long, directed against the eyes; antennal scape reaching the eye or nearly so; funicle 6 or 7 jointed; club elongate-ovate; elytra distinctly striate and punc- tured; pygidium in the male more or less ex- posed, femora usuallv toothed; front and middle tibiae with a hook, hind ones with a spine at tip; claws toothed. Weevils of the Nevada Test Site 23 KEY TO THE SUBGENERA OF AnthonOmilS AT THE NEVADA TEST SITE (After Dietz, 1891:189) 1. Posterior tibiae alike in both sexes Posterior tibiae of male curved Cnemocyllus 2. Eyes small, subrostral Anthonomorphus Eyes at least moderately large, position normal. Vestiture of derm not intermixed with erect setae. Prostemum not emarginate in front, last segment of funicle distinct from the club. Club elliptic or ovoidal, not very loosly articulate Anthonomus Dietz divided the subgenus Anthonomu.i into nine groups. Anthonomtts holteri confusus be- longs in the Sutiiralis group, A. histiis the Squa- mosus group, and the species described by Fall and ornatulus of Dietz in the subgenus Cnemo- cyllus. Subgenus Anthonomorphus Anthonomus peninsularis Dietz Reference. Dietz, 1891:195. Morphological characteristics. Oblong oval, color variable, black to reddish brown; clothed with white pubescence above and beneath; rostrum slender, with striae well developed; an- tennae inserted about two-fifths from apex; sec- ond segment of funicle longer than third. Eyes round. Head constricted behind eyes; coarsely punctate, sparsely subescent; frontal fovea deep. Prothorax much \vider than long, twice as wide at base as at apex; surface coarsely punctate, each puncture bearing decumbent white seta; more closely placed at base and median line. Elytra wider at base than prothorax, striae prom- inent with deep punctures; interspaces slightly convex and lightly punctate; surface wliite pu- bescence. Legs slender; femora bidentate; tarsi with bifid claws, tooth long. Length 5 to 5.5 mm, width 2.6 to 3 mm. Plant relationships. Seven specimens were collected in 1962 and 1965 (Table 7). Table 7. Collection records of Aiithonomus peninsularis at the Nevada Test Site. Plant Host" Month and Number Area or Community of Specimens 4 Mixed April 1 17 'Sphaeralcea sp. June 2 18 'Sphaeralcea sp. June 1 26 'Sphaeralcea sp. June 1 'Ephedra nevadensis Jvme 1 28 'Sphaeralcea grossu la riaefolia May 1 Subgenus Anthonomus Group Suturalis Anthonomus haematopus confusus Dietz Reference. Dietz, 1891:209. Morphological characteristics. Anthonomus confusus is now considered to be a subspecies of A. haematopus Boheman. It is similar in form to A. holteri Dietz, the tribe of which was describ- ed from specimens from New Mexico. It is little smaller, nifo-testaceous, with coarse conspicuous wliite pubescence. Rostrum stout and slightly curved; coarsely pimctured. Antennae stout, second segment of funicle longer than third. Club elongate and blackish in color. Eyes slight- ly convex. Head with coarse punctures and frontal fovea. Prothorax constricted at apex, punctate, with coarse pubescence; elytra \vith deep strial punctures; pubescence short and sparse. Length 2.5 mm, width 1.5 mm. Plant relationships. Six specimens were taken from Juniperus sp., and one from Artemisia tri- dentata, all in Area 16 in July, 1962. Subgenus Anthonomus Group Squamosus' Anthonomus liirtus LeConte References. LeConte, 1876:203; Dietz, 1891: 233. Morphological characteristics. Similar in form and sculpture to A. murinui Dietz; densely cov- ered with grayish and oclireous scales; body, antennae, and legs rufofemiginous. Rostrum slender and scarcely striate or punctured; second segment of funicle practically same length as third one. Tliree thoracic vittae, and along suture paler, legs nifopiceous, femora not strongly clavate, anterior armed with small tooth, middle and posterior mutic; tarsi rufus. Length 3 to 3.2 mm, width 1.3 mm. 24 Bbigham Young University Science Bulletin Plant relationships. Three specimens were taken in June and August, 1965, as follows: one from Sphaeralcea sp., Area 12; one from Eriogo- num iimbellatum. Area 16; and one from Arte- misia tridentata, Area 19. Subgenus Cncmocyllus Anthonomus ornatulus Dietz Reference. Dietz, 1891:241. Morphological characteristics. Elongate sub- ovate, piceous, antennae and legs reddish, dense- ly clothed with broadly oval scales, white and brown on upper surface and white beneath. Rostriun long, pinictured with distinct median carina. Antennae slender, club dark. Eyes more convex. Head short, punctures deep, fovea obso- lete. Prothorax wider than long. Punctures deep, surface covered with scales. Elytra oval, punc- tures concealed by white and some brownish scales, interspaces slightly convex; legs slender, clothed with white scales; anterior femur with scarcely perceptible tooth; tarsi slender and as long as tibiae; first segment longer than second in male, or equal to it in female; last segment long, distal end and claws blackish, latter armed with short obtuse tooth. Length 2.9 mm, width 1.1 mm. Plant relationship. One specimen was taken from Eurotia lanata. Area 16, in June, 1965. Subgenus Cnemoci/Ilus Anthonomus sphaeralciae Fall Reference. Fall, 1913:.55. Morphological characteristics. Elongate ob- long, pict'ous; rostrum, antennae and legs rufus; vestiture dense both above and below, consist- ing of pale ochreo-cinereous scales which are about twice as long as wide, almost uniform in color above except along median line and at sides of prothorax where they are whitish. Scutel- lum densely white. Body beneath covered with whitish scales; antennal funicle seven-segmented, second joint nearly twice as long as wide. Pro- thorax nearly as long as wide, subconical, apical constriction feeble. Front femora with small pointed tooth, middle and hind femora mutic. Length 3.2 to 3.4 mm, width 2 mm. Table 8. Collection records of Anthonomus sphaeralciae at the Nevada Test Site. Plant Host" Montli and Number Area or Community of Specimens 12 'Sallugia paradoxa ' Chrysothammts viscidiflonis 'Sphaeralcea sp. "Elymus cinereus June 1 Aug. 1 Aug. 3 Aug. 1 16 "Atriplex canescens 'Spliaeralcea sp. Aug. 1 Aug. 3 17 "Tetradtjmia glabrata July 2 18 'Oenothera calif ornica lune 1 "Artemisia sp. July 1 19 'Artemisia tridentata June 1 23 Mixed Aug. 1 28 'Sphaeralcea sp. June 1 Plant relationships. Seventeen were collected in 1965 (Table 8). specnnens Subgenus CncmoctjUus Anthonomus cycliferus Fall Reference. Fall, 1913:56. Morphological characteristics. Form oval, piceous, legs and antennae rufus, club fuscous; clothed with overlapping oval or nearly circular white scales, intermixed with scattered darker scales varying from pale brown to blackish purple in color; these darker scales aggregated most noticeably in two imperfect discal pronotal vittae and in elongate discal elytra spot at about postepinotal vittae and in elongate discal elytral spot at about posterior third, and less evidently in sub-basal spot on either side of suture. Ros- trum one-fourth longer than prothorax, finely punctate, polished; origin of antennae at point two-fifths from apex in male, just beyond middle in female; prothorax three-fourths as long as wide, sides arcuate and subparallel in basal half; surface completely concealed by vestiture. Elytra at base little wider than thorax, widest about middle, striae invisible. Front femora with small tooth; middle and hind femora not visibly toothed. Ungual teeth short and not approximate at tip. Length 2.7 to 2.8 mm, width 1.1 mm. Plant relationships. Four specimens were taken in June and August, 1965, as follows: one from Chiysothamnus teretifolius. Area 6; two from Artemisia tridentata. Areas 12 and 17; and one from Chaenactis stevioides. Cane Springs. Subgenus Cncmocyllus Anthonomus tenuis Fall Reference. Fall, 1913:57. Morphological characteristics. Narrowlv oval, piceous, scape and tarsi fusco-testaceous, club Weevils of the Nevada Test Site fuscous; vestiture wliite tliroughout, scales cir- cular, not dense or overlapping. Prothorax about tliree-fourths as long as wide; sides arcuate; ely- tra at base wider than prothorax; striae visible; femora without tooth. Tarsal claws with long teeth, approximate at tip. This species is closely related to A. cyclifcrtis, but is much smaller. Length 1.3 to 1.4 mm, width 0.9 to 1.0 mm. Plant relationships. Nine specimens were taken from 1960 to 1964 (Table 9). Table 9. Collection records of Anthonomus tenuis at the Nevada Test Site. Plant Host" Month and Number Area or Communit)' of Specimens 6 Grayia-Lycium Aug. 1 12 ' Chrtjsothatnnus paniculatus Aug. 2 16 'Artemisia tridentata Julys 'Jtmipertis sp. JvJy 1 26 Larrea July 1 Cane Springs Mixed July 1 Epimechus gracilis Fall Reference. Fall, 1913:59. Morphological characteristics. Form narrow and elongate, black; antennae, except club, legs and sometimes rostrum rufous; vestiture consist- ing of large, round, white overlapping scales. Rostrum longer than other specimens of this species before me, sejuamose at base, finely punc- tate. Antennae inserted near middle of rostrum; funicle six-segmented; first and second com- bined, as long as other four segments; prothorax wider than long, widest at about middle, apical constriction broad, surface coarsely punctate. Elytra twice as long as wide, and three times as long as prothorax; striae strongly punctured. Femora mutic; claws simple. Length 1.6 to 1.9 mm, width 0.70 to 0.80 mm. Plant relationship. Four specimens were taken from Chrsothamnus viscidiflorus, Areas 12 and 17, in August, 1965. Genus Brachijogmiis Linell Reference. Linell, 1897:51. Morphological characteristics. Type of the genus Bracht/ognuts is a small species; length 2.9 to 3.1 mm, width 1.3 mm; form convex; pro- thorax narrow; rostrum longer than head and prothorax; antennal scrobes straight, directed against the eyes, but abbreviated long before reaching them; hind tibiae mucronate; prostem- um short in front of the coxae; claws simple, divergent. Tliis genus is closely related to Epimechus from which it differs in the narrow prothorax and the gradually evanescent scrobes. Bracliiiogmus ornatus Linell Reference. Linell, 1897:51. Morphological characteristics. Form sub- ovate, piceous, body color concealed bv rounded scales, variegated \vith white, black and ferrugi- nous, broad blackish variable band across elytra at about the middle. Rostrum longer than head and prothorax, at base scaly with white and fer- ruginous, outwardly shining piceous. Scrobes commencing t\vo-fifths from apex, deep for about one-half distance toward eye, then grad- ually evanescent. Scape of antennae reaching eye, clavate at apex; funicle as long as scape, seven-segmented, each segment bearing whorl of long white hairs. Eyes roimd, front between eyes depressed and covered with ferruginous scales. Prothorax as broad as long, sides rounded, disc covered with white and ferruginous scales which are \'ariable in pattern; some with white median line bordered with dark bands. Scutellum conspicuous, with dense white setae. Elytra at base much wider than prothorax; twice as long as broad; striae deeply impressed, punctures concealed; intervals slightly convex, suture ele- vated toward apex; broad band across suture at middle witli numerous spots on other areas; scales white, blackish to ferruginous. Ventral scales white, mi.xed with ferruginous ones. Fe- mora clavate, toothed; tibiae mucronate at apex; tarsi piceous, clothed with white setae; first two segments equal, third emarginate, slightly wider; claws strong, black. Length 2.9 to 3.1 mm, width 1.3 mm. Plant relationships. Two specimens were taken in April, 1961; ten in July, 1960; and one in July, 1962, all from Li/cium pallidum in Area 5. Subfamily Tychiinae Tribe Tycliiini Genus Tijchius Germar Reference. Gennar, 1817:340. Morphological characteristics. The genus Tij- chius has the following distinctive character- istics: body elongate-oval, with robust recurved setae; eyes large, very nearly circular; head dor- sally not constricted behind eyes; antennal Bhigham Young University Science Bulletin funicle with six segments. Casey divided this genus into four subgcneric groups; T. prolixus is the only one placed in the third subgeniLs which is said to have but six segments in the antennal funicle. Tychitis prolixu.9 Casey References. Casey, 1892:419; Kissinger, 1964: 57. Morphological characteristics. Tychius pro- lixui- is a blackish, oblong-elongate, convex form; tip of rostrum and antennae pale reddish; vesti- ture dense, consisting of long, slender subrecum- bent squamules on pronotum, pale fulvous in color but wliitish along middle and sides; on elytra scales large, dense, rounded, imbricated with some reddish-gray in color; each interval with single series of coarse, recurved, reddish pointed setae. Head with large circular eyes; beak with prominent tuft above each eye; gla- brous and shining beyond antennae, equal in length to prothorax. Prothorax wider than long; sides feebly arcuate from base nearly to apex, then rather abruptly constricted. Elytra wider than prothorax and not (|uite three times as long; sides parallel and straight in basal tvvo-thirds; apex obtuse; depression at declivity. Legs and iinder-surface clothed with whitish scales. Length 3.3 mm, width 2 mm. Plant relationship. One beetle was taken from Astragalus lentiginosus. Area 17, in June, 1965. Subfamily Cryptorhynchinae Tribe Crj'ptorhynchini Genus Zascelis LeConte Reference. LeConte, 1876:256-258. Morphological characteristics. The genus Za- scelis in the United States is confined to lower California and the southwestern United States. The species are elongate, depressed, coarsely sculptured, pubescent and sometimes scaly. Ros- trum long, slender, and extends almost to meta- sternum. Antennae vary with species. Ventral sutures deep, first one slightly sinuate; first ven- tral segment longer than second which is equal to third; fourth and fifth equal in length. Meso- sternum elongate and deeply excavated almost to base. Femora anned beneath with small tooth; tibia broad, compressed, with row of teeth along anterior margin which is variable in species. The following is a key to the species of Zascelis: 1. Pubescence long, not mixed with scales 2 Pubescence shorter; tibiae feebly toothed 3 2. Tibiae very coarsely toothed; length 7 mm . serripes LeConte Tibiae very finely and sparsely toothed; length 4 to 4.8 mm ohlonga Horn 3. Pubescence short, erect, intermixed with large scales squamigera LeConte Pubescence prostrate, ssed. Legs slender, femora not clavate, tibiae feebly serrate on inner side, especially front pair; inner angle unguiculate; tarsi slender, third segment bilobed, not spongy beneath, but smooth and glabrous like others. Tlie outstanding characters of this genus are: eyes contiguoiLs beneath and widely separated above; antennal club is corneous sheath; small tliird tarsal segment not spongy beneath. Casey (1892) described three species and proposed the following key which may be used to separate them from Y. frontalis. 1 have added lentiginosus Csy. (1904) to the key: 1. Piceous-brown, punctation of the upper surface finer and more remote 2 Black, much larger, coarsely and deeply sculptured; legs and tarsi stouter grossus Csy. Black, smaller, finer and less deeply punctured; many of the punctures surround- ed by pale yellowish-white spots, legs moderate, shining, finely punctate Icntiptnosus Csy. 2. Body narrowly cylindrical, the elytra more than twice as long as wide; punctures of the elytral series becoming ver)' fine and feeble in apical half as usual; fifth and sixth series coalescent at base; humeri timiid and prominent frontalis Lee. Body much more robust, the elytra not quite twice as long as wide; punctures of the elytral series deep throughout, although small in apical half as usual; fifth and sixth series widely separated at base; humeri not tumid : sharpi Csy. Weevils of the Nevada Test Site I have studied specimens of this genus from Arizona, Nevada, and California, and agree \vitli Sleeper (1960) that Casey's Y. grossus is a valid species. The Nevada and Arizona specimens fit LeConte's description of Y. frontalis. Yuccaljorus sharpi Casey is a Mexican species. Yuccahorus lentiginosus specimens, due to the "elytra dull and with many of the punctures of the intervals surrounded by a pale yellowish-white modifica- tion of the surface, the punctures along each side of the pronotiim also so affected," smaller in size, with finer sculpturing and elytra only a fourth wider than the prothorax separates it as a species, in my opinion, from Y. grossus. Anderson ( 1948 ) reports that Barber con- sidered the galleries of Y. lentigino.sus similar to Dendroctonus. On the basis of the larvae, Boving and Craighead ( 1931 ) were able to separate the Curculionidae and Scolytidae. Crowson (19.55) includes the Scolytidae and PlatyjDodidae in the family Curculionidae. Yuccahorus frontalis (LeConte) Reference. LeConte, 1874:70. Morphological characteristics. Bodv black, shining; head sparsely, coarsely punctured, ros- trum straight, as long as prothorax, coarsely and densely punctured, sulcate and sparsely setose each side; frontal fovea distinct. Eyes contiguous beneath, widely separated above; prothorax longer than wide; apex tubularly constricted, disc coarsely but not densely punctured. Elytra with rounded humeri, sides parallel, striae finely punctiu-ed, intervals flat, with few small punc- tures; lateral margins and under-surface of pro- thorax densely and coarsely punctured. Legs slender, front tibiae with several small teeth on inner edge. Length 12.5 to 14.2 mm, width 4.9 to 5.5 mm. Plant relationship. Fi\e specimens were col- lected in a mixed community. Area 2.3, in August of 1961, 1964, and 1965. NOTES ON THE BIOLOGY OF SOME NEVADA TEST SITE WEEVILS The life histories and host plants of the weevils of the Great Basin are poorly known. In this study an attempt was made to bring to- gether information on the biologies of the weevils of this report. Many of the species dealt with are represented by one or not more tlian three or four specimens. This, no doubt, is due to lack of intensive collecting, such as beating and sweeping of the shrubs and smaller plants. The apterous species found on Atriplex, Coleo- gijne, Grayia. Larrea, Kochia, Lt/cium, Fran- scria and other shrubs of the area are not easily collected by traps or lights. They are taken main- ly by beating or close examination of the plants and by hand picking. I have searched the literature to find the re- corded host plants of the species discussed in this study but have met with little success. What information 1 have gleaned from the literature or the field notes taken at the Nevada Test Site are presented here. Species of Trigonorhiniis (Brachi/tarsus) were reported by Blatchley and Leng (1916) as breeding in stored corn, peas, and c-owpeas; stems of wide rye and in the smut of com and wheat; also taken on ragweed. Ambrosia artcmi- siaefolia, and from the pod of the bladdernut, Staphylcd trifolia. According to Valentine (1960), "Trigonorhinus species are plant feeders Hke the majority of the family, but the generical- ly distinct Old World species of Anthrihus Fos- ter, 1771 (formerly Brachylarsus Schoenherr, 1823), have larvae which are predaceous on the eggs of certain lecaniine scale insects." The speci- men collected at Mercury was on burr-sage, Franscria acanthicarpa. No reports of plant association for the species Thinoxenus ncvaclensis and Thricolepis inornata iiave been found in the literature. At the Nevada Test Site T. nevadensis was taken in a Pinyon- Juniper Community, but the specific host plant is unknown. Thinoxenus inornata was taken on Gamble oak at the test site; it is common on oaks in Utah. The Eucyllus species at the test site are wide-spread in most of the communities. One specimen was taken on winter-fat, Eurotia Janata. I have not found any reference to the host plants for this geims. Aragnomus hispiduhis specimens were taken on Coleogt/nc, Grayia, and Larrea plants. No other references to plant hosts for this species have been found. Van Dyke re- ported collecting Aragnonnts sctosus by beating manzanita. The Cn/ptolepidtis species are typical of this .xerophytic region. Some of the species such as C. Iceclii, C. planifrons, and C rugicoUis were reported by Ting (1940) as living on Atriplex and Sarcobatus vermictilatus. At the test site C. leechi, C. nevadicus, C. aridtis, and C. cazieri were taken in Atriplex associations. One speci- Bhicham Young University Science Bulletin men of C. nevadicus was taken on evening prim- rose, Oenothera californica. Other short-nosed weevils of the genera Ophrijastes, Orimodema, Paracimboccra, MiJo- deriis, and Dirotognathii.s are closely associated with the desert flora of the Great Basin. Fall, Van Dyke, Casey, Horn, and Davis, however, did not record plant hosts for this complex. Ting (1940) collected the holotype female of P. artemlsiae on Artemisia sp. Ophrijastes varlus and O. geminatus are common species on Larrea divaricata and Atriplex canescens in the test site area as well as areas of the Great Basin where these plants are found. Collier, according to Ting (1940), observed Paracimboccra artemisiae injuring the twigs of apple trees in Grand Valley, Colorado, in April, 1911. Orimodema protracta and Dirotognathus sor- didus were collected in the Pinyon-Jimiper com- munity, but a definite plant host was not deter- mined. Miloderes merctiri/ensis was collected on Sphacralcea ambigua and Grai/ia spinosa. One-half of all the specimens of weevils t-on- sidered in this study are Cleomis denticollis which was described from a series of five speci- mens collected at Peach Springs, Arizona. Casey ( 1891 ) did not indicate with what plants this species may be associated. The tribe Cleonini has adults of many species associated with water plants. According to Blatchley and Leng (1916) the larvae feed upon different kinds of Pohj- gonum (smart\veed) and other plants such as dock, ragweed, and wild sunflower. More than a hundred specimens of C denticollis were col- lected in the Grayia-Lycium community, and some specimens were taken on Htjmenoclea fas- ciculata and Larrea sp. at the test site. The host-plant records for Smicronyx imbri- catiis as reported by Anderson (1962), follow: Gutierrezia liicida, Pluchca sericea, Chn/sotham- nus speciosus, Prosopu.s juliflora, Covillea tri- dentata. Pinus edulis, and Pinus monophijUum. This species is common in the Mercury area, and Table 6 shows the wide variety of plants in most of the communities upon which S. imbrica- tits feeds. Promecotarsiis densus, which is includ- ed in the subtribe Smicronychini, has similar food plant preferences to that of species of Smicronyx, having been collected on Artemisia tridentata and Jtiniperus osteosperma. Larvae of species of the genus Apion were reported by Blatchley and Leng (1916) "to feed, for the most part, on seeds, principally those of legumes, though some form galls on the stems and leaves of plants, others knots on the roots, while a few bore into the pith and form a kind of cocoon of the gnawed particles." Specimens of A. sordidum are reported as forming galls on Artemisia. Specimens of A. varicorne, which is a wide-spread and variable species, were taken on Dalea pohjadenia and Eriogomim nodosum. Species of Aidetobius are reported by Blatch- ley and Leng ( 1916 ) as occurring on sweet- fern, Comptonia perigrina, bayberry, Mijrica cer- ifera, and leather-leaf, Chamaedaphne calyctd- ata. At the Nevada Test Site A. humeralis was collected on Ephedra nevadensis. Aidetobius sp. wa,s also found on E. nevadensis and Chrijso- thamnus viscidiflorus. Myrmcx 1. lineata was collected in a Larrea community. Magdalis lecontei tenebrosa was collected on Pinus monophijlla at the test site. Buchanan ( 1934 ) described M. piceae from specimens reared from Colorado blue spruce, Picea pungens glauca. All the species of Magdalis breed in the bark and wood of various coniferous and decidu- ous trees. All the species reported in the tribe Anthono- mini in this study breed and feed in the seeds of plants. Blatchley and Leng (1916) record them as living in the buds, flowers, fruits, and seeds of plants. Some species are plant-gall form- ers. Tlie larvae pupate in the feeding cell in- stead of entering the ground. Macrorhoptus his- pidus was collected on Sphaeralcea sp., Grayia spinosa and Artemisia sp. at the test site. An- thonomtis peninsularis was collected on Sphaer- alcea sp. and Ephedra nevadensis; A. haematop- iis confusus occurs on Juniperus sp. and Artem- isia tridentata; A. ornatulus was collected on Eurotia lanata; A. sphaeralceae was found on eight different plant species (Table 8); A. cycliferus was collected on species of Chryso- thamniis, Artemisia and Chaenactis; A. tenuis occurs on five different plant species at the test site. Epimechus gracilis was taken from Chryso- thamniis viscidiflorus. Brachyogmus ornatus breeds in the flowers and seeds of Lijcium pal- lidum at the test site. Tychus prolixus was collected on Astragalus lentiginosiis at the site. Zascelis irrorata occurs in the Juniper and Grayia-Lycium communities. Ceutorhynchini species, both native and intro- duced, are found in mesophytic to hydrophytic conditions. The larvae feed on "seeds, seed- stalks, or the stems of plants" (Blatchley and Leng, 1916). At the test site, Ccutorhynchus ad- junctus was collected on Stanleya pinnata and on an unknown plant species near Cane Springs. Ccutorhynchus tescorum was collected on prickle-poppy, Argemone corymbosa. Weevils of the Nenada Test Site 31 Tlie genus Omjchobaris in the tribe Madarini is represented by two species in tliis report. Species of this genus are found mainly in the western dry desert regions. Fall (1913) reports that he has taken O. densa on "flowers of a low fleshy-leaved plant just above the lieach near San Diego." Seven specimens of O. mystica col- lected in Area 5 at the test site were on Opuntia acanthacarpa. Specimens of Omjchoharus sp. near depressa were collected on five different species of plants (Table 10). LeConte (1874) described Yuccaborus fron- talis from a specimen found under a bark of Yucca in the Mohave Desert. At the Nevada Test Site this species was collected in a mixed community. REFERENCES Allred, D. M., D E. Beck, and C, D. Jorgensen. 1963a. Biotic communities of the Nevada Test Site, Brig- ham Young Univ. Sci. Bull, Biol. Ser., 2(2):l-52. . 1963b. Nevada Test Site study areas and specimen depositories. Brigham Young Univ. Sci. Bull, Biol Ser., 2(4): 1-15. Allred, D, M., D. E, Johnson and D E. Beck. 1965. A hst of some beeflies of the Nevada Test Site. Great Basin Nat., 25( 1-2) : 5-11. Anderson, D. M. 1962. The weevil genus Smicronyx in America, north of Me.vico (Coleoptera: Cur- culionidae). Proc. U,S, Nat. Mus., 113:185-372. Anderson, W. H. 1948. Lar\ae of some genera of Calendrinae ( Rhsnchophorinae ) and Strombos- cerinae (Coleoptera: Curculionidae ) . Ann. Entomol Soc. Amer., 41:413-437. Amett, R. H. 1960-1962. The Beetles of the United States; Fascicle 108; Curculionidae. Blatchley, W. S.. and C. W, Leng, 1916, Rhyncho- phora or weevils of north eastern America. Nature Pub. Co. Boheman, C. H. 1859, Coleoptera. Species novas de- scripsit, Kongliga Svenska Fregatten Engenies resaomkring Jorden, Insecta 1-218. Boving, A. C, and F. C. Craighead. 1931, An illus- trated synopsis of the principal larval forms of the order Coleoptera. Brooklyn Entomol. Soc, Brook- lyn, New York. Bradley, J. C. 1930. A manual of the genera of beetles of America north of Mexico. Daw, lUston and Co., Ithaca, New York, Bruhn, A. F. 1947. The external male genitalia of some Rhynchophora. Great Basin Nat., 8:1-35. Casey, T. C. 1888. On some new North American Rhynchophora. Ann. New York Acad. Sci., 4:229- 296. . 1891, Coleopterological notices. Ann. New York Acad. Sci., 6:176-180. , 1892. Coleopterological notices, Ann. New York Acad, Sci., 6:359-711. Crowson. R, A. 1955, The natural classification of the families of Coleoptera. Nathaniel Lloyd & Co., Ltd., London. Davis, A. C. 1947. Review of the weevils of the tribe Ophryastini of America, north of Mexico. Proc. U. S. Nat. Mus., 96:483-551. Dethlefsen, E. S. 1954. Revisional notes on the genus Brachytarsoides Pierce (Coleoptera, Platystonidae ) . 1. New North American species and subspecies. Wasmann J. Biol, 12(l):53-62. Dietz, \V. G. 1896, Revision of the genera and species of Ceutorhynchini inhabiting North America. Trans. Amer. Entomol. Soc, 23:387-480. . 1891. Revision of the genera and species of Anothonomini inhabiting North America. Trans. Amer. Entomol. Soc, 18:177-276. Fall, H. C. 1898. Revision of the species of Apion of America north of Mexico. Trans. Amer. Entomol. Soc, 25:105-184. . 1913, A brief review of our species of Magdalis. Trans. Amer. Entomol. Soc, 39:23-72. Fall, H. C, and T. D. A. Cockerell. 1907. The Cole- optera of New Mexico. Trans, Amer, Entomol. Soc. 33:260. Germar, E. F. 1817. Magazin fiir entomology (Ger- mar), Halle 2:339-341. . 1821. Genera quaedam curcuUonitum pro- posita, et speciebus observatis iUustrata. Mag. En- tomol, 4:291-345. 1824. Insectorum species novae ant minus cognitae, descriptionibus illustrate. Herbst, J. F. W. 1797. Natursystem Allen Bekannsten in-und auslandischen insecten, etc. Von C. G. Jab- lonsky, Beriin, 7:100. Horn, G, H. 1873. Contributions to a knowledge of the Curcuhonidae of the United States. Proc. Amer. Philos. Soc, 13:407-469. 1876. The Rhynchophora of America north of Mexico. Proc. Amer. Phil. Soc. 1.5:35. 1895. Coleoptera of Baja Cahfomia, Supplement 1. Proc. California Acad. Sci., 5:256- 257. Kissinger, D. G. 1964. Curculionidae of America north of Mexico. A key to the genera. South Lan- caster, Massachusetts. LeConte. J. L. 1853. Description of some new Cole- optera from Texas, chiefly collected by the Mexican Boundan' Commission. Proc. Acad. Nat. Sci. Phila.. 6:4,39-448. 1874. Description of new Coleoptera from the Pacific Slope. Trans. Amer. Entomol Soc, 5:43-72. LeConte, J. L., and G. H. Horn. 1876. The Rhyncho- phora of America north of Mexico. Proc Amer. Phil. Soc, 15:1-455. 32 BKIGHA^r ^ouNc: Univehsitv Science Bulletin Leng, C. W. 1920. Catalogue of the Coleptera of America north of Mexico. Supplements 1-5; 1927- 1947. Linell, M. L. 1897. New genera and species of North American Curculionidae. J. New York Entomol. Soc, 5:49-56. Marshall, Sir Guy. 1932. Notes on the Hylabiinae (Coleoptera Curculionidae). Ann. Mag. Nat. Hist., 9(52):341-355. Munz, P. A., and D. D. Keck. 1959. The California flora. Univ. of California Press, Berkeley. Pascoe, F. P. 1872. Contributions toward a knowledge of the Curculionidae. Joum. Linn. Soc. 2^1., 11: 440-492. Pierce, W. D. 1909. Studies of North American weevils. Proc. U. S. Nat. Mus.. 35:325-364. . 1930. Studies of the North American wet vils belonging to the subfamily Platvstomoidea. Proc. U. S. Nat. Mus., 77(17):l-34. Sleeper, E. L. 1957. Notes on the genus Macrorhop- ttis LeConte (Coleoptera Curculionidae). Ohio J. Sci., 57(2):70-74. . 1960. Notes on the Curculionidae. II. Ohio J. Sci., 60:83-88. Smith, ]. B. 1884. Synopsis of the Apioninae of the United States. Trans. Araer. Entomol. Soc., 11:41- 68. Schoenherr. C. J. 1826a. Curculionidum de.sDositio methodica. Lipsiar., p. 338. . 1826b. Curculionidum desposito methodi- cacum genera characterobus, 145. . 1833. Genera et species Curculionidum, Sturm. J. 1826. Catalog meiner insecten Sammlung, Erstes theil, Kafer, Numberg. Tanner. V. M. 1927. A preliminary study of the genitalia of female Coleoptera. Trans. Amer. En- tomol. Soc, 53:5-50. . 1934. Studies in the weevils of the western United States. No. I. Proc. Utah Acad. Sci., Arts, and Letters, 11:285-288. . 1942. Studies in the weevils of the west- em United States, No. V; A New species of Milo- deres. Great Basin Nat., 3:23-26. Tanner. V. M., and W. A. Packhiun. 1965. Tenebri- onidae beetles of the Nevada Test Site. Brigham Young Univ. Sci. Bull.. Biol. Ser., 1(6): 1-44. Ting, P. C. 1940. Revisional notes concerned with Cimhocera and related genera. Bull. Southern Cali- fornia Acad. Sci., 39(2) : 128-159. Van Dyke, E. C. 1930. New Rhynchophora (Coleop- tera) from western North America. Pan-Pacific Entomol., 6(4) :149-165. . 1936a. New species of North American weevils in the family Curculionidae, subfamily Brachyrhininae, V. Pan-Pacific Entomol., 12:73-86. 1936b. New species of North American weevils in the family Curculionidae, subfamily Brachyrhininae, V. Pan-Pacific Entomol., 12:73-86. . 1938. New species of Rhyncophora of western North America. Pan-Pacific Entomol., 14:1. 1939. The origin and distribution of the 1843. Genera et species Curculionidum, coleopterous insect fauna of North America. Sixth Pacific Sci. Cong., 4:255-269. Valentine. B. D. 1960. The genera of tlie weevil family Anthribidae north of Mexico (Coleoptera). Trans. Amer. Entomol, Soc. 86:41-85. Voss, E. 1934. Monographic des Rhynchitinae. Tribus Auletini. Stettinnes Entomol. Ziet., 94. WoUaston, T. V. 1861. Ann. Nat. HLst., 8(3):102. Weevils of the Nevada Test Site 33 Fig. 11. Map ot the area.s of the Nevada Te,st Site Bhigham Young University Science Bulletin A/ |Y^ Brigham Young University Science Bulletin ■-^^OWP^ zc UNIVERSITY A COMPARATIVE OSTEOLOGICAL STUDY OF TWO SPECIES OF COLUBRIDAE (PITUOPHIS AND THAMNOPHIS) by ROBERT E. BULLOCK and WILMER W. TANNER BIOLOGICAL SERIES — VOLUME VIII, NUMBER 3 ^y DECEMBER, 1966 I BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Dorald M. Allred, Department of Zoology and Entomology, Brigham Young University, Provo, Utah Associate Editor: Eabl M. Christensen, Department of Botany, Brigham Yoimg University, Provo, Utah Members of the Editorial Board: J. V. Beck, Bacteriology C. Lynn HArwAUD, Zoology W. Derby Lavs^s, Agronomy Howard C. Stutz, Botany WiLMER W. Tanner, Zoology, Chairman of the Board David L. Hanks, Botany Ex officio Members: Rudger H. > Science Ernest L. Olson, Chairman, University PubHcations Rudger H. Walker, Dean, College of Biological and Agricultural Sciences The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Publications, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah. Brigham Young University Science Bulletin A COMPARATIVE OSTEOLOGICAL STUDY OF TWO SPECIES OF COLUBRIDAE (PITUOPHIS AMD THAMNOPHIS) by ROBERT E. BULLOCK and WILMER W. TANNER BIOLOGICAL SERIES — VOLUME VIII, NUMBER 3 DECEMBER, 1966 s LIEf^ARY FEB 3 ^967 UNIVERSITY TABLE OF CONTENTS Page INTRODUCTION 1 MATERIALS AND METHODS 2 VERTEBRAL COLUMN 3 Atlas 7 Axis 7 First Thoracic Vertebra 7 Other Thoracic Vertebrae 9 Lumbar Vertebrae 9 Anterior Caudal Vertebrae 9 Middle Caudal Vertebrae 13 Posterior Caudal Vertebrae 13 Ribs 13 SKULL 15 Premaxilla 15 Nasals 15 Septomaxillae 15 Vomers 15 Frontals 15 Prefrontals 17 Parietals 17 Postorbitals 17 Supraoccipital 17 Prootics - 17 Exoccipitals 19 Basioccipital 19 Basisphenoid 19 MaxiUae 19 Ectopterygoids 19 Palatines 19 Pterygoids 21 Supratemporals 21 Quadrates 21 Mandible ^ 21 Stapes 22 Hyoid Apparatus 22 OSTEOLOGICAL COMPARISONS 22 Vertebral Column 22 Skull .■. 23 DISCUSSION 25 SUMMARY AND CONCLUSIONS 27 LITERATURE CITED 28 ABBREVIATIONS USED IN PLATES 28 LIST OF FIGURES Figure Page 1. Atlas 4 2. Axis 5 3. First Thoracic and Thoracic Vertebrae 6 4. Thoracic and Lumbar Vertebrae 8 5. Anterior Caudal Vertebra 10 6. Middle Caudal Vertebra 11 7. Posterior Caudal Vertebra and Rib 12 8. Skull, Dorsal View 14 9. Skull, Lateral View 16 10. Skull, Ventral View 18 11. Hyoid Apparatus, Mandible, and Cranium 20 A COMPARATIVE OSTEOLOGICAL STUDY OF TWO SPECIES OF COLUBRIDAE {PITUOPHIS AND THAMNOPHIS) by Robert E. Bullock' and Wilmer W. Tanner- INTRODUCTION Snake anatomy has been genei-ally neglected. Although some early writers recorded observa- tions on the anatomy of snakes, no attempt has been made to undertake a comprehensive study, and Cole (1944) has rightfully referred to this section of vertebrate anatomy as an "almost vir- gin field." Snakes deserve more intensive in- vestigation because of the considerable number of adaptations demonstrated by them, their peculiar types of locomotion, and methods of feeding. It is important to comprehend their anatomical specializations if we are to under- stand the evolutionary trends among modern reptiles. It also is evident that comparative de- scriptions of serpents, thus far based mainly on external characteristics, need to be supplemented by accounts of their internal anatomy. The purpose of this study is threefold: (1) To work out in detail the osteology of two common members of the family Colubridae. (2) To compare the bones and bone struc- tures of the two species in order to de- termine their anatomical similarities and differences, and to use these comparisons in determining the different speciahza- tions that have been made by two com- mon serpents. (3) To relate the structural differences of these genera, as far as possible, to some of the other generic groups previously reported. Our original intent was also to include the myology of the head, neck, and anterior tnmk region (first 10 vertebrae). This proved to be a major effort in itself, and such a report will ap- pear separately at a later date. The gopher snake, Pituophis catenifer Blain- N'ille, and the plains garter snake, Thamnophis radix (Baird and Girard), were chosen for this study because of their moderately large size and relative abundance. One of them (P. catenifer) cxjnstricts its prey. The other ( T. radix ) swallows it alive. This difference in mode of feeding may be responsible for some structural modifications. Although it is not the purpose of this study directly to resolve any phylogenetic or ta.xonomic problems, it is hoped that it will encourage other anatomical studies of reptiles, add to our present understanding of homologies, and indicate im- portant internal phylogenetic and taxonomic structures as suggested by Robinson and Tanner (1962). Although no previous investigator has studied the osteology of the two species included in this study, the general gross anatomy of various other snakes has been known for a long time. Ahrenfeldt (1955) stated that as far back as 1573 there were some fragmentary accounts of snake anatomy published in Europe, but Cole ( 1944 ) mentioned that it was not until 1683 that the first workmanlike and relatively accurate description of serpent anatomy was made by Edward Tyson on a "Timber Rattlesnake" said to have been collected in the West Indies. Owen ( 1866 ) gave one of the first detailed accounts of the osteology and myology of snakes based on his work with the species Crotalus horridtis, Pijthon tigris. Boa constrictor, Naja tripudians, and Deirodon scaber. Although Owen's work on this section of the vertebrates is a general account by our standards, in his time it was a major contribution to this area of anatomy. Another work dealing with snake anatomy is a laboratory dissecting guide by Kel- licott (1898) for the genus Heterodon. Unfor- tunately, the descriptions lack detail and are therefore somewhat superficial and in many areas incomplete. Huxley (1871), Gegenbaur (1878), Hoffman (1890), Sedgwick (1905), Wiedersheim and Parker (1907), VVilliston (192,5), Kingsley (1917), Goodrich (1930), and Versluys (1937) were some of the early textbook writers who ^Lethbridge College. Lethbridge, AJberta, Canad ^Department of Zoology and Entomology, Brigha Young University, Provo, Utah. Brigham Young University Science Bulletin dealt with the subject of snake osteology. Most of these writers restricted their studies to the skull of serpents, and on occasion included one or more generalized vertebrae. Romer's "Osteo- logy of the Reptiles" (1956) deals only briefly with serpents, but does bring some of the termi- nology and many of the homologies up to date. Two volumes by Ludicke (1962, 1964) de- voted entirely to the anatomy of serpents deal rather extensively with the developmental and adult anatomy of several species of snakes, in- cluding a few colubrid species. Many of the descriptions and comparisons in these works are of a general nature and, therefore, are of limited value in making comparisons with the colubrid species involved in this study. Sood (1941, 1948) published two papers dealing with the vertebral column of serpents. The first is concerned with the caudal vertebrae of the sandsnake, Eryx johnii (Russell), and illustrates many unique peculiarities in its verte- bral anatomv. The second studv includes both minute and gross vertebral anatomy for several other Asian species. An extensive work by Edgeworth (1935) describes and compares the development of the cranial muscles and associated skeletal structures in forms ranging from the fishes to the mam- mals. Haas (1930) and Cowan and Hick (1951) also dealt with the musculature of snakes, but included skeletal structures as well. Albright and Nelson ( 19.59 ) dealt with the cranial osteology and the musculature of the colubrid snake Elaphe obsolcta quadrivittata. Perhaps more osteology has been done with venomous species, particularly with the teeth (fangs) and sk-ull. The work of Klauber (1956: 712-743) is a good example. We made no major attempt here to e.xhaust the references which may refer to snake anat- omy. Those referred to above represent some of the more important works that we have seen. We were greatly surprised when we found rela- tively few studies dealing with either of the two common colubrid genera Pittiophis and Tham- nophis. Our prime concern, therefore, is to provide detailed descriptions and drawings which wiU set forth the skeletal anatomy as we found it in these genera. MATERIALS AND METHODS This comparative study utilized the species Pituophls catenifer deserticola Stejneger and Thamnophw radix haydeni Kennicott. The four- teen specimens of P. c. deserticola used were collected in Grand, Kane, Utah, San Juan, Gar- field, and Uintah Counties, Utah. The twelve specimens of T. r. haydeni came from Warner County, Alberta, Canada. However, two speci- mens of P. catenifer sayi Schlegel from Medicine Hat, Alberta, Canada, two T. elegans vagrans Baird and Girard from Utah County, Utah, and four T. sirtalis parietalis Say from Salt Lake County, Utah, were also dissected and studied. These additional species are similar within their respective genera; therefore, a description of these is not included. The skeletons were prepared in several dif- ferent ways, depending on whether fresh or preserved snakes were used, and upon the de- gree of articulation required. Disarticulated skeletons were prepared from fresh material by bacterial action. The snake was skinned, evis- cerated, and plac-ed in a container with enough water to cover the specimen so as to accelerate the decomposition process. After a few days in hot weather, the macerated material was placed on a fine mesh screen and a jet of water was used to clear away the remaining tissue. Some of the bones were bleached in a 3% solution of hvdrogen peroxide for ten to twelve hours. Most of the articulated skeletons were pre- pared from fresh material. The fresh, skinned, and eviscerated snake was placed in a 25% solution of ammonium hydroxide for one week, then boiled for t^vo to ten minutes until the tissues were loose but the ligaments still intact. A small jet of water was used to clear the soft tissue from the vertebral column. This jet proved to be too harsh on the more delicate and loosely connected bones of the skull. To remove the deeper skull muscles it was necessary to use forceps under a stereoscopic microscope. This latter method was essential in preparing skulls without distortion or loss of small bones. A few snakes preserved in alcohol were utilized in the preparation of skeletal material. These were cleaned by placing them for two days in a solution of two ounces of trisodium phosphate to each quart of water. Tlie material was then boiled in water for varying lengths of time, the time determined by the type of skeletal preparation desired. Soaking the preserved OsPEOLOGv OF Snakes material in concentrated ammonium livdroxide instead of trisodium phosphate before boihng proved to be almost as satisfactory. Specimens were studied and drawings made to scale with the aid of a hand micrometer. Literature dealing with the complete osseous skeleton of the snake is fragmentary and limited. Because the literature to date, as far as we were able to ascertain, reveals no account of the osteology of either P. catenifer or T. radix, the homologies were worked out and derived from work done by Owen (1866), Kellicott (1898), Romer (19.56). Sood (1941, 1948), and the more recent works by Albright and Nelson (19.59), and Ludicke (1962, 1964). The terminology for the skull structures used herein was adopted chiefly from Romer (19.56), and the terminology for the vertebral column from Sood ( 1941, 1948). The two snakes studied are anatomically similar; therefore, to avoid unnecessary repeti- tion a complete description of P. catenifer was prepared, but T. radix is discussed only in con- nection with comparative structures that differ in the two forms. Tlie plates were prepared with sufficient detail to aid in the comparisons of similar structures in the two species. VERTEBRAL COLUMN In P. catenifer as in other snakes, the verte- brae are numerous and procoelous, with ball and socket articulations wliich allow free movement. All of the vertebrae articulate with ribs except the atlas, axis, first thoracic, and caudals. The successive vertebrae not only articulate by the usual pre- and post-zygapophyses and by the procoelous centra, but also by additional articu- lations such as the zygosphenes and zygantra. The division of the ophidian vertebral col- umn into well-marked regions is less distinct than it is in other higher vertebrates. Owen (1866), Sedgwick (1905), Reynolds (1913), Williston (1925), and others considered the column to be made up of two regions — a pre- caudal region, and a caudal region. Five regions were distinguished by Roche- brune ( 1881 ) — the cervical, thoracic, pelvic, sacral, and cocygeal. Although these regions can be distinguished one from another, the differ- ences are slight and variable. Because this classi- fication was somewhat superficial and could not be regarded as ecjuivalent to the regions of the vertebral column of other vertebrates, it was presumably not adopted by later workers. Sood (1941, 1948) retained the division which separated the column into two regions (precaudal and caudal), but distinguished sev- eral subregions of each as follows: I. Tlie Precaudal Region A. Cervical Subregion, composed of the first two vertebrae, atlas, and axis. B. Thoracic Subregion, consisting of all the vertebrae that follow the axis and bear prominent hypapophyses. C. Lumbar Subregion, consisting of the vertebrae situated between the thor- acic subregion and the caudal region. The hypapophyses are reduced in size or completely lacking in this area. II. The Caudal Region A. Anterior Caudal Subregion, situated immediately behind the precaudal vertebrae. Vertebrae are provided with both fi.xed and articulating bi- furcated ribs called processi costo- transversii or lymphapophyses. B. Middle Caudal Subregion, containing fixed ribs which are unforked and differing from the posterior caudal subregion in the absence of haema- pophyses. C. Posterior Caudal Subregion compris- ing vertebrae with a pair of flat, platelike haemapophyses on the ven- tral aspect of their centra. This area shows a gradual reduction in the size and development of the vertebrae and vertebral processes from anterior to posterior until they are repre- sented near the end of the tail by extremely short vertebrae with al- most wholly vestigial processes. Because the division of the vertebral column as outlined by Sood is in general use, the pre- ceding vertebral classification will generally be followed; however, a few modifications have been made in order to adapt it to the species in- volved in this study. Bhigham Young University Science Bulletin A.INT.F. A.NEU.F. A.NEU.F Fig. 1. Atla-s. Pituophii catenifer, A-D, 9X: A, anterior; B, posterior; C, dorsal; D, lateral. Tliumiiophis radix, E-H, 15X: E, anterior; F. posterior; G, dorsal; H, lateral. Osteology of Snakes N.S. ZE. ^.J^. POZ. OPR T.PR. / ^■•.fe.'C^-vt' \ ^^^Yt^ HYP.F. P.HYP. CEN. C X:V\\ P07F ^^^^^ ^-77 N.S. ZE. O.PR. POZ. HYP.F. \fe 1 jp^%, X CEN. A.HYP. _ P.HYP. ^'^- ]\,y^t' P'["°P^"^''"'"'f^'' A-D, 8X: A, anterior; B, posterior; C, dorsal; D, lateral. Thammphis radix E-H l^X: h, anterior; F, posterior; G, dorsal; H, lateral. Bricham Young University Science Bulletin PRZ.F Fig. 3. Pituophts catenifer, A-D, 7X: A-B, first thoracic vertebra: A, dorsal; B, lateral; C-D, thoracic vertebra: C, anterior; D, posterior. Thamnophis radix, E-H, 9X: E-F, first thoracic vertebra: E, dorsal; F, lateral; G-H, poi thoracic vertebra: G. anterior; H, posterior. Osteology of Snakes Fig. 1-A, B, C and D The first cervical vertebrae or atlas differs widely from the general vertebral pattern. It is a relatively narrow ring composed of tliree separate bones fused together by means of su- tures. Tlie two lateral neural arches or neura- pophyses join dorsally as the sagittal neural su- ture; thev form a pair of neurocentral sutures at their ventral borders where they unite with the intercentrum. Tliere is no neural spine, but a nidimentary dorsal crest is developed. The centrum of the atlas actually coalesces with that of the axis leaving the atlas without a true cen- trum, its place being taken by the autogenous hypapophysis (Owen, 1866). Romer (1956) stated that the atlas has been robbed by the a.xis of its centnmi, but has retained its intercentnmi. Anteriorly the intercentrum presents a concave articular surface which articulates with the basioccipital tubercle of the occipital condyle. Posteriorly a similar surface adjoins the ventral surface of the odontoid process, as well as the interior facet of the anterior hypapophysis of the axis. Posteroventrally the intercentrum develops a small conical hypapophysis. The base of each neural arch half possesses an anteromesial, concave articular surface which receives the exoccipital tubercle of the occipital condyle, and a posteromesial surface which ar- ticulates with the dorsal and lateral convex por- tions of the odontoid process. Each neural arch half carries a short postzygapophysis from the lateral border of its dorsoposterior region, and from its ventrolateral border a short conical transverse process points posteriorlv. The neural canal is formed by the dorsal expansion of the neural arch, while the smaller ventrally located condylar or odontoid canal is formed between the mesial concave articular surfaces of the arch and the dorsal surface of the intercentrum. The two canals are partially separated by a tough transverse ligament extending from the dorso- mesial border of one neiu-al arch half facet to the other. The intervertebral foramina are formed when the notches in the posteroventral edges of the neural ring of the atlas come in contact with the anteroventral neural ring notches of the adjacent axis. AXIS Fig. 2-A, B, C and D The axis has a very stout and elongate cen- trum which protrudes anteriorly, forming the odontoid process, and terminates posteriorly as a ball-like articulating condyle. The odontoid process is a separate cone-shaped bone con- nected to the main centrum and the anterior hypapophvsis by a broad, curved suture. The apex of the odontoid has a rounded process which extends through the odontoid cavity of the atlas and rests lightly upon the basioccipital portion of the occipital condyle. The lateral ar- ticular surfaces of the process are c-onvex and are received between and articulate with the pos- teromesial neural arch facets of the atlas. The neural arch of the axis develops an elon- gate, posteriorly-projecting, ribless, transverse process from each side of its base, a partially developed zygosphene from the anterior dorso- lateral border of each side, and a moderately long, posteriorlv-projecting neural spine from its coalesced dorsal surface. The posterolateral ex- pansions of the neural arch form the zvgantrum internally and the postzygapophyses externally. The postzygapophyses contain flat, oval-shaped facets pointing ventrolaterallv from the expand- ed surfaces, whereas the zygantrum has two similarly shaped articulating surfaces excavated from the inner surface of these same expansions, which point in a dorsomesial direction. The post- zygapophyses articulate with the prezygapo- physes of the first thoracic vertebra while the zygentral facets articulate with the zygosphenal surfaces of the same thoracic vertebra. There are two hvpapophyses developed on the ventral surface of the axis. The anterior one is sutured to the ventral border of the odontoid process as well as the anterior border of the pos- terior hypapophysis. It is somewhat triangular in shape with its curved dorsal suture represent- ing the base of the triangle, and its apex point- ing ventrally. The anterior vertical border con- tains an elongate, oval facet which articulates with the intercentrum of the atlas. The posterior hypapophysis is an elongate spinelike process developed on the midventral aspect of the cen- trum which points obliquely posterior. FIBST THORACIC VERTEBRA Fig. 3-A and B The first thoracic vertebra lacks free ribs and is also otherwise modified. It differs from the other thoracic vertebrae in possessing a rather elongate, transverse process much like that of the axis, a fairly long, narrow, posteriorly-pointing, bladelike neural spine, and in the absence of articular surfaces for the attachment of ribs. The transverse process of these vertebrae is consid- ered by Romer ( 19.56 ) to be partially composed of short fused ribs. He therefore states that they can be referred to as "cervical" vertebrae. Other structures developed on this first thoracic verte- Brigham Young Univebsity Science Bulletin Fig. 4. Pituophis catenifer, A-D: A-B, thoracic vertebra, 7X: A, dorsal; B, lateral; C-D, lumbar vertebra, 6X: C, dorsal; D, lateriil. Thamrwphis radix, E-H: E-F, thoracic vertebra, 9X: E, dorsal; F, lateral. G-H, lumbar vertebra, 7X: G, dorsal; H, lateral. Osteology of Snakes bra are similar to the other more typical verte- brae of tliis subregion and are discussed in c-on- nection with them. OTHER THORACIC VERTEBRAE Figs. 3-C and D, 4-A and B The typical thoracic vertebrae are strong and blocklike, being wider than they are long. Tlie number of these vertebrae varied from fifty to fifty-three in the specimens of P. catenifer studied, but averaged approximately fifty-t\vo in number. Tlie centrum is not round, but ratlier compressed dorsoventrally. Its anterior face bears a concavity, the vertebrae being procelous. Tlie socket faces a little ventrad from the greater prominence of the upper border; the prominent ball terminates the back part of the centrum rather more obliquely, its aspect facing some- what upward. Ventrally the centrum bears an elongate median hypapophysis e.vtending pos- teroventrally, and terminating slightly posterior to the condyle of the centrum. Tlie transverse processes are short, bilobed structures arising from the anterolateral portions of the centrum, and extending ventrally in an obliquelv posterior direction. The major portion of each transverse process is covered by the rib articular surface. The dorsal lobe of each process is a convex sur- face, whereas the ventral area is flattened and even slightly concave. The area of the transverse process which extends ventrally and anteriorly below the level of the centrum is referred to as the parapophysis. The neural arches are broad, swollen struc- tures facilitating the arrangement of the zygapo- physes, whose nearly horizontal articular sur- faces are placed far apart from each other at a level not far above the floor of the jaeural canal. The oval, transversally elongate facet of each prezygapophysis is supported by a lateral pro- cess arising from the dorsal aspect of the trans- verse process, and is facing dorsad. Sood ( 1948 ) noticed that each prezygapophysis supported a lateral projection pointing outwardlv and slightly forward, serving as a point of attach- ment for muscles, and seemed to correspond to the metapophysis of mammals. A wedge-shaped process, the zygosphene, is developed from the anterior border of the biuse of the neural spine and bears two smooth, oval, flat articular surfaces. These zygosphenal facets extend from the ventral apex of the wedge (sloping dorsolaterally ) to its dorsolateral bor- ders. This wedge Ls received into the cavity (zygantrum) containing the zygantral facets, wliich is excavated in the posterior ex-j^ansion of the neural arch of the preceding vertebra. The zygosphenal surfaces are adapted to articulate with those of the zygantra. The postzygapophyses and zygantra of the thoracic vertebrae are very similar to those of the axis except for their being slightly larger and more horizontally placed. The neiu-al spine is of moderate height, placed about equidistant in its anteroposterior extent on the vertebra, and is laterally com- pressed and tnmcate. There are typically two sets of foramina in the vertebral column — the intervertebral and the intra vertebral (Sood, 1948). Tlie intervertebral foramina have already been mentioned in con- nection with the atlas and a.xis, but they are found along the entire column between con- tiguous vertebrae. When viewed laterally each foramen is composed of two apertures — a dorso- lateral, superior inters'ertebral foramen and a ventrolateral, inferior intervertebral foramen. The zygapophysial articulation separates the single intervertebral foramen into its superior and inferior external apertures. A feature not noted previously is the presence of several pairs of intravertebral foramina in addition to the pair of minute apertures situated ventrally in the middle of each centrum on either side of the median longitudinal line. These additional in- travertebral foramina are located as follows: a foramen on the dorsal surface of the proximal portion of each metapophysis, a foramen in the middle of the base of each lateral wall of the neural arch, a foramen or pair of foramina on each side of the anterior central articular socket, and several pairs of minute foramina on the roughened ridge dorsal to each postzygapo- physis. These foramina are found not only in the thoracic vertebrae, but arc common to most vertebrae in all of the subregions. LUMBAR VERTEBRAE Fig. 4-C and D The lumbar vertebrae vary in number from 183 to 187 in the specimens studied. These ver- tebrae lack hv-papophyses, but possess a small, rudimentary mid-ventral ridge. Most vertebrae iire slightly larger and heavier than those of the thoracic subregion, but with the exception of lacking hypapophyses, they are morphologically very similar. ANTERIOR CAUDAL VERTEBRAE Fig. 5-A. B, C and D In general structure these five or six vertebrae resemble those of the precaudal series. The Bhigham Young University Science Bulletin POZ.F. Fig. 5. Anterior caudal vertebra, 6X. Pituophk Mtenifer, A-D: A, anterior; B, posterior; C. dorsal; D, lateral. Thamnophis radix, E-H: E, anterior; F, posterior; G, dorsal; H. lateral. Osteology of Snakes 11 '* rJ::^^^.7S^.z:::::'r;s:s::t2:j:r;:s;^:'- — ^' ^"* °^ '■-"'■ Bhigham Young University Science Bulletin HAEM HAEM. POZ.F. HAEM. CO.PR ^'^' LerJl"Z^%hT'''7-^'°J-^'^-^°''''"°'^^"'^''^'^^^^ 1«'^- A' ^"t^™--^ B. dorsal; C, lateral; D, rib, lateral, 5X. Thamrwphis radix, posterior caudal vertebra, E-G, 12X; E, anterior; F, dorsal; G, lateral. Osteology of Snakes 13 region can be distinguislied, however, by the ab- sence of free ribs and tlie presence of fused or articulating bifurcated ribs called lymphapo- physes. In the younger specimens studied the lymphapophyses of the first anterior caudal ver- tebra articulate with the body of the vertebra, as do the preceding ribs of the lumbar verte- brae. In the majority of the older and more ma- ture specimens the articulating surfaces were somewhat fused, and in most incidences the lymphapophyses were not free to move in the same manner as articulating ribs. In a few of the mature specimens the only evidence of the pre- vious articulation is a groove aroimd the base of the lymphapophyses. The lymphapophyses of the remaining vertebrae of this subregion do not articulate or demonstrate any superficial indica- tion of such. These bifurcated structures (lymphapo- physes) are associated with muscle attachment as well as with the large lymph hearts on either flank; the divided processes extend outward, dorsal and ventral to the hearts. Tlie more ven- tral branch is an elongate hornlike process pointing posteroventrally. Tlie dorsal branch is much shorter and arcs in a ventrolateral direc- tion. The dorsal and ventral branches of these vertebrae, however, do show some degree of morphological modification from vertebra to vertebra and from specimen to specimen. Both branches show continuity in becoming reduced in size as they progress posteriorly. The dorsal branch is reduced to the point of becoming non- existent in the middle caudal subregion, but the ventral branch remains in its reduced form throughout the remaining caudal region. The first one or two vertebrae of this subregion have developed a small ridgelike hypapophysis, but are devoid of haemapophyses. In P. catenifer, however, the last few vertebrae of tliis section possess a pair of flat, platelike, curved haemapo- physes projecting ventrallv from the posterior part of each centrum. These structures are con- sidered to be chevrons by Romer (1956), and serially homologous with the hypapophyses of the precaudals. There was no apparent indication of sexual dimorphism in the vertebrae of this (cloacal) area. MIDDLE CAUDAL VERTEBRAE Fig. 6-A, B, C and D These three or four vertebrae differ from the anterior caudals in possessing unforked, fixed ribs which are relatively long and only moder- ately curved. Although in Sood's criteria for this subregion the vertebrae lack haemapophyses, both colubrid snakes involved in this study possessed them. Tlie possession of haemapo- physes by middle caudals makes it rather diffi- cult to distinguish between the last few verte- brae of this group and the first few vertebrae of the posterior caudal subregion. POSTERIOR CAUDAL VERTEBRAE Fig. 7-A, B and C The number of vertebrae involved in this subregion vary greatly from specimen to speci- men; however, an average range is fifty-five to sixty-five. These vertebrae are similar in general pattern to the middle caudals, but they gradual- ly diminish in size until the last few are rudi- mentary. Their processes also show a gradual reduction in size until they are almost vestigial near the tip of the tail. Although the demarcation between these vertebrae and the middle caudal group is some- what arbitrary, it is possible to separate the two areas by the length and direction of the fixed ribs (transverse processes) as well as the struc- ture of the haemapophyses. In this latter caudal group the transverse processes point aiitero- ventrally; the fixed ribs of the middle caudals point either ventrally or posteroventrally, and are longer. The apices of the haemapophyses ap- proach each other mesially until they almost touch in the anterior portion of the posterior caudals, whereas they are farther apart and more rudimentary in the middle caudals of the majority of the specimens studied. About mid- way along the posterior caudals, the haema- pophyses undergo a transition whereby they pro- ject almost straight downward in a posteroven- tral direction with little or no inward curve, re- maining rather vddely separated at their distal ends. Fig. 7-D All of the precaudal vertebrae, except the atlas, a.xis, and first thoracic, articulate with a pair of vertebral ribs ( pleurapophyses ) . The ribs in the middle of the body are longer and heavier than the first and last few pairs. Tliey are terete, curved, pointed, and are so articulated that they move freely in an anteroposterior plane. Each articulates with the anterior edge of the cen- trum, the head being connected with the bilobed transverse process. Although the rib head is single, the articular surface is divided into a flattened ventral portion and an adjacent dorsal cupped area. Tlie rib head does not develop a 14 Brigham Young University Science Bulletin EC. PO. PAL. SM.PR. Fig. 8. Skull, dorsal. A, Pituophis catenifer, 4X. B, Thamnophis radix, 5X. Osteology of Snakes 15 true capitiilum or tubt'rciilum as is characteristic a posterodorsally-directed costal process for the of higher vertebrates. Distal to the head there is attachment of muscle fibers. SKULL Tlie skull of P. catcnifcr is well ossified and the bones are dense; the cranium is relatively long and broad, and the brain cavity extends Ix'tween the orbits. The apparent large size of the head is partially due to the arrangement of the supratemporals, cjuadrates, and mandibular structures. Tlie bones making up the skull are for the most p;irt loosely articulated, thus making possible a greater flexibility in the mouth region. PREM AXILLA Figs. 8-A, 9-A. 10-A The premaxilla is a single, small median bone which terminates the snout. Its anterior surface is an inverted Y-shaped ridge, the prongs of which extend in a ventrolateral direction. Tlie dorsal process of the ridge curves posteriorly and wedges between the anterior portion of the t^vo nasals. From the ventral crotch of the "Y" there is a horizontal platelike process, terminally bi- furcated, ex-tending posteriorly. Its dorsal surface articulates with the ventral surfaces of the an- terior septomaxillary processes, thus forming the roof of the anterior portion of the mouth. Tlie prema.villa does not bear teeth. Figs. 8-A, 9-A, 10-A From a dorsal view the articulated nasals appear as an ov^alshaped structure between the premaxilla and the frontals, and are loosely con- nected to them by connective tissue. The nasals separate anteriorly to form an interspace, which receives the dorsal process of the premaxilla, but remain connec-ted posteriorly, forming a pointed process which is received into the an- terior cleft between the two frontals. Dorsally the nasals do not articulate with the frontals. The nasals sheath the nasal cavities dorsolaterally, and form a double median septum between them. This vertical septum extends postero\'en- trallv as a process which comes in contact with the fused anteroventral borders of the frontals. The posterior portion of the septum is in contact with the dorsal surface of the septoma.xillae. SEPTOMAXILLAE Figs. 8-A, 9-A. 10-A The elongate, horizontal, platelike septo- maxillae form the floor of the internal nares. Tliey are in contact dorsally with the nasals, anteriorly with the premaxillae, ventrally with the vomers, and posteriorly with the frontals. The septoma.xillae lie close together near the mid-dorsal line, but do not suture. Their posterior ex-tremities turn ventrad and come in contact with the frontals. A pair of horizontal winglike processes extend laterally from about the middle of each bone, become narrow, and turn dorsally until they almost touch the descending dorso- lateral borders of the nasals. Figs. 8-A, 9-A, 10-A The edentulous vomers are connected to the ventral surfaces of the septomaxillae; at this union they fomi a pair of hollow, spherical vomeronasal organs which open by paired ori- fices into the buccal cavity. The inner margins of the vomers do not suture, but are joined to adjacent elements only by connective tissue. Each vomer has three vertically-flattened pro- cesses extending from its flat mesial border — an anterior process connected to the septomaxilla, a posterior process connected to the mesial sur- face of the posteroventral process of the nasal and the posterior process of the septomaxilla, and a larger posteroventral process with a large triangular fenestra occupying most of its area. The posteroventral process does not articulate with any skull structures, but is connected by connective tissue to other adjacent elements. FRONTALS Figs. 8-A, 9-A, 10-A The frontals are highly developed and form a complete enclosure for the anterior portion of the brain. Tliey remain separate, but are joined mesially by the sagittal suture. The dorsal sur- face of each frontal is flat, subquadrate, longer than broad, and there is a groove in each, parallel with the indented orbital edge. This groove is perforated with several supraorbital foramina in the adult forms. In younger snakes some of these foramina are not completely formed, but are mere indentations from the orbital margin. The anterolateral edge has a small, depressed articu- lar surfac-e to which the prefrontal is loosely at- Brigham Young University Science Bulletin SOC. PR SM.PR. Fig. 9. SkuU, lateral. A. Pituophis catcnifcr, 4X. B, TJwwnoplm radix, 5X. Osteology of Snakes 17 tached. The lateral surface, considered to be the orbitosphenoidal plate of the frontal, joins the dorsolateral margin of the frontal at an acute angle, thus forming a major portion of the mesial wall of the orbit. Ventrallv the descending walls rest upon the presphenoidal prolongation of the basisphenoid, completing the orbital septum and cranial floor. Each wall is notched posteroven- trally where it sutures with the anterior wall of the parietal, thus forming the anterior jx)rtion of each orbital or optic foramen. Through these foramina pass the optic nerves, eye-muscle nerves, and blood vessels. Anteriorly, where the frontals contact the nasals and septomaxillae, two large canals or ethmoid foramina, to facili- tate the passage of olfactory nerves, are formed by way of a vertiaxl, median double lamina (one from each frontal). PREFRONTALS Figs. 8-A, 9-A, 10-A Although some early workers considered the lacrimals to be present in snakes, Romer (1956) stated that "lacrimals are absent; a foramen or groove for the lacrimal duct is present in the prefrontal." According to Gregory (1913), the prefrontal of reptiles is not homologous with the lacrimal of mammals. From a lateral view each prefrontal is an irregular cone-shaped structure with a laterally compressed anterior process (forming the apex of the cone) extending laterally to the posterior portion of the nasal, septomaxilla, and vomer. The prefrontals loosely articulate with the anterolateral surfaces of the frontals, forming the anterior edge of each orbit. A rather large lacrimal foramen or duct is located on the ventral border of each prefrontal near its articulation with the dorsal surface of the maxilla. Figs. 8-A, 9-A, 11-C Posteriorly the frontals articulate with the fused (coalesced) parietals, there being some degree of motility between the frontals and the parietal. The parietal is the largest of the cranial elements and forms the greater part of the braincase roof, but like the frontals, extends far down either side of the brain, reaching ventrally to the basisphenoid and forming the posterior portion of each orbit. The posterolateral borders of the parietal suture with the prootics. The tri- angular-shaped, dorsal surface has its narrow apex attached to the supraoccipital posteriorly. Dorsally a V-shaped pair of crests converge pos- teriorly near the supraoccipital articulation. The posterodorsal surface of each orbit is expanded laterally by the parietal into a flattened lobe- like process, the lateral border of which articu- lates with the curved mesial surface of the post- orbital. A smaller coneshaped process, ventral to the lobelike process, articulates dorsally with the posteroventral border of the postorbital. About midway down the coalesced sagittal sut- ure, there are two small parietal foramina im- mediately lateral to the midline. POSTORBITALS Figs. 8-A, 9-A, 11-C The small postorbitals, considered to be post- frontals by Owen (1866) and Kellicott (1898), are narrow, elongate flattened bones which ar- ticulate with the anterolateral surfaces of the parietal, and form the dorsoposterior boundary of each orbit. A strong ligament connects the posterior part of each postorbital with the an- terodorsal surface of the ectopterygoids. SUPRAOCCIPITAL Figs. 8-A, 9-A, 11-C The small butterfly-shaped supraoccipital is medially placed and unites anteriorly with the parietal, laterally with the prootics, and pos- teriorly with the exoccipitals. It forms the roof of the posterior part of the brain cavity, and is internally ex-panded from its lateral wings to form the dorsal portion of each oHc capsule. The lambdoidal ridge or nuchal crest is prominent, and extends obliquely posterolaterally down each side of the posterior portion of the cranium. Each lateral branch of the crest diminishes in size near the exoccipital articulation, b t con- tinues as a rather sharp ridge along the anterior border of the exoccipital, terminating at the posterior border of the fenestra ovalis. Dorsally a short median longitudinal ridge extends pos- teriorly, ending where the mid-dorsal suture joins the exoccipitals. Figs. 8-A, 9-A, 11-C Each prootic is an irregular quadrate-shaped bone forming the anterior part of the internal otic capsule and the jx)sterolateral wall of the braincase. It is bordered by the parietal an- teriorly, supraoccipital dorsally, exoccipital pos- teriorly, and the basisphenoid and basioccipital ventrally. The anterior half of each fenestra ovalis is formed by the prootic; the exoccipital completes the fenestra posteriorly. The foramen is basically for the passage of the trigeminus, but Brigham Young University Science Bulletin BO.HYP. Fig. 10. Skull, ventral. A, Pituophis catenifer, 4X. B, Tliamnophis radix, 5X. Osteology of Snakes 19 also carries some facial nerve fibers. It is divided externally into two openings, both anterior to the fenestra ovalLs. The posterior foramen is larger than the anterior one, and has a lateral proct*ss projecting posteriorly from its anterior border, therefore partially obscuring the open- ing. Other smaller foramina pierce each prootic, carrying cranial nerve fibers. EXOCCIPITALS Fig.s. 8-A, 9-A, 10-A, U-C The exoccipital bones form the posterolateral walls of the braincase as well as part of its roof. They are joined together by a mid-dorsal suture, connected to the supraoccipital and pro- otics anteriorly, and resting upon the basioc- cipital ventrally. The opisthotics are fused with the exoccipitals, the combined bone thus sur- rounding the jugular foramen and extending forward to form the [X)sterior border of the fenestra ovalis (Romer, 19.56). This jugular foramen, which is internally subdivided into two or more smaller aspects, is just posterior to the fenestra ovalis, and is used for the passage of the ninth and tenth cranial nerves. Posteriorly the exoccipitals form the entire dorsoventrally- compressed, oval foramen magnum, except for a small ventral portion of the occipital condyle. Posteroventrally the exoccipitals terminate as a pair of articulating tubercles which form the lateral portions of the crescent-shaped occiptal condyle. BASIOCCIPITAL Figs. 9-A, 10-A, 11-C The basioccipital is a pentagonal-shaped bone forming the floor of the posterior part of the brain ca\'itA'. The ventral surface has a low, sagittal ridge with a laterally depressed area on either side. Posteriorly a midventral tubercle forms the ventral portion of the occipital con- dyle, thus completing the foramen magnum. The basioccipital is bordered laterally bv the exoc- cipitals, and is joined anteroventraliy with the basisphenoid. BASISPHENOID Figs. 9-A, 10-A In the skull of adult serpents, the basisphen- oid is applied without a suture anteriorly to the elongate parasphenoid, forming a single bone. The basisphenoidal portion of the bone is a flat, hexagonal plate, bounded dorsally at its margins by prootic and parietal walls, and pos- teriorly by the basioccipital. A pair of prominent transverse processes extend obliquely and ante- riorly toward the midline from its ventrolateral surfaces. Paired carotid foramina pierce the bone laterally to each process to enter the pituitary fossa. This fossa is seen ventrally as a convex area, and is the floor of the sella turcica in which the pituitary gland lies. A pair of foramina also pierce the posterolateral borders of this bone. The narrow elongate parasphenoidal portion of the basisphenoid extends anteriorly into the rostrum area, and becomes the floor of the orbit and optic foramen. A median longitudinal groove extends the entire length of the ventral surface, whereas a dorsomesial keel is wedged between the ventral connection of the f rentals. MAXILLAE Figs. 8-A, 9-A, 10-A Each maxilla is a short, curved bar that con- forms to the shape of the head. On the mesial border, about midway, there is a small horizon- tal dorsoposteriorly-pointing process that articu- lates with the ventral surface of the prefrontal. The posterior end of the maxilla is broadened somewhat, and is received by the flattened club- shaped ventral surface of the ectopterygoid. Each maxilla bears sockets for about fifteen or sixteen curved teeth, but not all of the sockets contain rigidly fixed teeth; therefore, some are usually lost in the preparation of the skull. In P. catenifer about every other tooth on the maxillae, as well as the other tooth-bearing structures , was only loosely lodged in the grooves and could be removed very easily. The maxillae are connected by fibrous tissue to the premaxillae, and not by an osseous articulation as in most vertebrates. ECTOPTEBYGOroS Figs. 8-A, 9-A, 10-A The ectopterygoids, which were sometimes referred to as the transpalatines by earlier work- ers such as Kellicott (1898) and Wiedersheim and Parker (1907), are small club-shaped, flat bones connecting the maxillae to the pterygoid, and overlying each at its ends. Each flattened spatulatelike anterior process is notched ante- riorly where it ligamentously articulates with the maxilla, and an elongate, posterior end articulates with the dorsolateral groove in the pterygoid which runs parallel with the curved contour of tliis bone for its entire length. The ectoptery- goids do not bear teeth. PALATINES Figs. 9-A, 10-A Tlie palatines are short bars placed anteriorly to the pterygoids, and mesially to the ma.xillae, Brigham Young University Science Bulletin SOC. FO.M. OT.C. BR.COR. BO. HYP. Fig. 11. Pituophis catenifer, A-C: A, hyoid apparatus, ventral, 4X; B. mandible, medial, 5X; C. cranium, po.sterior. 5X. D, Thamtiophis radix, cranium, posterior, 9X. Osteology of Snakes each bearing nine or ten teeth. The posterior end of each pahitine articulates with the ante- rior end of the pterygoid. About midway each palatine possesses a broad, flattened, horizontal, median process with apex directed anteriorly, and a similar but narrower lateral process ar- ticulating with the ventral border of the pre- frontal. The palatines are connected by fibrous connective tissue to the snout elements. PTERYGOIDS Figs. 8-A, 9-A, 10-A Each pter)'goid is a curved, flattened bar extending from the palatine posterolaterally to the angle of the mandible. Anteriorly it narrows and forms a loose articulation with the posterior end of the palatine. From this articulation a row of eight to twelve teeth extends posteriorly along the medial border, ex'tending about mid- way. The pterygoid receives the ectopterygoid into an elongated, dorsolateral groove. Posterior- ly the pterygoid narrows into a slender, curved process that re.iches the base of the c|uadrate as well as the mandible. SUPRATEMPORALS Figs. 8-A, 9-A, lO-A The supra temporals are not true squamosals as some of the earlier writers indicated (Romer, 1956). Owen (1866) referred to them as mas- toids, but this terminology has been dropped in favor of the more applicable term (supra- temporal) used in this paper. The problem of the homologies of the scjuamosal, supratemporal, etc., is, however, far from being solved in the various reptile groups, as pointed out by Jollie (1960). Each supratemporal is a short, harrow, flat- tened bone connecting the posterolateral dorsal part of the skull with the proximal end of the quadrate. Anteriorly each supratemporal over- lays a portion of the parietal, prootic, supra- occipital, and exoccipital, and is attached to these cranial elements by fibrous connective tissue. The two bones are almost parallel to each other, converging only sbghtly as they extend posteriorly. QUADRATES Figs. 8-A, 9-A, 10-A Each quadrate is a strong, rectangular bone; its proximal surface articulates with the postero- lateral border of the supratemporal, and its dis- tal, notched, condylar surface with that of the mandibular condyle. Its proximal surface is twisted to become obliquely aligned along the sagittal plane, whereas its concave distal articu- lating surface is extended transversely to form an intercondyloid fossa for mandibular articu- lation. About midway down the mesial side of each quadrate there is a small, rectangular, raised process which articulates with the distal end of the stapes. MANDIBLE Figs. 8-A, 9-A, 10-A, 11-B The mandible is composed of two jaw bones which lack a firm anterior symphysis, but are connected by an elastic ligament anteriorly, and is held together posteriorly to the symphysis by the transverse muscles. Each jaw bone is long, curved anteromesially, and is separable into two major bones in the adult snake; how- ever, the splenial and angular bones can some- times be separated in addition to these two major elements. The longer pro.ximal part of each jaw is composed of angular, prearticular, articular, surangular, and splenial aspects, and is without teeth. Although the splenial and an- gular are the only bones that can be separated from this proximal portion of the jaw, the other bones are coalesced and form the following areas: the articular and prearticular form the mandibular condyle area; the surangular ex- pands into a longitudinal dorsomesial crest near the condyle, a deep lateral groove is formed at the base of the crest, and the inferior dental foramen opens at the bottom of the groove. The splenial and angular bones are triangular shaped with their bases together and their apices point- ing anteriorly and posteriorly from their mid- ventral location on the mesial surface of the jaw. The apex of the splenial points anteriorly; and owing to the reduction in its size, the anterior part of the meckelian canal becomes an open mesial groove in the dentary just anterior to the splenial. Bogert ( 1943 ) uses the term postarticular when referring to the process of the articular that extends posteriorly from the mandibular articulation with the quadrate. His work was with the cobra and other elapids. The dentary forms the anterolateral portion of the jaw, and contains a row of sixteen to eighteen of the usual hooked or recurved teeth. It is short, curved, pointed anteriorly, and bifur- cated posteriorly on the lateral surface, thus articulating with the anterior projection of the proximal portion of the jaw. This contact be- tween the two bones is not a close one; there- fore, it gives a degree of flexibility to the jaw. The mental foramen is located near the middle of the lateral surface of the dentary. Bbigham Young University Science Bulletin STAPES Figs. 8-A, 9-A Each stapes (columella) is a delicate, slen- der, rodlike stylus connecting the otic capsule, by way of the fenestra ovalis, to the rectangular articular surface of the quadrate. Its pro.ximal end is enlarged to forni a footplate that fits into the foramen. The stapes is believed to transmit sound vibrations from the various jaw bones to the otic capsule. HYOID APP.\RATUS Fig. 11-A The hyoid apparatus is a cartilagenoiis, V- shaped structure with its vertex or base (basi- hyobranchial ) attached to the midventral raphe between the lower jaws; each rodlike branchial cornu e-vtends posteriorly along the lateral bor- der of the elongate tongue, being inserted into the base of one of the elongate retractor muscles. Suj^erficial, ventral cranial muscles are attached to the basihyobranchial and the anterior portion of each comu. The hyoid apparatus is not at- tached to any of the skull elements as it is in most other vertebrates, but lies imbedded in the muscle and fascia of the intermandibular region. The tongue, along with its posteriorly con- tinuing retractor muscles, is more than twice as long as the skull. The hyoid apparatus, there- fore, is modified into a very long structure in this species even though it is a rather delicate organ. OSTEOLOGICAL COMPARISONS The two species considered in this study (Pituophis cafenifer and Thamnophis radix) show many osteological similarities. There are, however, a few significant structural differences existing between them. Some of these differ- ences appear to be highly correlated with the special adaptations made by the two species in becoming more compatible with their specific environments and food habits; however, there are many minor structural differences that would seem to be nonadaptive, or perhaps they ma\' have other effects of survival value not apparent to the authors. Only those differences which appear to be of sufficient magnitude to be com- pared, and perhaps have some bearing on the adaptation of these two species, will be dis- cussed. A complete set of osteological illustrations for Thamnophis radix, corresponding to those of Pituophis catenifer, has been included and will be referred to in the course of the compara- tive discussion. VERTEBRAL COLUMN Precaudal Vertebrae The atlas of T. radix (Fig. 1-E, F, G, and H) is proportionately broader along the sagittal, neural suture line ( Fig. 1-G and H ) than in P. catenifer (Fig.l-C and D), but its dorso- posterior projections containing the postzygapo- physes are not as prominent. The hypapophysLs of the atlas of T. radix (Fig. 1-H) does not point posteroventrally as it does in P. catenifer ( Fig. 1-D ) , but instead is represented as a small vertical process extending ventrad from the ventral portion of the intercentrum. The major differences in the axis of the two species are found in the anterior and posterior hypapophyses. In T. radix the axis (Fig. 2-E, F, G and H) develops a stout, hatchet-shaped, anterior hypapophysis (Fig. 2-H) with its an- terior articular facet placed vertically on its anterior border, whereas the previously-men- tioned, wedge-shaped, anterior hypapophysis of P. catenifer (Fig. 2-D) possesses a narrower posteroventrally situated facet. The posterior hyjiapophysis of T. radix (Fig. 2-H) is not as spineUke or tapered as in the other species, but is more trapezoid in shape with its ventral margin being somewhat truncate. Some of the less obvious differences existing in these vertebrae are the pro[X>rtionally narrow- er odontoid process (Fig. 2-G), the more pos- teriorly placed, dorsoventral odontoid suture (Fig. 2-G and H), and the greater anteriorly projecting portion of the neural spine (Fig. 2-H) in T. radix. In T. radix the first thoracic vertebra (Fig. 3-E and F) lacks free ribs, just as it does in P. catenifer. The fixed ribs or transverse processes, however, are short, and extend posteriorly in both species. In the majority of the specimens studied, the transverse processes of the first thoracic vertebrae were a little thicker at the base, and less spinelike in T. radix (Fig. 3-F). This species also differs from P. catenifer in that the neural spine is narrower and slightly longer (Fig. 3-F), the anterior zygapophyses are placed Osteology of Snakes 23 more mesially and are partially covered by the zygosphenes when viewed dorsally ( Fig. 3-E ) , and there is a complete lack of metapophyses. The other thoracic or typical t'loracic ver- tebrae show few noticeable differences between the two species. It can be observed, however, that in T. radix these twenty-two or twenty- three thoracic vertebrae ( Figs. '3-G and H; 4-E and F) develop proportionally smaller metapo- phvses but larger transverse proc-esses (Fig. 3-G and H). The dorsal convex portion of the trans- verse prcK-ess, bilobed rib, articular surface pro- jects more laterally (Fig. 4-E and F), and the facets of the anterior zygapophyses are tilted slightly more oblicjuely dorsad (Fig. 3-G) in T. radix than those of P. catenifer (Fig. 3-C). The lumbar vertebrae of T. radix (Fig. 4-G and H) retain relatively large hypapophyses (Fig. 4-H), whereas the hypapophyses of P. catenifer (Fig. 4-D) are lacking in this region, and are represented only by rudimentary mid- ventral ridges. The presence of hypapophyses on the lumbar vertebrae of T. radix makes it very difficult to distinguish the first few vertebrae of this subregion from the last two or three thor- acics. However, there is a decisive reduction in the length of the hypapophyses extending over three or four vertebrae, making it jx>ssible to distinguish between the two vertebral areas with a relative degree of consistency. In T. radix there is an average of approximately 136 or 137 lumbar vertebrae, compared to 185 or 186 found in P. catenifer. The lumbar vertebrae of T. radix (Fig. 4-G and H) develop transverse processes much like those of the thoracic vertebrae. The dorsal rib facets are placed more dorsally (Fig. 4-H), and are more prominent laterally (Fig. 4-G) than in P. catenifer. In tliis species the ventral apex of each transverse process (para- pophysis ) is devoid of a rib articular surface, and thus forms an anteroventrally projecting process similar to those found on the thoracic vertebrae of botli species (Fig. 4-B and F). Caudal Vertebrae The anterior caudal vertebrae of T. radix (Fig. 5-E, F, G and H) number five or six in the specimens studied. In addition to their total number being essentially the same as in P. catenifer, they also develop a ridgehke hypapo- physis (Fig. 5-E and F) on the first one or two vertebrae of this subregion, and rather rudi- mentary haemapophyses on the remaining three or four. The dorsal branch of the lymphapo- physis does not arc as much ventrally in T. radix (Fig. 5-E and F), and the ventral branch does not attain as great a length proportionally, sweep back posteriorly, or curve as much ventrally at its apex. The neural spine is also narrower and projects more posteriorly (Fig. 5-H) in T. radix. There are three or four vertebrae belonging to the middle caudal subregion in T. radix ( Fig. 6-E, F, G and H), the same number as was found to be present in P. catenifer. These verte- brae are similar in the two species, but there are noticeable differences in their fixed ribs (trans- verse processes ) and haemapophyses. In T. radix the fixed ribs curve posteroventrally (Fig. 6-G) instead of anteroventrally ( Fig. 6-C ) as they do in P. catenifer. The haemapophyses of this species are more delicate in structure, not as widely arched from each other at their midpoint (Fig. 6-E and F), and they project more pos- teriorly (Fig. 6-G and H) than in P. catenifer. The vertebrae of the posterior cauda sub- region of T. radix (Fig. 7-E, F and G) vary greatly in number from sjxjcimen to specimen just as in P. catenifer; however, an average would fall somewhere between fifty and sixty (an average of about five less than in P. cateni- fer). For the most part, the same interspecific differences found in the middle caudal vertebrae are carried over into the posterior subregion; however, the neural spine slopes more obliquely anteriorly ( Fig. 7-G ) , and the transverse pro- cesses curve more anteriorly at their apices and are more pointed and less foothke (Fig. 7-G) in T. radix. There were no noticeable differences in the ribs of the two species except for size. Tham- nophis radix, being the smaller serpent, de- veloped proportionately smaller ribs. The skulls of both species are composed of similar bones and bone structures; however, there are a few morphological and size relation- ship differences that do exist. The most apparent of these differences and those thought to be of some significance will be discussed. A dorsal view of the skull of T. radix (Fig. 8-B), a lateral view (Fig. 9-B), a ventral view (Fig. 10- B), and a posterior view of the cranium (Fig. 11-D) have been included to facihtate the com- parative discussion. The premaxilla of T. radix is more com- pressed dorsoventrally, having a distinct depres- sion or fossa about midway along its dorsal surface (Fig. 9-B) in contrast to the ridgelike process in P. catenifer ( Fig. 9-A ) . This postero- dorsal process does not extend dorsally to wedge 24 Brigham Young University Science Bulletin between the anterior portions of the nasals as it does in P. catcnifer. The septomaxillae are heavier and more ex- tensive in T. radix. They cover a greater pro- portion of the ventrallv-attached vomers in this species, and the lateral winglike processes are proportionately broader and turn posterodorsal- ly (Fig. 9-B) instead of anterodorsally (Fig. 9- A) as they do in P. catcnifer. The elongate, flat anterior processes from the septoma.xillae are less tapered and more horizontally situated in T. radix. Although tlie frontals and prefrontals are more rigidly attached in T. radix, there are no significant differences in these bones in a com- parison of the two species. The parietal and postorbitals, however, show a major structural modification in T. radix. The parietal is narrow- er proportionately, develops more prominent dorsolateral crests, and lacks the small, paired foramina on its dorsal surface (Fig. 8-B). In P. catcnifer the parietal foramina (Fig. 8-A) are quite prominent. In comparing the length of each parietal with its width, it was found that in T. radix the parietals were 1.1 times as wide as they were long, whereas in P. catcnifer they were 1.3 times as broad as long. The postorbitals of T. radix are not just narrow, elongate bones attached to the antero- lateral margins of the parietals as they are in P. catcnifer (Figs. 9-A, 11-C). b'lt thev ex- tend ventrally, coming in contact with the dorsal surfaces of the anterior spatulate processes of the ectopterygoids (Figs. 9-B, 11-D). Some of the less obvious differences of the bones comprising the braincase are the narrow- er basisphenoid (Fig. 10-B) in the case of T. radix, with its transverse processes more pos- teriorly situated and extending laterally (perpen- dicular to the midline) instead of obliquely and posterolaterally as in P. catcnifer (Fig. 10-A). The posteroventral surface of this bone presents a convexity instead of a concavity as it does in P. catcnifer. In T. radix the prootics have a prominent crest running longitudinally along their lateral borders just dorsal to the large prootic foramina. This crest is somewhat hidden by the lateral border of the supratemporal. The lateral process which projects posteriorly, par- tially obscuring the opening to the posterior prootic foramen, is not present in T. radix. Therefore, the foramen appears round, from a lateral view (Fig. 9-B), instead of kidney- shaped (Fig. 9-A) as in P. catcnifer. The basioccipital of T. radix develops a prominent, median, crestlike process or hypapo- physis (Fig. 11-D) for the attacliment of ventral neck muscles; this structure is not found on the basioccipital of P. catcnifer. From a dorsal view (Fig. 8) the size of the po,steromesial notch in the dorsal surface of each exoccipital, where each sutures in the midline, is greater in the case of T. radix (Fig. 8-B). Both the quadrate and the supratemporal are proportionately heavier and broader (Figs. S-B, 9-B) in T. radix. Tliev do, however, occupy approximately the same position in both species. The maxillae, ectopterygoids and palatines (Fig. 10-A and B) are similar structurally in the two species, but there is a difference in the number of teeth on the tooth-bearing structures ( maxillae and palatines ) . The maxillae of T. radix bear about twenty-two or twenty-three curved teeth as compared to the fifteen or six- teen in P. catcnifer; the palatines bear about eighteen teeth as compared to the nine or ten teeth developed in P. catcnifer. The large, flat- tened, median process of the palatine is directed more dorsally in T. radix, and in most cases the dorsal apex of this process arches mesially, ter- minating in a ventral direction. The pterygoids (Fig. 10-A and B) are pro- portionately broader, their longitudinal ventral grooves deeper, and the number of teeth is greater in T. radix. This species bears about twenty-seven or twenty-eight teeth (more than twice as many as found in P. catcnifer) in a longitudinal median row which extends almost twice as far posteriorly as it does in P. catenifer. The pterygoid is abruptly curved laterally at its posterior end, making the mandibular articulation in T. radix; whereas in P. catcnifer the entire bone is gradually curved laterally until it articu- lates posteriorly with the angle of the jaw. The mandible of T. radix is curved a Httle more mesially at its middle dentary articulation (Fig. 10-B), and it contains from twenty-eight to tliLrtv teeth on the dentary portion, or about twice as many as previously recorded for P. catenifer. Both species have splenial and angu- lar bones that can be separated from the an- teromesial border of the proximal half of the jaw bone (Fig. 11-B). Although the shape and structure of the hyoid apparatus (Fig. 11-A) is very similar in the two species, it is proportionately longer in T. radix, being appro.ximately 2.2 times longer than its mandible. In P. catenifer the hyoid is only approximately 1.4 times longer than the Osteology of Snakes 25 mandible. Both specit's, however, develop a rather large and functional hvoid apparatus. According to Albright and Nel.son (1959), Elaphe ohsoleta has a hyoid apparatus about twice the length of its skull. Smith and Wanier (194S) figured a series of ophidian hyobranchia and included the genus Pituophi.s, but not Thamnuphis. Their Fig. 1-R of Pituophii- is similar to our Fig. 11-A, but dif- fers in that it is smoothly rounded anteriorly, whereas we find Pituophis to exhibit a small anterior protuberance which we have desig- nated as the basihyobranchial. The specimens of T. radix used in this study, however, did not possess this anterior protuberance. Although Cowan and Hick (1951) did not discuss the osteology of the hyoid apparatus of Thamnophis siitalis as such, they indicated its structure by figures and descriptions concern- ing the hyoid musculature. According to these authors the basihyal (basihyobranchial) por- tion of the hyoid of T. sirtalis is smoothly rounded anteriorly, and does not possess an an- terior protuberance. DISCUSSION From the foregoing comparisons it is evident that there are osteological differences existing in the two species. In order to provide a com- plete comparison of the differences it would be necessary to correlate and compare as many of these interspecific differences as possible with other serpent groups (particularly colubrids) in order to arrive at some osteological continuity in the species studied, and to postulate some reasons for these differences. This is obviously beyond the scope of this study. There are, how- ever, a few osteological differences that can be discussed, and these will be considered insofar as our data will permit. The ribless condition of the first thoracic vertebrae, as demonstrated by P. catenifer and T. radix, has not been reported in the species studied by the majority of workers ( op. cit. ) . Owen (1866) stated that the ribs commence in the cobra, as they do in other serpents, at the third vertebra from the head. Kellicott (1898) mentioned short transverse processes in connec- tion with the heads of the ribs as being present on all of the body vertebrae ( not including atlas and axis ) in Heterodon. A general statement was made by Gadow (1901) concerning snakes, in which he states that "all the vertebrae, except the atlas, carry ribs." Albright and Nelson (1959) reported short ribs on vertebrae 3 and 4 of Elaphe, the first full-length rib being on the fifth vertebra. Ludicke (1962), on the other hand, reported that all of the vertebrae, with the exception of the atlas, can bear free ribs, but the axis and the first two "cervical" or thoracic vertebrae may also lack ribs in some species. The latter statement appears to coincide more fully with the findings of our study in which the first thoracic vertebra was lacking free ribs. Because of the close adherence of the atlas to the cranial elements and the difficulty in- volved in separating it from the skull intact, it may be that some of the above statements are not based on sufficiently meticulous dissection to be complete. Possibly the true atlas has been overlooked in some cases, and has not been in- cluded as part of the anterior vertebrae. A de- tailed study of many different genera would be necessary in order to determine the extent of the ribless first thoracic vertebrae in the various types of colubrid snakes. The presence of metapophyses on the first thoracic vertebra of P. catenifer, along with their greater development on the other thoracic ver- tebrae, seems to indicate a more extensive area for the attachment of lateral trunk muscles, giv- ing this constriction species an advantage in both strength and motility in the neck and anterior trunk region. Tltatnnophis radix, being the smaller of the two species, has fewer precaudal vertebrae, but the proportionate number of the various types of vertebrae in this region vary in the two species (Table 1). There is a decrease of ap- proximately 48 (26%) lumbar and 28 (29%) thoracic vertebrae in T. radix when compared to P. catenifer. It is evident from the above data that the larger snake (P. catenifer) owes much of its greater precaudal body length to the great- er number of lumbar vertebrae, but proportion- ately or percentage-wise, there is an addition of about twice as many thoracic vertebrae as com- pared to the lumbar. Owen (1866) stated that hypapophyses are developed on the first seventy-four of 253 body vertebrae in Python tigris, the first sixty anterior l)ody vertebrae (305 total, counting caudals) in Brigham Young University Science Bulletin Table 1. A comparison of the number of vertebrae in each of the vertebral regions and subregions of fourteen Pituophis catenifer and twelve Thamnophis radix. Both male and female specimens are included. Average Number of Vertebrae Vertebral (Ran^ ;es in Parentheses ) Colunm Areas P. catenifer T. radix Atlas 1 1 Axis 1 1 First Thoracic 1 1 Other Thoracic 52 23 Lumbar 185 137 Anterior Caudal 6 6 Middle Caudal 4 4 Posterior Caudal 61 56 Precaudal 240 (237-244) 163 (160-165) Caudal 69 (60-78) 64 (58-69) Precaudal and Caudal 309 (297-322) 227 (218-234) Boa constrictor, and from the thirty-two anterior vertebrae of Dasypeltis scahra, which has a total of 256 vertebrae. The anterior ten hypapophyses of thLs latter snake are directed posteriorly, whereas the posterior ten are pointed anteriorly. Owen also reported that all of the 168 body ver- tebrae of Crotalus horridus develop hypapo- physes, and Ukewise, all of the body vertebrae in Naja tripudians. Tliese hypapophyses, however, were found to be shorter in the more posteriorly located body vertebrae. Heterodon, however, possesses hypapophyses only on the first twenty- two of 135 body vertebrae (Kellicott, 1898). From the few species mentioned above, it would appear that when using the presence or absence of hypapophyses as a criterion, P. caten- ifer is similar to the other constricting species such as P. tigris and B. constrictor. Heterodon and D. scaber, not being constrictors and yet lacking hypapophyses on the lumbar vertebrae, appear to contradict the highly speculative as- sumption above, or possibly one could infer that these snakes are closely related to the constric- tor-type serpents. A much more intensive inves- tigation including many different species of Colubridae would have to be made before such a hypothesis could be verified. Although P. catenifer has an average of about five more caudal vertebrae than T. radix (Table 1 ) , its tail is actually proportionately shorter because of the greater number of precaudal vertebrae. The ratio of precaudal vertebrae in P. catenifer is 3.5:1, that of T. radix being 2.5:1. Ludicke (1962) states that Colubridae and Elapidae have a ratio of precaudal to caudal vertebrae of 2:1. The t\vo colubrid species in- volved in this study appear to have a higher proportion of body or precaudal vertebrae than the group of colubrids from which Ludicke's ratio was calculated. The added length in the tail region (proportionately) in T. radix over P. catenifer might be of some advantage to this semi-aquatic species when taking to the water. According to Brongersma ( 1938 ) the char- aoteristic of well-developed hypapophyses under the posterior precaudal vertebrae is a variable charac^:er in some species. Such species as Chrijsopelea ornata (Shaw) and Boiga irregu- laris (Merr. ), collected from the Indo-Australian Archipelago, the Philippines and Siam, were found to vary from specimen to specimen in the presence or absence of well-developed hypapo- physes. The presence of parietal foramina, in the case of P. catenifer, is a pecuhar phenomenon if the statements made by some writers are to be interpreted rigidly. Kingsley (1917) and Willis- ton (1925) stated that Ophidia lack parietal foramina. Gadow ( 1901 ) quahfied his statement somewhat, and mentioned that there is no inter- parietal or pineal foramen in serpents. In dealing with Elaphe, Albright and Nelson (1959) found a small foramen in the posteroinferior surface of the parietal, and stated that it is the only one of any size in the entire wide expanse of the bone. We assume that their posteroinferior sur- face of the parietal is our posterior lateral or ventral surface. In none of their plates did they figure this foramen. According to WiUiston (1925) the parietals are always fused in ser- pents. Romer (1956) stated that parietals, as in most lizards, are fused into a single bone in nearly all snakes. The two species involved in this study demonstrate this fused parietal phenomenon; however, Mookerjee and Das ( 1932 ) recorded that distinct paired parietals were present in Tijphlops braminus, and that perhaps this species had been overlooked by some previous writers. The osseous connection between a cranial bone ( postorbital ) and the upper jaw structures, as found in T. radix, has less flexibility and is more rigid than the ligamentous connection in P. catenifer. Perhaps this structural rigidity is needed in the case of T. radix as it eats live, nonconstricted prey which would be struggling during the engulfing process. It should be men- tioned, however, that Python, a constrictor, has a postorbital which contacts the ectopterygoid. The postorbitals of Boa and some poisonous ser- pents, according to Owen (1854), do not reach the ectopterygoids. Albright and Nelson (1959) indicate that in Elaphe obsoleta the small, nar- Osteology of Snakes 27 row postorbitals approach but do not articulate with the upper jaw structures. Such an osseous connection is also lacking in Heterodon. Tlie proportionately heavier and broader (juadrate and supratemporal in the case of T. radix seems to c-oincide with the previously stated conjecture concerning rigidity in the jaw structures necessary for the feeding habit of this species. According to Kellicott (1898) the basiocci- pital hv'papophysis, which is present in T. radix but missing in P. catenifer, is also lacking in Heterodon. Owen (18.54) stated that the python has a prominent hypapophvsis with a recurved point, and that the rattlesnake also develops a long, strong, recurved hypapophvsis from its basioccipital which acts as a point of insertion for the powerful ventral neck muscles, by which the downward stroke of the head is performed in the act of inflicting a wound. In both species studied, there were occasional specimens which demonstrated an abnormally large number of teeth on some of their toothed structures. It was noted that the bases of the extra teeth were lined up along the sides of the teeth in the normal tooth row, one tooth being firmly fi.\ed in the socket, and the more medially placed tooth being only connected to the bone by fascia. It appeared as if the loosely connected teeth were being replaced by the firmly fixed ones, thus maintaining a complete set of func- tional teeth at all times. This observation, per- taining to the continual replacement of teeth in serpents, was noted in the works of West (1898), Kingsley (1917), Wiedersheim and Par- ker ( 1907 ) , Gadow ( 1901 ) , and more recently by Edmund (1960), to mention a few. Almost all reptiles are polyphyodont ( number of denti- tions indefinite), but in some cases such as Typhlopidae, certain teeth are not replaced and others undergo reduction (Wiedersheim and Parker, 1907). The greater number of teeth found on all of the tooth-bearing structures in T. radix ( Table 2) seems to give this species an advantage in grasping and retaining struggling prey such as frogs, toads, fish, grasshoppers, mammals, and other regularly eaten animals. Tliis adaptation to the nonconstrictor habit of food-getting was noted previously in connection with the post- orbitals, quadrates, and supratemporals. The proportionately narrower posterior por- tion of the skull, due mainly to the narrower parietal, has possibly resulted from selection for more streamlined forms, in the case of T. radix, which would have an advantage for life in the swampy areas where diving and swimming be- come an integral part of its life. Table 2. A comparison of the number of teeth de- veloped on die various tooth-bearing structures of four- teen Pituophis catenifer and twelve Thamnophis radix. Tooth-bearing Structures Average Number of Teeth ( Ranges in Parentheses ) P. catenifer T. radix Maxilla 15 (14-17) 23 (22-25) Palatine 9 (8-11) 18 (16-19) Pterygoid 10 (8-12) 28 (26-29) Mandible 18 (15-20) 28 (26-30) SUMMARY AND CONCLUSIONS The complete skeletal anatomy of Pituophis catenifer deserticola Stejneger is described in detail. The osteology of this species is com- pared with that of Thamnophis radix haijdeni Kennicott in the following areas: atlas, axis, first thoracic, other thoracic lumbar, anterior caudal, middle caudal, and posterior caudal vertebrae, as well as the skull and hyoid apparatus. Many structural similarities exist in these two species of colubrids. Tlie more important struc- tures or aspects in which similarities were ob- served are: (1) the natural division of the verte- bral column into regions and subregions, (2) the gross morphology of the various types of vertebrae, (3) the number and ty|:>es of bones comprising the skull, (4) the number and loca- tion of the majority of vertebral and skull fora- mina, (5) the structure of the hyoid apparatus and the ribs, and (6) the position and shape of the stapes. The major structures or aspec-ts in which there are differences encountered between the two species are: (1) the shape of the anterior hypapophysis of the atlas, (2) the e.vistence or lack of metapophyses on the first thoracic verte- bra, (3) the presence or absence of hypapo- physes on the lumbar vertebrae, (4) the rela- tive size and shape of the lymphapophyses, (5) the structure of the caudal haemapophyses, (6) the number of vertebrae in each region and sub- region, (7) the relative size and shape of the 28 Bricham Young University Science Bulletin cranium and its component bones, (8) the shape and character of the septomaxillary process, (9) the existence or lack of paired parietal foramina, (10) the shape and extent of the postorbitals, (11) the presence or absence of basioccipital hypapophyses, (12) the number of maxillary, pterygoid, and mandibular teeth, ( 13 ) the rela- tive size and character of the supratemporal, quadrate, and mandible, and ( 14 ) the relative size of the hyoid apparatus. LITERATURE CITED Ahrenfeldt, R. H. 1955. Two Briti.sh anatomical studies on American reptiles (1650-1750) II. Ed- ward Tyson: Comparative anatomy of the timber rattle,snaike. Herpetologica, 2(l):49-69. Albright, R. C, and E. M. Nelson. 1959. Craniiil kinetics of the generalized colubrid snake Elaphii obsoleta qtiadrivittata. I. Descriptive morphology, and II. Functional morphology. J. Morph., 105(2): 193-492. Bogert, C. M. 194.3. Dentitional phenomena in cobras and other elapids with notes on adaptive modifica- tions. Bull. Amer. Mus. Nat. Hist., 8 1 ( 3 ) : 285-360. Brongersma, L. D. 1938. On the presence or absence of hypapophyses under the posterior precaudal ver- tebrae in some snakes. Zool. Meded. Leiden, 20: 240-242. Cole, F. J. 1944. A history of comparative anatomy. Macmillan, London. Cowan, I., and W. Hick. 1951. A comparative study of the myology of the head region in three species of Thamnophis. Trans. Roy. Soc. Canada, ser. 3, 45(3). Edgeworth, F. H. 1935. The cranial muscles of ver- tebrates. Univ. Cambridge Press, Cambridge. Edmund, A. G. 1962. Se{juence and rate of tooth replacement in the Crocodilia. Life Sciences Divi- sion— Contribution 56, Toronto. Gadow, H. 1901. Amphibians and reptiles. Macmil- lan, London. Gegenbaur, C. 1878. Elements of comparative anat- omy. Macmillan, London. Goodrich, E. S. 1930. Studies on the structure and development of vertebrates. Macmillan, London. Gregory, W. K. 1913. Homology of the "lacrimal" and the "alisphenoid" in the recent and fossil rep- tiles. Bull. Geol. Soc. Amer., 24:241-246. Haas, G. 1930. Ueber das kopfskelett und die haut- muskulatiu- der Typlilopiden und Glauconiiden. Zool. J;ihrb., Abt. Anat., 52:1-94, 95-218, 347-404. Hoffman, C. K. 1890. Reptilien. In Bronn's Kla.ssen und Ordmmgen des Tier Reich, 6(3) :1420-1465. Hu.\ley. T. H. 1871. A manual of anatomy of verte- brated animals. J. and A. Churchill, London. Jollie, M. T. 1960. The head skeleton of the hzard. Acta Zoologica, 41:1-64. Kellicott, D. S. 1898, The dissection of the Ophidi;in. Gen. Biol. Supply House, Chicago. Kingslcy, J. S. 1917. Outline of comparative anatomy of vertebrates. P. Blakiston, Philadelphia. Klauber, L. M. 1956. Ratdesnakes. Univ. California Press, Los Angeles, 2:710-1476. Ludicke, M. 1962. Ordnung der Khisse Reptiha, Serpentes. Handbuch der Zoologie, 7(5): 1-128. . 1964. Ordnung der Klasse Reptilia. Ser- pentes. Handbuch der Zoologie, 7(6) : 129-298. Mookerjee, H. K., and G. M. Das. 1932. Occurrence of a paired parietal bone in a snake. Nature, 130: 629. Owen, R. 1854. The principal forms of the skeleton and of the teeth. Blanchard and Lae, Philadelphia, , 1866, The anatomy of vertebrates, Long- man, Green and Co,, London, 1:53-57, Reynolds, S. H, 1913, The vertebrate .skeleton. Uni- versity Press, Cambridge. Robison, G. W., and W. W. Tanner, 1962. A com- parative study of the species of the genus Crota- phi/tus Holbrook ( Iguanidae ) , Brigham Young Uni- versity Sci, Bull,, Biol, Series, 2(1):1-31, Rochebrune, A, F, 1881, Memoire .sur les vertebres des ophidiens, J, Anat, Physiol. (Paris), 17:185-229. Romer, A. S. 1956. Osteology of the reptiles. Univ. Chicago Press, Ilhnois. Sedgwick. A. 1905. A student's text-book of zoology. Swan Sonnenschein and Co., Ltd., London, v, 2, Sood, M. S. 1941. The caudal vertebrae of Erijx jolmii (Russel). Proc. Indian Acad. Sci., 14:390- 394. . 1948. The anatomy of the vertebral column in serpents. Proc. Indian Acad. Sci., 28:1-26. Versluys, J. 1937. Kranium und visceralskeUett der Reptilien, In Bolk et al. Handbuch der verg- leichenden anatomic der WirbelHer, 4:780-7, West, G, S, 1898, Tooth succession, J. Linn, Soc, 26:517-526. Wiedersheim, R,, and W. N. Parker. 1907. Compara- tive anatomy of vertebrates. Macmillan, London. Williston, S, W. 1925. The osteology of the reptiles. Harvard Univ. Press, Cambridge. ABBREVIATIONS USED IN PLATES VEHTEBR.\E D.CR. dorsal crest. A.HYP. anterior hypapophysis. F.RB. fi.xed rib. A.INT.F. anterior intercentral facet. HAEM. haemapophysis. A.NEU.F. anterior neurapophysial facet. HYP. hypapophysis. CEN. centrum. HYP.F. hypapophysial facet. CO.PR. costal process. INT. intercentrum. Osteology of Snakes 29 LP.D.B. LP.V.B. MET. N.C.S. NEU. N.S. N.SU. O.CL. O.PR. PARP. P. HYP. P.INT.F. POZ. POZ.F. PRZ. PRZ.F. RB.F. RB.H. SH. T.PR. ZE. ZE.F. ZM. ZM.F. A. AR. BH. BO. BO.HYP. BO.T.PR. BR.COR. B.S. l)Tnp]iapophysis dorsal branch. lymphapophysis ventral branch. metapophysis. neiiro-central suture. neurapophysis. neural spine. neural suture. odontoid canal. odontoid process. parapopliysis. posterior hypapophysis. posterior intercentral facet. postzygapophysis. postzygapophysial facet. prezygapophysis. prezygapophysial facet. rib facet. rib head. shaft. transverse process. zygosphene. zygosphenal facet. zygantrum. zygantral facet. angular. articular. basihyobranchial. basioccipital. basioccipital hypapophysis. basioccipital transverse process. branchial comu. basisphenoid. D. D.F. EC. EO. FO.M. FOV. FR. J.F. M. ME.F. NAS. o.c. OP.F. OT.C. PAL. PAR. PARA. PAR.F. PER. PM. PO. PR. PR A. PT. QU. SA. SM. SM.PR. SOC. SP. ST. SUP. vo. V. dentary. dental foramen. ectopterygoid. exoccipital. foramen magnum. foramen (fenestra) ovalis. frontal. jugular foramen. ma.villa. mechelian foramen, canal or groove, nasal. occipital condyle, optic foramen, otic capsule, palatine, parietal, parasphenoid. parietal foramen, prefrontal, premaxilla. postorbital. prootic. prearticular. pterygoid, quadrate, surangular. septoma.xilla. septomaxillary process, supraoccipital. splenial. stapes. supratemporal. vomer, fifth cranial nerve. ^-ha^ Trco^o] MUS, COMF L12RAK , Brigham Young University "^^ Science Bulletin H- J MALE SPHAEROPTHALMINE MUTILLID WASPS OF THE NEVADA TEST SITE by WILLIAM E. FERGUSON BIOLOGICAL SERIES — VOLUME VIII, NUMBER 4 JUNE, 1967 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Dorald M. Allred, Department of Zoology and Entomology, Brigham Young University, Provo, Utah Associate Editor: Eabl M. Cheistensen, Department of Botany, Brigham Young University, Provo, Utah Members of the Editorial Board: J. V. Beck, Bacteriology C. Lynn Hayward, Zoology W. Derby Laws, Agronomy Howard C. Stutz, Botany WiLMER W. TAN^fER, Zoology, Chairman of the Board David L. Hanks, Botany Ex officio Members: RUDGER H. > Science Ernest L. Olson, Chairman, University Publications RuDGER H. Walker, Dean, College of Biological and Agricultural Sciences The Brigham Young University Science Bulletin, Biological Series, pubhshes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Publications, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University. Orders and materials for hbrary exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah. Brigham Young University Science Bulletin MALE SPHAEROPTHALMINE MUTILLID WASPS OF THE NEVADA TEST SITE by WILLIAM E. FERGUSON BIOLOGICAL SERIES — VOLUME VIII, NUMBER 4 JUNE, 1967 MUS. COMP. ZOOL. LIBRARY OCT 30 1967 HARVARD UNIVERSITY TABLE OF CONTENTS INTRODUCTION 1 ECOLOGY OF THE TEST SITE 2 MATERIALS AND METHODS 2 HISTORICAL REVIEW 4 MORPHOLOGICAL CHARACTERS 6 ACKNOWLEDGMENTS 7 SPECIES ACCOUNTS 8 AcropJtotopsis eunjgnuthus 8 Dihphotopsis concolor crassa 8 D. concolor puron 9 AcanthopJiotopsis falcifomus falciformis 9 Sphaeropthalma (Micromutilla) acontia 9 S. becki 9 S. brachyptera II S. difjicilis 11 S. macsicaini 12 S. pallida 13 S. parapenalis 14 S. sonora 16 S. yumaella 17 Sphaeropthalma (Photopsis) angulifera 17 S. blakeii 17 S. ferruginea 17 S. helicaon 17 S. unicolor 18 Sphaeropthalma (Photopsioides) amphion 20 Odontophotopsis (Odontophotopsis) armata 21 O. clypeata 21 O. cookii 21 O. inconspicua inconspicua 21 O. microdonta 22 O. obliqua , 23 O. qtiadru^pinosa 24 O. serca 24 O. setifera 24 Odontophotopsis (Periphotopsis) mamata 25 Dasymtttilla gloriosa 25 D. paemdata 25 D. satanas 25 LITERATURE CITED 26 MALE SPHAEROPTHALMINE MUTILLID WASPS OF THE NEVADA TEST SITE' by William E. Ferguson- INTRODUCTION In 1959, ecological studies of the fauna at the Nevada Test Site were begun by Drs. Dorald M. Allred and D Elden Beck of the Department of Zoology and Entomology, Brigham Young Uni- versity'. In 1962 I was asked to identify the mutillid wasps which had been taken in large numbers, chiefly from can pit-traps (Fig. 1) that Fig. 1. Can pit-trap with open end flush with the ground surface. Masonite cover is raised out of position at left. had been in place at the same localities for several years. However, only the females of noc- turnal species were well represented. In August, 1964, I was invited to study the mutillid fauna at the test site, and collect the males of the species. The females remain largely unstudied, unnamed, and have been grouped as one genus, although the males are much better known and are separable into several genera. Only three published accounts of se.x correlations among the nocturnal mutillids appear in the literature. It was assumed that with the large numbers of females at hand, the sexes of die species could be correlated in spite of the great se.xual dimor- phism. Attempts to correlate the se.xes at the test site were unsuccessful. Possible reasons are as follows: (1) The apterous females were col- lected between 1959 and 1962, but most of the males were taken only in August, 1964, under different ecological conditions and by different collection methods, even though attempts to col- lect them were made at other times of the year. (2) Cycles of abundance are known to change considerably over the years (Ferguson, 1962). (3) Light traps are highly selective for males, and attract them from large areas. (4) Can pit- traps are nonattractive to females and males, but since the females must travel on the ground, they are more liable to find the covered traps by chance, and fall into the cans. (5) Males can fly out of the cans, whereas the females cannot es- cape. (6) Males which fall into the cans are more likely to be eaten by lizards and arthropods also caught in the traps, since the male mutillids are more active, less heavily sclerotized, and in con- trast to the females, cannot sting. The best method of asscK-iating the males and females at the test site apparently must involve a thorough study of females from areas outside of the test site. Through study of the distributional pat- terns of the females, it may be {x>ssible to cor- relate them with males by a process of elimi- nation based on geographical occurrence. The large number of unnamed females must be de- scribed, and all named females must be re- described. This will be a considerable task that may take several years, and naturally is not within the scope of the present study. Although the diurnal mutillids in this study are rather well known, even the males of noc- turnal species are pB.Y.U. AEG. Report No. €00-1326-4. Field work related to this study was done under A. EC. ronlraits .\T{ 1 1-1 )78ri and AT Ol-DUJiJ awarded to Brigham Young University. Study of the five species of Sphaeroplhalma. subgenus Pholopsh, reported herein, was greatly aided by N.S.l'. Grant GB-27+5 to the author, for "A Biosysteniatic Study of Nocturnal Mutillid Wasps in the Subgenus Pholopsis of North Anierica." ^Associate Professor of Entomology. Department of Biological Sciences, San Jose State College, San Jose, California, 95114. "For a detailed discussion on the ecology of tlie lest site, refer to Allred, Beck, and Jorgensen (1963a, 1963b). Brigham Young University Science Bulletin and distributionally. This study not only makes known the sphaeropthalmine Mutilhdae of the test site, but contributes importantly to our knowledge of the taxonomy and distribution of the nocturnal Mutillidae. Furthermore, it has shown that even the former "ground zeros" of nuclear detonations are reoccupied by the mutil- lids within a few years following a blast or series of blasts. The barren area surrounding ground zero apparently is attractive as a nesting site for various species of flying aculeate Hy- menoptera, which become hosts of the para- sitic mutillids. The host species may find food for provisioning their brood cells in the soil, either inside or outside of the barren area. The flightless female mutillids then have reinvaded the barren area in one or both of the following ways: (1) by walking from the undevastated perimeter back into the barren area, or (2) by surviving nuclear blasts while in nests of their hosts deep in the soil, and emerging into the devastated area after the blast. There is interest as to whether or not nuclear testing will cause genetic abnormalities that will result in defor- mation, "monsters," and other unusual genetic traits. A careful search was made for such an- atomical changes among the Mutilhdae col- lected at the test site. Apparently the Mutilhdae are not anatomically different from those outside of the test site. An organism's genetic adjust- ment to its environment is like an accurately adjusted chronometer. Anything diat happens to that adjustment will most likely be deleterious. Hence, one would e.xpect that genetic changes in creatures of the harsh desert environment would prove fatal in an extremely high percent- age of instances. Abnormalities (unfavorable genetic traits) should neither survive nor spread commonly throughout a normal, surrounding jjopulation. This appears to be true of the mutillid population samples, if such genetic aberrations have occurred. ECOLOGY OF THE TEST SITE The Nevada Test Site of the Atomic Energy Commission is located in southern Nye County, Nevada, at elevations from approximately 2800 ft, at Fortymile Wash, to 7694 ft on Rainier Mesa. The vegetation ranges from halophytes typical of the Mojave Desert playa margins, to the open stands of Pintis monophylla and Arte- misia tridentata at higher elevations. The area is especially interesting because the boundary of the Mojave and Great Basin Deserts cuts in an east-west direction through the test site near its middle. More detailed discussions of the ecology of the test site were prepared by Allred, et ai. (1963a, 1963b). MATERIALS AND METHODS Specimens from the test site were collected principally in sheltered can pit-traps in the soil (Fig. 1) and from light-traps of three kinds: ( 1 ) a 200-watt incandescent bulb suspended from a tripod above a ground-level, white cloth sheet; (2) a one- or two-mantle Coleman gaso- line lantern above a white sheet or water-filled trap (Fig. 2); and (3) an ultraviolet hght source (Fig. 3). Gasoline lanterns and ultra- violet lights appear to be equally effective in attracting mutillid wasps. The use of lights above sheets permitted hand-catching of the mutillids which were attracted to the lights. These were placed separately into cyanide killing jars, thus avoiding the disadvantage of moth scales and damage by other insects as is typical of most light traps. Specimens were returned to the laboratory and pinned before drying, and the genitalia were extruded. Ultraviolet and incandescent light-traps were operated all or part of the night. Insects of many kinds were caught as they fell into deter- gent water beneath the light sources. The deter- gent used was American Cyanamid's Aerosol- OT. However, most other detergents were al- most equally effective in reducing surface ten- sion, thus permitting the insects to "drown" quickly, rather than struggle on the water sur- face. The insects apparently were immobilized by the greatly reduced oxygen supply in the water, but after submergence during the night, most of the insects became active again within a few minutes to several hours after removal from the water. The nocturnal mutillids were MUTILLIDS OF THE NEVADA TeST SiTE Fig. 2. Gasoline lantem light-trap. A large aluminum salad-ring mold is buried with its edge flush with the ground surface, and is filled with detergent water. The lantem covers the center hole and pro- vides hght all night. washed individually in clean detergent water, rinsed for a few minutes in 70%' and 90% ethyl alcohol, dried briefly on a paper towel, and pinned. This procedure prevented matting of the pubescence and wings, especially when the pinned specimen was blown upon with a strong air stream from the lips. The unattended, all- night light-traps were especially effective for trapping the small, hard-to-catch individuals, for collecting a series of a common species, and for catching individuals of species which were present in low densities. Although females are positively phototaxic at night, few were attracted into the light-traps. Probably this was due to ground-level shadows of surface irregularities and plants, and to low mobility of the females. Can pit-traps were outstandingly effective in trapping female mutillids, but either were much less effective in trapping males, or allowed them to escape after entering the traps. Speci- mens from can pit-traps had been preserved in alcohol, which is unsuitable for mutillids. Fur- thermore, some of the vials lost alcohol through their cork stoppers, thus concentrating the glue- like dissolved body fluids and dirt. All alcoholic specimens required washing in 70? and 90f ethyl alcohol, and ether, prior to pinning and identification. Nevertheless, some could not be restored adequately for certain identification be- m>^ Fig. 3. Generator-operated, 40-watt ultraviolet Ught- trap supported on folding wooden legs. The shal- low depression beneath was formed with a shovel, lined with a plastic sheet, and filled with deter- gent water. cause of altered integumental color, matted hairs and wings, and insoluble "glue" and dirt on the integument. Each specimen was assigned a reference num- ber and a code number, both of which relate to collection records. Because of inadequate knowledge of female behavior, it is impossible at the present time to determine the influences of certain ecological factors on the two sexes. However, during the light-trapping in August, 1964, surface activity of the males was greatly reduced or even stopped by bright moonlight, rain, strong winds, or low temperature. Nocturnal tempera- lures in the range 80° to 100° F apparently are optimum for the flight of the males, but activity was noticeably reduced in the range of 70° to 80° F, and was insignificant below that tempera- ture range. Distribution records for the test site are frag- mentary and are localized because of pre- planned, intensive collections in selected areas, especially those closer to headquarters at Mer- cury, and adjacent to roads. iVIore informative can pit-trap and light-trap survey data for the Mutillidae should include replicated transects between altitude extremes and in the same vege- tation types at comparable sampling stations. All-night, gasoline lantern, detergent-water light- traps operated once or twice each week would catch males and some females, but permanently installed can pit-traps for females could be oper- ated continuously at each station. Such a samp- ling method would provide much more trust- worthy information on sex associations and relative abundance. Brigham Young Univebsity Science Bulletin Specimens on which this study is based came primarily from Frenchman Flat and the hillsides north-northwest of Mercury. Locality IB, in Yucca Flat, and Cane Springs were secondary collection sites, and collections from the higher valley slopes and top of Rainier Mesa were relatively few. Figure 4 will assist in locat- ing the areas referred to in this paper. For more exact collection code interpretation, refer to Allred, et al (1963b). The light-trap collections of nocturnal male mutillids were limited at the test site, but several factors permit making generalizations about the distribution of species there. First, maximum and minimum temperatures, which were re- corded for several years at a number of localities at the test site, indicate that August is the most favorable time for nocturnal activity of mutillids. Consequently, one can expect the largest num- bers of individuals and species to be active dur- ing that month, when most of the light-trapping was done. Furthermore, I have collected exten- sively to the north of the test site in the Great Basin Desert, and to the soutli in the Sonoran Desert. Collections from these two deserts have been compared with each other, with smaller collections from the Mojave Desert, and with collections from the test site, as a basis for specu- lating or generalizing about the occurrence of mutillids at the test site. These speculations must not be confused by the reader with the presently known facts of distribution, which are simply the raw collection data. HISTORICAL REVIEW Prior to 1958, the classification of nocturnal Mutillidae and the description of species were based largely upon morphological characters which today are not recognized as reliable indi- cators of phylogenetic relationship. Schuster's important paper was published in 1958, but had been written approximately twelve years earlier. It included many new, very distinct species, focused attention on newly discovered taxonomic characters, and included a new classification. However, several characteristics of his work have impeded rapid improvement of our taxo- nomic concepts and classification of the nocturn- al Mutillidae. These impediments include ( 1 ) validation of most specific names only in keys rather than in adequate descriptions, (2) lack of precise type locality or distribution data, (3) lack of information on the range of variability of each species, (4) lack of designated holotype specimens for many names, and (5) apparent Key to the important collection localities shown in Fig. 4. 1. Mercury — incandescent and ultraviolet light collections 2. Hillside 0.8 mi NNW of Mercury — incandescent and ultraviolet light 3. lOS study area — can pit-traps 4. Yz mi E of Mercury Hwy. on Kay Bunker Rd. — light collections 5. Cane Springs — can pit-traps and light collections 6. 0.3 mi W of Y on Rainier Ntesa — light collections 7. Rainier Mesa Rd. at ponds near Tunnel E — light colections 8. 1 mi W of Y on Rainier Mesa — light collections 9. 0.9 mi W of Area 12 residences — light collections 10. 3 mi SSW of Area 12 residences on new road to Fortymile Canyon — light collection 11. IB study site — can pit-traps, one light collection at 1BB25 12. 0.2 mi E of Mercury Hwy. at Checkpoint Pass — light collection 13. Old Mercury Hwy., 1 mi from Mercury Hwy., at 3400 ft (extreme NE corner of Specter Range Quad- rangle)— light collection 14. 5A study area — can pit-traps and light collections 15. Old Mercury Hwy. in barren, extensive sandy wash, 2 mi from intersection with Mercury Hwy. in French- man Flat — light collection 16. 5E study area — can pit-traps and light collections 17. 4A study area — can pit-traps 18. ECB study area — ultraviolet light collections 19. CP study area — Allrcd live-catch, box-t)'pe traps 20. 6A study area — can pit-traps 21. IF study area — can pit-traps MUTIILIDS OF THE NEVADA TeST SlTE PAHUTE ME^A i ? SHOSHONE M UNTAIN LiRRE4-FRAN •-□ OTRIPLEX-KOC ... □ ~-- ■ D.-- n COLEOGYNE □ ""-;'"" SALS0L4 Bi ^ ' MOUNTAINOUS *■" D _,_ 1^ r NEVADA TEST SITE SCALE 0 1 2 3 4 B 1 L_J \ \ ^MLE Fig, 4. Principal collection sites of mutillid wasps at the Nevada Test Site. See key on facing page. Brigham Young Univebsity Science Bulletin loss of some holotypes and lack of information about locations of many others. In July, 1959, one year after publication and approximately thirteen years after preparing the first manuscript, Schuster attempted to find and label type specimens representing the names that he had validated in his keys. Some were found and deposited in major repositories, some were not found, whereas others apparently remain in the Schuster collection although they are the property of institutions from which they were borrowed many years ago. Because Schuster largely neglected publica- tion of tyjx; data, it is included here, when known, for all species. Missing type specimens, inaccessibility of others, and unpublished type data have affected this study in several ways: ( 1 ) inability to verify the accuracy of some steps in Schuster's key leaves identification of a few species in question, (2) a few specimens from the test site have been retained by me without identification because they represent un- described or inadequately described species, and type material is unavailable for comparison. MORPHOLOGICAL CHARACTERS Certain terminology and measurements used by Schuster have never been adequately ex- plained; hence, I have included herein my inter- pretation of these, and an alteration of some other aspects of his terminology. Measurements are most meaningful to mutil- lid taxonomists when expressed in units rather than in a ratio or other expression. The taxono- mist must be able to judge the accuracy of mea- surement when necessary, and construct what- ever ratios may be desirable from the unaltered measurements. Consequently, in descriptions of new species I have given most measurements in ocular micrometer units, which equal one-sixti- eth of a millimeter. The limits of accuracy in using my ocular micrometer at 60x magnifica- tion are plus or minus one-half unit. Measure- ments on small individuals, therefore, are sub- ject to relatively greater error. Relative to pubescence, I have used the temi simple hair to describe hairs which have smooth rather than barbed surfaces. Brachy- plumose refers to hairs which have many barbs, the lengths of which are appro.ximately equal to or less than the diameter of the hair at the place where each barb is located. Plumose is reserved for hairs which have longer barbs on at least part of their length. Body size is variable and dependent on the amount of food available to the mutillid larva (Ferguson, 1962). Sculpture is most regularly and deeply expressed in large specimens, but becomes more irregular and less distinct among the smallest individuals of a sj>ecies. Directions of structures on the head have traditionally been expressed as though the mutillid head were prognathous. Since the head actually is hyjxjgnathous, I have given direc- tions on the head accordingly, and hope tliat this practice will be followed in future studies of the nocturnal Mutillidae. Head shape has been described variously by Schuster, including "evenly semicircular behind the eyes," "temples bulging," "subquadrate," and others. In many cases, species which according to Schuster have the head other than "evenly semicircular" actually have the head outline, dor- sal to the eyes, conforming almost exactly to an arc of a circle, rather than as described in one of the above phrases which indicates otherwise. Head shape can be determined precisely by drawing the head outline, in frontal view, with the aid of an ocular grid or camera lucida. On the basis of the height and width of the head above the eyes, one can determine geometrically the center of an arc with those proportions, and trace a true arc with a compass, over the head outline. I have done this with type specimens and have shown elsewhere in this paper that Schuster has imperfectly described head shapes of some species. The interpretation "subquad- rate" probably is an illusion, at least sometimes, based on large head size, which results in rela- tively longer radius curvature, but not the sub- quadrate condition. Eye length, as used here, includes measure- ment of the eye to the margin of the ocular sclerite, which is larger than the black area be- neath it. Ocellar length is the measurement of the greatest dimension of the ocellus. Ocellocular distance is interpreted here as the shortest distance from the margin of the ocellus, to the margin of the ocular sclerite, but not to the margin of the black eye pigments. NfuTILLIDS OF THE NEVADA TeST SiTE New temiinologv also is introduced here for tlie mandibular teeth. Usually the mandibles are tridentate apically, as in Figures 5, 6, and 7. The apical tooth, which is an extension of the pos- terior margin (ventral margin of Schuster), is termed here the 1°, or primary t(K>th. The t(K>th which is an extension of the anterior margin (dorsal margin of Schuster) is termed the 3°, or tertiary tooth. The smallest, often extremely small tooth between, is termed the 2°, or sec- ondary tooth. The mandibular tooth which often is found near the midpoint of the jx)sterior margin and basad of a reduction in mandible width (excision of Schuster), is here considered the basal tooth of the posterior margin (as op- posed to the apical teeth). Thorax is used here as .\Iickel has used it, to include both the true thorax and the propodeum, and apparently is synonymous with Schuster's term alitrunk. The pterostigmal cell length is measured from its separation at vein R + M + Sc, to its proximal margin, but not including the vein which encloses it distally. Marginal cell length on the costa is measured on the costal margin between the points where the enclosing veins bend most definitely toward the posterior mar- gin, but none of the posteriorly directed veins is included in the measurement. The abdominal segments are numbered as in other literature on Mutillidae, with the petiole or morphological second segment considered the first abdominal segment, and the first seg- ment of the gaster considered the second ab- dominal segment. The length of the second ab- dominal tergum is measured through the mid- line of the felt line. Penis valves, as defined by Michener (1944), are the paired structures between which the membranous endophallus issues apically. The pair is clasped laterally by parapenal lobes of the parameral plates. From the latter, die para- meres extend apicad, and the cuspes are paral- lel to and between these, but are separated from each other basally by the penis valves. The coarseness of the punctation depends on the diameter and depth of individual punctures in relation to the size of the punctured sclerite. In this paper, the following terms express the degrees of punctation in the order of decreasing coarseness: reticulate, coarse, moderate, small, fine, micropunctate. Tlie latter refers to punc- tures which are smallest of all, extremely shallow, and do not have vertical walls or sharp margins. There is a definite need for revision and modernization of mutillid terminology, as well as alteration of the description format, now that our nocturnal mutillid fauna is relatively much better known. ACKNOWLEDGMENTS I wish to thank Drs. D Elden Beck and Dorald M. Allred of Brigham Young University who made this study possible. Clive D. Jorgen- sen, field director for some of the B.Y.U. pro- jects at the test site, was directly responsible for coordinating on-site activities. Mr. Clyde Pritchet, Research Associate, was especially help- ful in providing technical assistance. I am also grateful to the following who facilitated the study of type specimens in their care: Drs. Harry Allen and the late Harold J. Grant, Jr., Academy of Natural Sciences at Philadelphia; Dr. K. C. Kim, University of Minnesota; Dr. Karl V. Krom- bein, U.S. National Museum; Mr. Hugh B. Leech, California Academy of Sciences; Dr. W. R. M. Mason, Canadian National Collection; and Dr. Clarence E. Mickel, Tucson, Arizona. Of special significance was my opportunity to study some type s{5ecimens as a part of the Brigham Young University cooperative study. I acknowledge here the loan and ownership, by the following persons and institutions, of specimens involved in this study. The following abbreviations are used throughout the text: AMNH, American Museum of Natural History; ANSP, Academy of Natural Sciences at Phila- delphia; BM, British Museum (Natural History); BYU-AEC, Brigham Young University-Atomic Energy Commission Projects; CAS, California Academy of Sciences; CDA, California Depart- ment of Agriculture; CIS, California Insect Sur- vey, University of California at Berkeley; CNC, Canadian National Collection; CU, Cornell Uni- versity; DAG, D. A. Garner; JZVV, J. Z. Warren; LACM, Los Angeles County Museum; OSU, Ohio State University; RMS, R. M. Schuster; TAM, Texas Agricultural and Mechanical Col- lege; UA, University of Arizona; UCD, Univer- sity of California at Davis; UCR, University of California at Riverside; UI, University of Idaho; Brigham Young University Science Bulletin UM, University of Minnesota; USNM, United States National Museum; USU, Utah State Uni- versity; UU, University of Utah, and WEF, Wil- ham E. Ferguson. All other records not desig- nated by the above initials were taken as part of the Brigham Young University study at the Nevada Test Site. I am especially grateful to my wife, Stephenie, for her assistance in preparing the manuscript. SPECIES ACCOUNTS In the treatments of new species of male mutillids, I have departed from the usual for- mat used for descriptions of the diurnal species. As new taxonomic characters are discovered and other characters are considered to be of little importance, species concepts change. Conse- quently the nature of the descriptions has changed and will continue to change. Coloration of the diurnal species often is helpful in identification, but in the nocturnal species it is much more uniform, and therefore much less useful for identification with the un- aided eye. Since one must use the microscope to begin identification of the nocturnal forms, even to genus, the taxonomist is aided most by having the key morphological characters mentioned in the diagnosis of each species. In order to keep the diagnosis of males brief yet meaningful, I have included descriptions of certain features of the following: (I) size, (2) mandibles, (3) clypeus, (4) mesostemal tu- bercles or processes, (5) processes of the coxae and trochanters, and (6) felt lines. The genitalia usually are of great importance taxonomically, but since the description of these structures ap- pears last in each species description, the infor- mation is found there easily. In the lists of type data, a specimen that is not definitely known to be a holotype, as dis- tinguished from lectotype or syntype, is listed thus: holotype (?). Acrophotopsis eurygnathus Schuster Acrophotopsis eurygnathus Schuster, 19.58. Ento- mol. Amer., 37(n.s.):10, 68. Type data. Holotype d" Globe, Arizona (CAS). Paratopotypes: 2 cT same data as holo- type. Paratypes: 44 cT (see Schuster, 1958:68). Discussion. Nineteen specimens were taken in June, July, and August in both can and light- traps from only two localities at the test site: between Mercury Highway and the Frenchman Flat playa (about 3200 ft), and at the Cane Springs area (4000 ft). Both of these localities are in the Larrea-Franseria biotic communinty (Allred, et ah, 1963a). These are the first rec- ords of occurrence in the Mojave Desert, and probably represent the northern limit of the species. Other published and unpublished rec- ords indicate that eurygnathus occurs in the Sonoran Desert of Arizona, but apparently not in the Great Basin Desert. Dilophotopsis concolor crassa (Viereck), new combination Odontophotopsis crassus Viereck, 1924. Can- adian Entomol., 56:112. Dilophotopsis concolor utahensis Schuster, 1958. Entomol. Amer., 37(n.s. ):87. New synonymy. Type data. Odontophotopsis crassus, holo- type cT Oliver, British Columbia, Canada (CNC type no. 754). Dilophotopsis concolor utahensis, holotype d Delle, Tooele Co., Utah (CU). Para- type cT Wadsworth, Storey Co., Nevada ( errone- ously published as New Mexico, where no such county exists). Although stated in publication that the holotype belongs in the Cornell Uni- versity collection, it probably is still in the Schu- ster collection. The new synonymy listed above is based on examination of the holotype of O. crassus, which is typical of this Great Basin Des- ert form, and readily keys to D. concolor utahen- sis in the key by Schuster (1958). Discussion. One specimen without darkened gaster was taken at study site 6A about 2 mi north Yucca Playa, and 0.6 mi east Mercury Highway (4000 ft). I have taken this subspecies commonly in all of the Great Basin Desert states. It is signifi- cant, therefore, that crassa occurs extremely rarely at the test site, and the single collection locality is within the Great Basin Desert por- tion. Dilophotopsis concolor paron (Cameron) is almost equally rare, and apparently does not occur outside of the Mojave Desert portion. These are the distributional relationships one would expect based on collections that have MUTILLIDS OF THE NeVADA TeST SiTE been made to the north and south of the test site. Dilophotopsis concolor paron ( Cameron ) SphaerophthaJma [sic] paron Cameron, 1896. Biol. CentraH-Amer., Insecta, Hymenoptera, 2:381. Dilophotopsis concolor sotwrcnsis Schuster, 1958. Entomol. Amer., 37(n.s.):88. Dilophotopsis concolor paron Mickel, 1965. Proc. Entomol. Soc. Wash., 67(1 ):1. Type data. Sphaerophthahna paron, holo- type cT (BM). Dilophotopsis concolor sonoren- sis, holotype d Gila Bend, Arizona (UM). Para- type cf Chiricahua Mts., Arizona (depository unknown ) . Discussion. This subspecies is represented at the test site by three sf>ecimens taken near porch lights on the CETO^ laboratory building at Mercury (.3800 ft). Apparently it does not even occur at lower elevations in Frenchman Flat. This form with the darkened gaster is characteristic of the Sonoran Desert, and this is the first published record of its occurrence in the Mojave Desert. Acanthophotopsis falciforniis falciformis Schuster Acanthophotopsis falciformis falciformis Schu- ster, 1958. Entomol. Amer., 37(n.s.):13, 108. Type data. Holotype c? Palm Springs, Cali- fornia (UM). Paratypes: 2 d Palm Springs, California (one at UM, location of other un- known); 1 cT 15 mi E Sombrerete, Mexico (de- pository unknown ) . Discussion. Of the seven specimens taken at the test site, three were attracted to porch lights on the CETO building at Mercury ( .3800 ft) in August. At locality 5A (.3200' ft) in Frenchman Flat, two came to ultraviolet light and one to incandescent light in August, but a fourth specimen was taken from a can pit-trap in June. No records are known to me for the Great Basin Desert. Earlier published records include only the type series. Sphaeropthalma (Micromutilla) acontia (Fox) Photopsis nanus Ashmead, 1896. Trans. Amer. Entomol. Soc, 23:181. Preoccupied. *C;ivil Etfecti Test Operations of the U.S. .\tomic Energ.v Comni Mutilla acontius Fox, 1899. Trans. Amer. Ento- mol. Soc, 25:266. Mutilla Ashmeadii Fox, 1899. Trans. Amer. En- tomol. Soc, 25:289. New name for P. nanus Ashmead. New synonymy. Type data. Photopsis nanus, holotype cT Tucson, Arizona (USNM type no. 3279). Mutil- la acontius, holotype d Las Cruces, New Mexi- co (ANSPtypeno. 4&14). Discussion. Twenty specimens of this small- sized species were taken in both incandescent and ultraviolet lights in the Mojave Desert por- tions of the test site in late July and August. Sf>ecific localities include only Mercury, Cane Springs, and Rock Valley. Individuals of this species are among the smallest noctunial male Mutillidae in North America, averaging .5 mm in length. Consequent- ly they are inconspicuous and usually are not captured unless a special effort is made to catch the smallest mutillids flying among the other insects around a light source. With such effort I collected 17 specimens in one month, whereas other collectors at the test site routinely col- lected only five in a period of about five years. New synonymy is based on personal exami- nation of the two type specimens. The genitalia of acontia are like those of S. (Micromutilla) pallida (Blake), and are unmistakably distinct from those of all other small nocturnal Mutil- lidae known to me. The cuspis is very short, scarcely extending beyond the tip of the penis valves. This character, coupled with the scarcely excised mandibles, relatively slender petiole, and parameres not apically divergent, facilitates identification of the species regardless of slight differences in size and sculpture. The only reliable records of the distribution of acontia include the type localities of Tucson, Arizona, and Las Cruces, New Mexico. La Cueva, New Mexico (5300 ft) was cited by Melander (1903:318) as a collection locality; however, the altitude probably is too high for acontia, and Melander 's mention of long pubes- cence suggests that some other, hairier species was misidentified. Sphaeropthalma (Micromutilla) becki, new species Diagnosis. Male: Length 7 mm. Integument of antennae and body testaceous, legs slightly lighter. Mandibles with posterior margin excised to apex, reducing width of apical one-half to ap- Brigham Young Uniyersity Science Bulletin proximately one-half of basal width, with sub- tending tooth angulate and without a deep, rounded notch distally. Clypeal disc longitudin- ally concave near apical margin, transversely straight, with a pair of blunt, short teeth apical- ly, basally not produced into a carina or tubercle. Mesosternum moderately, shallowly punctate, without tubercles or processes. Coxae and tro- chanters unarmed. Sternal felt lines subequal in length to those on second tergum. Pygidium polished, very shallowly micropunctate on apical one-fourth. Description. Head: Integument polished, with outline dorsad of eyes forming an arc of a circle in frontal view; punctures fine, sparse. Measurements in micrometer units: eye length 39, ocellocular distance 17, ocellar length 15. Interocellar area darkened. Mandibles overlap dorsoventrally, anterior margin of distal one- half not twisted dorsad; bearing a distinct carina from base to tertiary apical tooth (Fig. 5). Apical mandibular teeth with 1° tooth basal Fig. 5. Mandible of Sphaeropthalma (MicTomutiUa) becki. width Ix and length 2x the 3° tooth; 2° tooth basal widtli Hx the 3° tooth. Clypeal surface polished, with sparse micropunctures bearing long or short simple hairs; apex not impressed below anterior margin of mandibles. Antenna with first flagellar segment short, 0.6x lengtli of second segment. Frontal line and preocellar pit distinct. Thorax: Pro- notum dorsally and laterally shallowly reticulo- punctate, except polished and shallowly rugose on posterolateral margin. Mesonotal punctures fine, shallow, one puncture width apart, with flat, pohshed interspaces. Parapsidal lines with posterior one-third impressed. Mesopleuron widi anterior impressed area polished, extremely shallowly reticulate, with sparse, simple hairs; f>osterior raised area reticulate anteriorly, im- punctate, polished posteriorly. Propodeum coarsely, shallowly reticulate. Pterostigmal cell 0.38x length of marginal cell on costa. Abdomen: Plumose fringes dense only on second tergum apically. First tergum polished, finely punctate medially, moderately punctate laterally, apically 0.56x greatest width of second tergum. First sternum arcoiate in cross sec-tion, anterior one- half with a mediolongitudinal carina not termi- nating anteriorly in a distinct tooth. Second ter- gum polished, finely, sparsely punctate with sparse, brachyplumose, recumbent hairs and long, simple, erect hairs; felt lines 0.4x length of tergum laterally. Second sternum moderately, shallowly, c-ontiguously punctate with sparse, brachyplumose, recumbent hairs and long, simple, erec-t hairs; anteromediall}' tumid. Geni- talia: Parameres arcuate dorsad, laterally straight, glabrous. Cuspis 0.4x free length of paramere, rodlike, with approximately 20 mesal- Iv-directed short hairs on mesal surface. Female. Unknown. Discussion. Sphaeropthalma becki most close- ly resembles acontia in external features, but the apical one-half of the mandibles is more tapered in the latter species. The very short marginal cell in becki facilitates distinction from acontia, which has the marginal cell approximately equal to the length of the stigma. Furthermore, the genitalia of the two species are very different. In becki the cuspis is rodlike and approximately four-tenths the free length of the parameres, but in acontia the cuspis scarcely exceeds the length of the penis valves. The holotype was selected for its large size, which best expresses the characteristics of the species. Size ranges from 3 to 7 mm, with the smaller specimens generally having both sculpture and abundance of plumose hairs reduced. In many of the smaller specimens, the frontal line is not visible. Type data. Holotype cT (USNM): Hillside 0.85 mi NXW Mercury, Nye Co., Nevada, at light, W. E. Ferguson. Actual label data: Mer- curv, Nevada, N.T.S.; BYU-AEC Code MMT; \Tn-23-64; ref. no. 1263. Six d paratopotypes same data. Eleven paratopotvpes same locality, at light; 1 c^ VlII-3-64 (no. 1107); 2 d VIII-10- 64 (no. 1189); 4 d" VIIl-20-64 (no. 1232); 4 d" \TII-21-64 (no. 1251). Twenty-three paratypes: CALIFORNIA. Riverside Co.: 6 mi S Pabn Springs. 1 c? VI-20-64, at light (WEF). San Bernardino Co.: 12 mi N Earp, 2 d V-5-53 (WEF). NEVADA. Clark Co.: 8.4 mi SE Indian Spring, 9 cT \TII-22-64, at Hght (WEF). Nye Co.: West side Frenchman Flat, 3 d" VIII-6- 64, at light (no. 1117); 1 d" VH-13-61, can trap (no. 25); 2 d' VII-19-65, ultraviolet hght (nos. 91 and 101); Cane Springs vicinitv. 2 d VHI-7-64, at light (no. 1134); Jackass Flats, 3 d VIII-7-62, ultraviolet light (nos. 1086-1088); 5 d ¥11-20-62, MllTILLIDS OF THE NeVADA TeST SiTE 11 ultraviolet light (nos. 1101-1105). Washoe Co.: 1 mi NW Nixon, 2 d VIl-8-61, at light (WEF). Distribution. This small mutillid is reasonably common at least locally throughout the Sonoran Desert of California, the Mojave Desert, and at least the lower valleys of the western Great Ba- sin Desert. Its abundance is not indicated by most collections because its small size makes it inconspicuous at light-traps, and a special effort must be made to catch individuals as they fly among the larger insects. Spliaeropthalma (Micromutilla) brachijptera (Schuster) Photopsis brachijptera Schuster, 1945. Pan-Paci- fic Entomol., 21:149. Spliaeropthabna (Micromutilla) yavapai Schu- ster, 1958. Entomol. Amer., 37(n.s. ):19. New synonymy. Type data. Photopsis brachijptera, holotype cT Berkeley, California. Sphaeroptlialma yavapai, holotype cf Kirkland, Peeples Valley, Yavapai Co., Arizona (data provided verbally by Schu- ster). Both specimens are the property of Cor- nell University, but were still in the Schuster collection in 1959. A paratype of yavapai is in the University of Minnesota collection. The unique type of brachyptera appears to be an intersex, because the reductions of wings, eyes, and ocelli are comparable reductions in male- ness. Such an individual might result from either genetic, nutritional, or other environmental in- fluences. I extracted the genitalia far enough to ascertain that they are identical to those of the normal yavapai. The small size of the brachyptera holotype (4.7 mm), as in small specimens of other species, also is the cause of reduced sculpture and pubescence. Consequent- ly, this specimen should be compared with larger specimens in the 7-9 mm range as well as with a series in the intermediate range, so that the transition of character expressions can be observed. Discussion. Thirty-one specimens of this widespread species were collected from the test site at incandescent and ultraviolet light-traps, but not in can pit-traps. Although brachyptera occurs in both the Great Basin and Mojave Desert portions of the test site, it was not col- lected with other species on the gently sloping floors of the valleys. All collections were made during August, which correlates with the time of intensive light-trap collecting rather than with the activity cycle of brachyptera. Collection data follow: 3 d hillsides near Mercury (4000-4200 ft); 14 d 0.9 mi VV area 12 residences (.5500 ft); 1 d" Rainier Mesa road (5800 ft); 10 d" 3 mi S area 12 residences (6200 ft). The vegeta- tion at these localities included Quercus gambelii and Pinus monophijlla, Junipcrus osteosperma and Coleogijne ramosissima, nearly pure C ram- osissima, and "mixed vegetation," but not Larrea- Franseria, and other communities of the valley floors. Specimens in my collection are from the area including the Coast Ranges of California, east to iVIono Lake and the Nevada Test Site, south to the Rio Mayo of Sonora, Mexico, and eastward in Arizona to the Chiricahua Mountains. In spe- cimens from the latter area the gaster, legs, an- tennae, and pterostigma are considerably dark- ened. This correlates with the integumental darkening of other species which occur in the same area (see discussion of S. pallida). Al- though brachyptera has not been collected from the valley floors at the test site, I have taken it in similar places where Larrea divaricata grows at Organ Pipe National Monument in Arizona, and 8 mi NE of Mesa, Arizona. Sphaeroptlialma (Micromutilla) difficilis ( Baker ) , new combination Photopsis difficilis Baker, 1905. Invert. Pacifica, 1:114-115. Sphaeroptlialma (Micromutilla) maricopella ma- ricopella Schuster, 1958. Entomol. Amer., 37 ( n.s. ) : 17. New synonymy. Spliaeropthalma (Micromutilla) maricopella pur- isimella Schuster, 1958. Entomol. Amer., 37 ( n.s. ) : 17. New synonymy. Sphaeropthalma (Micromutilla) maricopella cas- tanea Schuster, 1958. Entomol. Amer., 37 (n.s. ):17. New synonymy. Sphaeropthalma (Micromutilla) californiense californiense Schuster, 1958. Entomol. Amer. 37 (n.s.): 18. New synonymy. Sphaeropthalma (Micromutilla) californiense fuscatella Schuster. 1958. Entomol. Amer., 37 (n.s.): 18. New synonymy. Sphaeropthalma (Micro mitt ilia) quijotoa quijotoa Schuster, 1958. Entomol. Amer., 37(n.s.):18. New synonymy. Sphaeropthalma (Micromutilla) quijotoa parrasia Schuster, 1958. Entomol. Amer., 37(n.s.):18. New synonymy. 12 Brichaai Young University Science Bulletin Type data. Photopsis difficilis, holotype d Claremont, California (CU); Sphaeropthahtui maricopella maricopella, holotype d California (CU); paratypes 2 d" Hopkins Well, Riverside, California (CIS). Sphaeropthalma californiense calif orniense, holotype d Mt. Diablo, Contra Costa Co., California (CAS); paratypes: d Antioch, Contra Costa Co., California (WEF); d Antelope Is., Davis Co., Utah (CIS). Al- though type localities were given in Schuster's key for each of the other new taxa (except fuscatella) listed above in synonymy, apparently type specimens are not available for study be- cause they have not been designated for those Discussion. Twenty-seven specimens were collected from Mercury, Frenchman and Yucca Flats, and Cane Springs at the test site, but not in the Pinyon-Juniper community on Rainier Mesa. This is a very widespread species which is represented in my collection by specimens from many localities in the Great Basin Desert and throughout the southwest, from coastal Cali- fornia and Mexico east to Las Cruces, New Mexico. The taxonomic treatment of this species given by Schuster (1958) overlooked the valid name difficilis, and stressed differences without equal emphasis on similarities. Consequently, the | small, geographically isolated samples which he studied appeared to him distinct enough to bear different names. The unifying characteristics of the many slight morphological and color variants are the convex clypeus, deeply excised mandibles, more or less petiolate condition of the first ab- dominal segment, and the form of the genitalia. The cuspis is more or less rodlike, slightly bulb- ous at the apex, and approximately two-thirds the length of the parameres. It bears numerous ven- trally-directed long hairs on its entire ventral | surface. In addition, the parameres at about mid-length bear at least a few, and usually many long hairs directed mesally and attached to the mesal surface. I have studied the holotypes of difficilis and californiense californiense, and paratypes of maricopella maricopella. On the basis of a two- week visit with Schuster in 1959 and subsequent correspondence with him, I judge that type specimens have never been designated for the other names listed in synonymy above. Schuster's 1958 key, therefore, provides the only avail- able descriptive information relating to these names. The range of eye sizes given by him clearly indicates that eye size differences form a continuum rather than distinctly different categories. He expressed the distance between the eye and the lateral ocellus as unity, and compared the longest dimension of the eye to that unit of length, with the following results. Eye Size Range Taxa 1.6 - 1.9 2.1 - 2.2 2.2 - 2.35 2.5 - 2.6 2.6 - 2.8 californiense californiense and c. ftiscatella maricopella maricopella and m. castanca quijotoa quijotoa quijotoa parrasia maricopella purisimella It is especially significant that the center of the range of eye size in "subspecies" of marico- pella is not occupied by a subspecies of marico- pella, but a different "species. " Sphaeropthalma (Micromutilla) macswaini, new species Diagnosis. Male: Length 7.5 mm. Integu- ment testaceous except head, antennae and legs slightly lighter. Mandible with posterior margin deeply excised, with subtending tooth large, pro- tuberant, rounded (Fig. 6). Clypeal disc longi- Fig. 6. Mandible of Sphaeropthalma (Micromutilla) macswaini. Numbers indicate transverse measure- ments. tudinally concave, transversely straight, with margin truncate and carinate apically, and median base not protuberant or carinate. Meso- sternum with one or two tiny denticulations on each side of mid-ventral line. Coxae and tro- chanters unarmed. Sternal felt lines approximate- ly two-thirds length of those on second tergum. Pygidium polished, with extremely shallow micropunctures. Description. Head: Integument polished, with outline dorsad of eyes inflated dorsolateral- ly, not forming an arc of a circle in frontal view; h-tuTlLLIDS OF THE NeVADA TesT SiTE 13 punctures fine (diameters 1-2 units), sparse. Measurements in micrometer units: eye 42, ocell- ocular distance 19, ocellar length 15. Interocel- lar area slightly darkened. Mandibles overlap dorsoventrally; distal one-half not twisted on its a.xis; carina of anterior margin well developed, complete, expanded anteriorly at bend; mea- surements given in Fig. 6. Apical mandibular teeth with 1° tooth basal width 0.6x and length 2.x the 3° tooth; 2° tooth basal width 0.25x and length Ix the 3° tooth. Clypeal surface jx>lished, glabrous, impunctate, ajiex not impressed below anterior margin of mandibles. Thorax: Pronotum dorsally and laterally coarsely, shallowly punc- tate except rugose on posterolateral margin. Mesonotum finely, shallowly punctate, the punc- tures approximately four diameters apart, with interspaces polished, glabrous. Parapsidal lines with posterior one-half impressed. Mesopleuron with anterior impressed area finely, shallowly, sparsely punctate, the pimctures bearing long and short, brachyplumose hairs; posterior raised area reticulate. Propodeum coarsely reticulate. Pterostigmal cell testaceous, 1.2x length of marginal cell on costa. Abdomen: Plumose fringe moderately dense on apex of second ter- gum only. First tergum polished, minutely and sparsely pimctate dorsally, finely punctate lat- erally, apically 0.52x greatest width of sec- ond tergum. First sternum arcuate in cross sec- tion, anterior one-half with a mediolongitudinal carina, not terminating in a tooth anteriorly. Second tergum polished, moderately punctate, the pimctures one to two diameters apart ex- cept contiguous anterolaterallv, with short, brachv|:>lum()se, recumbent hairs and long simple hairs; felt lines one-half length of tergum lateral- ly. Second sternum moderately, closely punctate, with brachyplumose recumbent hairs and sparse, erect simple hairs, anteromedially tumid. Genitalia: Parameres arcuate dorsad and mesad. Cuspis length two-thirds free length of para- mere, basaliy large, gradually tapered to a jx)int apically, arcuate mesad, with mesal surface con- cave and bearing short, mesally directed hairs. Female. Unknown. Discussion. Although the mesostemum of macsicaini bears tiny denticulations, it is not re- ferable to either Photomorphus or Odontopho- iopsis on the basis of the peculiar genitalia. By ignoring the tiny denticulations, macsicaini keys readily to Micromutilla, species group hyalina, in Schuster's 19.58 key, but does not fit either alternative of couplet three because the petiole is not obviously petiolate, but it has a well- developed clypeus. Since the species of Micro- mutilla have more varied genitalia than Photo- morphus and Odontophotopsis, macsicaini seems to fit best into Micromutilla, at least until more of the undescribed species have been studied and described. Size ranges from about 4 to 9 mm. None of the specimens has well-developed abdominal plumose hairs, even on the second tergum. The cuspis length varies between two- thirds and three-fourths the free length of the paramere. The combination of the distinctive mandibles, with the broad tertiary tooth and the \'ery prominent, rounded tooth on the posterior margin, plus the unusual genitalia, permit quick differentiation of macswaini from all of the other described nocturnal mutillids. Type data. Holotype d (USNM): 2.1 mi NE Mercury, Nye Co., Nevada; VHI-24-64; at light, W. E. Ferguson. Actual label data: Mercury, Nevada, N.T.S.; VIII-24-64; BYU-AEC Code 5CH(T); ref. no. 1277. Fifteen paratypes: ARI- ZONA. Yuma Co.: Dome, 1 d ¥11-21-24 (CAS). CALIFORNIA. Imperial Co.: Kane Springs, 1 d" X-3-23 (CAS); 2 d" foot of mts. W Salton Sea beach, VII-23-52 (CAS). Riverside Co.: Magnesia Canyon, 1 d VII-2-52, 4 d ¥11-20-52 (UCD); Dead Indian Canvon, 3 d VHl-6-65, at hght (WEF); Palm Desert, 1 d IV-11-50 (UI); Pahn Springs, 1 d V-29-39 (CAS). San Bernardino Co.: Cronise Valley, 1 d lV-29-56 (CIS); 12 mi N Earp, 2 d \'-5-53 (WEF). NEVADA. Nye Co.: Mercury, code MMT, 1 d VlI-10-64, no. 1183, 1 d VIII-21-64, at light, no. 1249. Distribution. Although large collections of nocturnal Mutillidae have been made in the Great Basin Desert, macsicaini has not been found there. Collection records cited above sug- gest that macswaini is a hot-desert species, and therefore might not occur outside of the Sonoran and Mojave Deserts. At the test site it occurs as high as 4200 ft. Sphaeropthalma (Micromutilla) pallida (Blake) Agama pallida Blake, 1871. Trans. Amer. Ento- mol. Soc, 3:263. Sphaeropthalma (Micromutilla) arizonae Schu- ster, 1958. Entomol. Amer.. 37(n.s.):16. New synonymy. Type data. Agama pallida, holotype cf Texas (ANSP type 4552). Sphaeropthalma art- Brigham Young University Science Bulletin zonae, holotype cT Tucson, Arizona, June 5, 1935, Bryant (UM). Discussion. Seven specimens were taken at the test site in August between approximately 4000 and 5500 ft. Although a large number of mutillids in the small end of the size range were obtained during the limited light collecting at the test site, this species apparently does not occur at lower elevations there, and is not very abundant. The new synonymy cited above is based on examination of the type specimens and a series of specimens in the University of Minnesota collection. The genitalia of the holotypes are identical as described above in the discussion of acontia, but the differences described by Schuster (1958:16) in the key which validated the name arizonae simply are expressions of the range of variation that occurs in this species. The holotype arizonae is only 3.5 mm in length. With size reduction, sculpture is reduced, and the length of the first flagellar segment becomes slightly reduced in relation to the length of the second segment and pedicel. Coloration also is variable. In the lower elevations ( below approxi- mately 4000 ft) the body usually is uniformly testaceous with slightly lighter legs and anten- nae. However, there is a tendency toward dark- ening of the second abdominal segment, and this can be detected even at the lower elevations. At 5400 ft in the Chiricahua Mts. of Arizona, the head, thorax, and petiole are ferruginous and the gaster castaneous, but the legs are light- er than the thorax. At higher elevations (5900 ft in the Guadalupe Mts. of New Mexico), the legs and antennae are darkened also, and the humer- al angles of the pronotum are darkened in some specimens. Farther east, in Nolan County on the Texas plains at 2000 ft, pallida has the colora- tion of test site specimens. Integumental darken- ing with increasing altitude, as described here, is not restricted to pallida, but is found in most other species which occupy a wide altitudinal belt. Extensive collections will be required to determine more completely the range of pallida. Distribution. Mojave and Sonoran Deserts, east to Texas. Sphaeropthalma (Miciomutilla) parapenalis, new species Diagnosis. Male: Length 9 mm. Integument entirely ferruginous. Mandibles without poster- ior margin excised, without a tooth on the pos- terior, basal margin, more or less parallel-sided from base to base of apical teeth. Clypeal disc longitudinally concave, transversely straight, with a pair of blunt, short teeth apically, basally produced into a prominent tubercle. Mesostern- um moderately, shallowly punctate, without tubercles or processes. Coxae and trochanters un- armed. Sternal felt lines reduced to tufts ap- proximately 0.2x the length of tergal felt lines. Pygidium polished, impunctate. Description. Head: Integument with outline dorsad of eyes fonning an arc of a circle in frontal view; punctures moderate, confluent. Measurements in micrometer units: eye length 42, ocellocular distance 22, ocellar length 16. Interocellar area not darkened. Mandibles over- lap anteroposteriorly; posterior margin of apical one-half twisted ventrad, anterior margin bear- ing a distinct carina from base to tertiary apical tooth; measurements given in Fig. 7. Apical mandibular teeth with 1° tooth basal width Ix and length 3x the 3° tooth; 2° t(K)th basal width 0.75x and length 1.3x the 3° tooth. Clypeal sur- face polished, with sparse micropunctures bear- Fig. 7. Mandible of Sptiaeropthalrna (Micromutilla) parapenalis Ferguson. Numbers indicate transverse measurements. ing short or long hairs; apex not impressed be- low anterior margin t)f mandibles. Antenna with first flagellar segment 0.77x the length of sec- ond segment. Frontal line and preocellar pit absent. Thorax: Pronotum shallowly, coarsely, confluently punctate, except polished and shal- lowly rugose on posterolateral margin. Meso- notal punctures moderate, shallow, one half puncture widtli apart, with flat, polished inter- spaces. Parapsidal lines with posterior one-half impressed. Mesopleuron with anterior impressed area polished, with small, scattered punctures, and micropunctures bearing fine, simple liairs on the interspaces; posterior raised area reticu- late, with narrow, polished interspaces anterior- ly, impunctate, pohshed posteriorly, with fine, scattered punctures bearing short, simple hairs. Propodeum c-oarsely, deeply reticulate. Ptero- stignial cell castaneous, O.S2x length of mar- MUTILLIDS OF THE NEVADA TeST SiTE 15 ginal cell on costa. Wing distinctly infuscated apicad in a band posterior to stigma. Abdomen: Plumose fringes dense only on second tergiim apicallv. First tergum moderately, confluently punctate throughout, apically 0.49x greatest width of second tergum, first sternum arcuate in crt)ss section, anterior one-half without a medio- longitiidinal carina and distinct basal tooth. Sec- ond tergum polished, finely, sparsely punctate, with sparse, short, recumbent, and long, erect simple hairs, except more coarsely and closely punctate anterolaterally; subapical margin finely, confluently punctate, with short, simple hairs; felt lines 0.64x length of tergum laterally. Second sternum moderately, shallowly, subconfluently punc-tate, with long, erect, simple hairs, slightly tumid anteromedially. Genitalia: Parameres ar- cuate dorsad, laterally straight, glabrous. Cuspis 0.38x free length of paramere, rodlike, with ap- proximately 12 mesally-directed short hairs at- tached to the mesal surface. Female. Unknown. Discussion. In Schuster's paper (1958:14), this species was misidentified as Sphaeropthahna (Micromutilla) juxta (Blake) to which it keys. The name jiixta appears again, on page 32 in Schuster's key to the species of the subgenus rhotopsis, with which it should be associated. The name then should properly read Sphaerop- ihalma (Photopsis) juxta (new combination). Sphaeropthahna parapenalis is distinct from all other North American nocturnal mutillids in having the parapenal lobes of the parameres impressed, concave, membranous dorsally, and not closely clasping the penis valves laterally. This species usually can be recognized readily without the use of the genital characters by means of the following combination of charac- ters: mandibles with uninterrupted posterior marginal carina, wings with distinct infuscation extending from stigma to posterior margin and more or less distinctly to the apex of the wing. Furthermore, the pubescence of the last two segments of the abdomen is distinctly infuscated, whereas the remaining pubescence is hyaline, and the petiole is more coarsely sculptured than in most species. As is usual in the nocturnal Mutillidae, the smallest specimens have the in- fuscation and coarseness of sculpture consider- ably reduced. The distinctness of this species warrants placing it in a group by itself, as Schuster has done. However, since the name juxta belongs to another species in another sub- genus, the group name parapenalis should now be used for the species parapenalis. Type data. Holotype d (CAS): Mt. Diablo, Contra Costa Co., California; VII-11-54; at light, W. E, Ferguson. Paratypes: ARIZONA. Cochise Co.: Chiricahua Mts., 1 d" VI-27-49 (OSU); Chiricahua Mts., Stewart For. Camp, Cave Creek Canyon, 3 d IX-(13-14)-52 (CAS); Chiricahua Mts., S. VV. Research Sta., 2 d IX-I- 58, I d IX-6-58 (CIS); Chiricahua Mts., Pinery Cn., I d VII-30-57 (UA); Chiricahua Mts., 5 mi W Portal, 1 d VIII-5-58 (UCD); 1 d IX-9-59, 1 d VII-2-61, 1 d VII-7-61 (WEF); 3 d VIII-28-57 (UI); 1 d VI-13-58 (WEF); Chiri- cahua Mts., 3.5 mi W Portal, 3 d VIII-13-52 (CAS); Chiricahua Mts., 15 mi W Portal, 1 d VIII-4-58 (UCD); Portal, 1 d VI-4-59; 1 d VI- 23-59 (UC;D); 9 mi W Portal, 3 d VII-1-64 (WEF); Huachuca Mts., 1 d VII-36 (CAS); Huachuca Mts., Carr Canyon, 1 d VIII-29-52 (CAS); 8 mi \ Bisbe^-, 1 d VIII-lI-52 (CAS); 9 mi S McNeal, 1 d VIII-30-58 (CIS). Coconino Co.: Oak Creek Canyon, Midgley Bridge, 2 d VIII-25-52 (CAS). Graham Co.: Graham Mts., Noon Creek, 3 d Vni-8-55 (UA, WEF); Thatch- er, 2 d VI-18-51, 1 d VI-20-50, 1 d VI-24-50, 1 d VI-26-51 (UCD). Pima Co.: Catalina Mts., mile 6, 2 d" VII-14-55 (WEF); Santa Catahna Mts., Mt. Lemmon Lodge, 2 d VII-24-52 (UCD). Santa Cruz Co.: Santa Rita Mts., Ma- dera Canyon, 1 d Vni-15-40 (CAS); 1 d VII- 10-57 (UCD), 1 d VII-I4-59 (UA). Yuma Co.: San Luis, 1 d VIII-11-40 (CAS). CALIFORNL\. Alameda Co.: Berkeley, 1 d VII-14-31 (UCD). Butte Co.: Oroville, 1 d \TI-13-26 (CDA). Cala- veras Co.: San Andreas, 1 d VI-16-34 (CAS). Colusa Co.: Colusa, 1 d VI-29-59, 1 d VII-1-59, 1 d VII-15-59 (UCD); College City, 1 d \TI-9-59 (UCD). Contra Costa Co.: 5 mi W Brent^v(Kxi, 2 d IX-5-58 (WEF); Danville, 1 d VII-8-49, 2 d XII-17-50, 1 d VIII-27-50 (CAS); Martinez, 1 d IV-8-11 (CAS); Mt. Diablo, 1 d VI-11-60, 4 d VII-11-54, 1 d VII-11-56, 2 d VII-19-61, 1 d VII-29-56, 3 d VIII-16-58, 1 d IX-22-56, 4 d IX-24-60, 1 d IX-29-58 (WEF); Pleasant Hill, 1 d VII-19-61, 1 d VII-28-58, 1 d VII-18-52 (WEF); Somersville, 2 d VI-27-56 (WEF); Walnut Creek, I d VlII-6-61 (WEF). Fresno Co.: Pine Flat Dam, 1 d VIII-5-52 (WEF). Kern Co.: WtK)dy, 3 d \'II-30-59 (UCD). Kings Co.: Lemoore Id VIII-11-59 (LACM). Lake Co.: Lucerne, 1 d VII-7-51 (CAS); Soda Bay, 3 d VII-25-58, 1 d VII-17-59 (UCD); Lower Lake, 1 d VII-3-59 (UCD). Los Angeles Co.: Pasadena, 1 d VlII-4-15 (UCR). Mendocino Co.: Ukiah, 1 d XII-22-59 (CDA); Rancheria Creek at Yale Creek, 5 d \TI-25-54 (CAS). Merced Co.: Dos Palos, 1 d VI-26-57 (WEF). Mono Co.: Cole- 16 Brigham Young University Science Bulletin viUe, 1 d" VII-2-48, 1 d VI-29-48, 1 d VII-10-48, 1 d VII-12-48, 2 d VII-16-48, 2 d VII-17-48, 1 d VII-19-48, 1 d VII-28-48, 1 d VII-31-48 (WEF). Monterey Co.: Jamesburg, 1 d VII-9-58 (UCD). Napa Co.: Mt. George, 1 d" no date (WEF). Sacramento Co.: Rio Linda, 4 d VI- 26-59, 1 d VIII-23-57, 3 d VII-15-55, 2 d VIII-9-55, 4 d VI-29-56, 1 d VII-6-56, 1 d IX- 7-56, 2 d VI-28-57, 1 d VII-5-57, 7 d VII-11- 57, 8 c^ VII-19-57, 1 d VII-26-57, 2 d VIII-7-57 (UCD); Carmichael, 1 d VI-23- 31, 1 d VI-27-55, 1 d VIII-1-55 (UCD); Fair Oaks, 1 d VI-25-36 (UCD). San Mateo Co.: Menlo Park, 1 d 1-05 (CAS). Santa Clara Co.: San Antonio Val, 2 d VIII-17-49, 1 d VII-30- 49 (CIS); New Almadeh, 1 d IX-10-64, 1 d VIII-5-64 (WEF). Shasta Co.: Redding, 1 d 1958 (UCD). Solano Co.: Vacaville, 1 d VII- 28-48, 1 d VIII-30-49, 2 d IX-19-52, 1 d VIII- 15-53, 1 d VI-9-54, 1 d VII-16-54, 1 d VI-29- 56 (UCD). Sonoma Co.: Preston, 2 d VII-16-I7 (CAS). Tehema Co.: Dairwille, 1 d VII-9-56 (UCD); Red Bhiff, 3 d Vn-9-56, 2 d VII-20- 56, 1 d VIII-16-56, 2 d IX-7-56, 1 d IX-21-56 (UCD); Los Molinos, 2 d VII-20-56, 1 d VIII- 23-56 (UCD). Tulare Co.: Pixley, 1 d VIII-7-59 (LACM). Yolo Co.: Davis, 1 d VI-17-59, 1 d VI- 22-59, 1 d VII-8-59, 1 d* VIII-19-50, 1 d VI-22- 54, 1 d VII-17-59, 2 cT VIII-4-55, 2 d VIII-10- 55, 1 d VIII-5-58, 1 d VIII-12-53, 1 d IX-23-46 (UCD); 4 mi SW Dunnigan, 1 d IV-25-59 (UCD); 4 mi SE Dunnigan, 2 d VII-1-59, 1 d VII-8-59, 2 d VII-28-59 (UCD); 7 mi NW Dunnigan, 3 d VII-1-59, 2 d VII-6-59, 1 d VII-14-59, 4 d VII- 12-59, 1 d VII-16-59, 1 d \'II-21-59, 1 d VII- 28-59, 1 d IX-2-59, 1 d IX-10-59 (UCD); 2 mi SE Dunnigan, 1 d VI-17-59 (I'CD); 8 mi i\W Win- ters, 4 d VII-13-59, 1 d VII-28-59, 1 d VIII- 5-59, 2 d VIII-28-59, 1 d IX-2-59 (UCD); 9 mi W Zamora, 1 d \'II-16-59, 1 d VIII-28-59, 1 d VII-1-59 (UCD); Rumsey, 3 d VII-23-55, 1 d VIII-5-55, 1 d VIII-11-55 (UCD); Esparto, 1 d VII-3-56, 1 d VIII-13-59 (UCD). IDAHO. Ada Co.: Boise, 2 d VII-26-53 (CAS). Cassia Co.: 5 mi NE Malta, 2 d VII-16- 52 (UI). OREGON. Umatilla Co.: Umatilla, 6 d VII-10-58 (WEF); Cold Springs Junction, 4 d VIII-9-60 (WEF). NEVADA. Eureka Co.: 27.5 mi W Carlin, 1 d VII-10-59 (CAS); Nye Co.: 35 mi NNW Mercury, Nevada Test Site, 1 d VIII-11-64 BYU- AEC code 12M(T), no. 1201; 1 d VIII-11-64 BYU-AEC code 12CC(T), no. 1206; 1 d VIII- 5-65 BYU-AEC code ECH(TB), no 1085. TEXAS. Nolan Co.: Sweetwater, Te.xas Exp. Sta., 1 d VIII-3-37 (TAM); 1 d VII-28-37 (WEF). UTAH. Juab Co.: Ibapah, Callao Pass, 1 d VIII-3-53 (UU). Millard Co.: Delta, 1 d VIII- 3-49 (USU). Weber Co.: Ogden, 1 d VII-8-59 (CIS). WASHINGTON. Asotin Co.: 2 mi S Aso- tin on river bank, 2 d VII-23-63 (DAG); 1 d VII-23-63 (WEF); 1 d \'II-23-62 (JZW). MEXI- CO. Chihuahua: Catarinas, 1 d VI 1-25-47 (AM \H). Coahuila: Cabos, 1 d VIII-21-47 (AMNH). Durango: Nombre de Dios, 1 d VIII-6-51 (CIS). Zacatecas: 15 km E Sombrerete, 1 d VIII-28-51 (CIS). Distribution. This is one of the most dis- tinct nocturnal mutillids of North America, mor- phologically as well as distributionally. In Cali- fornia it occurs through the coastal valleys, the Central Valley, and the coastal as well as Sierran foothills. Farther inland it is found throughout the Great Basin Desert, but apparently only characteristically on the slopes of the ranges rather than in the basins. At the test site this is illustrated by only three specimens captured at three places along the ba.se of Rainier Mesa, 35 miles NNW of Mercury. The dominant vegeta- tion there is Coleogyne ramosissima, with sparse Junipcrus osteosperma. Even large collections at altitudes from 3800 to 4200 ft at the test site did not contain parapcnalis. In Arizona, para- pcnalis has been found onl\' within the lower edge of the tree belt which usually is associated with the mountains that rise rather abruptly from the desert. Its distribution extends farther east onto the Texas plains at altitudes of at least 2000 ft, and south into Mexico at altitudes up to at least 7500 ft. I have not seen any specimens from the Mojave or Sonoran Deserts. SpJ}aeropthahna (Micromutilla) sonora Schuster Spliaeropthalma (MicromutiUa) sonora Schuster, 1958. Entomol. Amer., 37(n.s.):16. Type data. Holotvpe d Tucson, Arizona (UM). Para types: 2 d Palm Springs, Riverside Co., California (LACM); 1 d Borego, San Diego Co., California (CIS); 2 d" Hopkins Well, Riverside Co., California; 1 d Coalinga, Fresno Co., California ( CIS ) ; 1 cf locality unknown to me (UM). Discussion. Ninety specimens were collected at the test site from ten different localities of the desert slopes and valleys below the lower limit of the Coleogyne and the Pinyon-Juniper communities, or approximately 4.500 ft. Although sonora is locally abundant in the Sonoran and VIojave Deserts, it occurs at least as far north as Fort Churchill and Pyramid MUTILLIDS OF THE NEVADA TeST SiTE 17 Lake, Nevada, where I have collected it com- monly at light. The existence of sonora at Yucca Flat thus fits this more extensive, though very imperfectly known, distribution pattern. Sphaeroptliabiw (MicioinutiUa) yumaella Schuster Sphaeropthahiw (Micromutilla) yumaella Schu- ster, 1958. Entomol. Amer., 37(n.s.):19. Type data. Holotype d Wellton, Yuma Co., Arizona. The specimen is the property of Cor- nell University, although it probably still is lo- cated in the Schuster collection. Discussion. SL\ of ten specimens were c-ol- lected at incandescent light at Mercury. The other four were taken at incandescent and ultra- violet lights, and from a can pit-trap in study area 5A on the west side of Frenchman Flat in July and August. Tliis species should be present in the Great Basin portion of the test site, since specimens in my collection indicate that the distribution includes the area from Pyramid Lake, Nevada, south into Baja California, Mexi- co, and east to Portal, Arizona. Representatives of yumaella usually are among the smallest nocturnal mutillids, and hence probably were not captured in numbers proportionate to their actual abundance at the test site. Sphaeropthalma (Photopsis) angulifera Schuster Sphaeropthalma (Photopsis) angulifera Schuster, 19.58. Entomol. Amer., .37(n.s. ) :32. Type data. Holotype d Bakersfield, Cali- fornia (CAS). Paratype d Aberdeen, Owen's Valley, Inyo Co., California (ANSP). Discussion. Seven specimens were collected at the test site from late June to early Septem- ber within the altitudinal range of 3900 ft near Cane Springs, to 5400 ft at the base of Rainier Mesa. It is surprising that angulifera males have not been collected at lower elevations at the test site, because I have taken them at lights in the Mojave Desert near Lancaster and Ivanpah, but never in large numbers. Sphaeropthalma (Photopsis) hlakeii (Fox) Photopsis Blakeii Fox, 1893. Proc. California Acad. Sci., (Ser. 2), 4:6. Mutilla ceyx Fox, 1899. Trans Amer. Entomol. Soc., 25:262. Type data. Photopsis Blakeii, lectotype cT San Jose del Cabo, Lower California, Mexico (CAS type no. 292). Mutilla ceyx, lectotype d" Calmali Mines, Lower California, Mexico (ANSP type no. 4653). Syntype d El Paraiso, Lower California, Mexico (ANSP type no. 5054). Discussion. Only three specimens were taken at light from Mercury, and from locahty 5A on the west side of Frenchman Flat. Sphaeropthal- ma blakeii occurs rather commonly in the lower altitude deserts south into Mexico, but I have not collected individuals of this species farther north in the Great Basin Desert. This suggests that blakeii is at the limit of its distribution in the Mojave Desert portion of the test site. The synonymy listed above was based on examination of all three of the cited type speci- mens, and comparisons of these with a long series of specimens from other localities (Fer- guson, 1962:10). Sphaeropthalma (Photopsis) ferruginea (Blake) Aganm ferruginea Blake, 1879. Trans. Amer. En- tomol. Soc, 7:254. Type data. Holotype d Nevada (ANSP type no. 5615). Discussion. Four of six specimens were taken at light in August between 5400 and 6100 ft from two localities at the foot of Rainier Mesa on the test site in typical Great Basin Desert vegetation. At the lower site, Coleogyne ramosis- sima was predominant with scattered juniperus osteosperma. A fifth specimen was taken at light in August, approximately 2.3 mi NE of Mercury and 0.2 mi E of Mercury Highway, at 4200 ft. The sixth specimen was taken at light in July from locality 18M. Specimens in my collection indicate that this species is characteristic of the Great Basin Des- ert. It also occurs westward to the Coast Ranges, and southward into the mountains of southern California where it occurs in the Pinyon-Juniper and the Oak-Grassland commu- nities. Sphaeropthalma (Photopsis) helicaon (Fox) Mutilla helicaon Fox, 1899. Trans. Amer. Ento- mol. Soc, 25:254. Photopsis lingulatus Viereck, 1903. Proc. Acad. Nat. Sci. Phila., .54:737. New synonymy. 18 Brigham Young Univebsity Science Bulletin Sphaeropthalnui (Photopsis) carinata Schuster, 1958. Entomol. Amer., 37(n.s.):34. New synonymy. Sphaeropthalnui (Photopsis) heUcaon coahuilae Schuster, 1958. Entomol. Amer., 37(n.s. ):34. New synonymy. Sphaeropthalma (Photopsis) heUcaon diegueno Schuster, 1958. Entomol. Amer., 37(n.s.) :3.5. New synonymy. Type data. Mutilla heUcaon, holotype d" Nevada (ANSP type no. 4642). Photopsis litigu- latus, holotype cf La Jolla, San Diego Co., Cali- fornia (ANSP, type not numbered). Sphaerop- thalma carinata, holotype cT Purissima, Baja California, Mexico (property of USNM, unique type not yet deposited by Schuster). Sphaero- pthalma coahuilae, type material not designated, but presumably from the state of Coahuila, Mexico. Sphaeropthalma helicaon diegueno, un- til June 28, 1959, was represented by a male specimen on loan to Schuster from Cornell Uni- versity and bearing the following label data: Porter 20; Cornell U., Lot 709, Sub.; Arizona; Specimen B 12 F; Holotype, Sphaeropthalma helicaon ssp. diegueno Schu. At that time Schu- ster discovered that this specimen had deeply excised mandibles with a large basal tooth, and therefore did not represent a variation of helicaon, nor could it be keyed to helicaon diegueno in his key which validated the latter name. Schuster then selected a new holotype male which bears the following label data: S. Carlos, Ariz., 12, 13 May 18, J. Ch. Bradley; Holotype, Sphaeropthalma helicaon ssp. die- gueno Schu. This specimen also belongs to the Cornell University collection, but has not yet been deposited there by Schuster. Discussion. One specimen was taken at light in June at Mercury. Label data from other specimens in my collection suggest that this species is characteristic of the Mojave and Sonoran Deserts, but not the Great Basin Desert. The rarity of helicaon at the test site apparently is indicative of the marginal nature of the environment there tor this species. The new synonymy listed above is based on a preliminary study of this species as a part of a revision of the subgenus Photopsis. I have examined all holotype specimens except that of coahuilae, which Schuster could not find as of July, 1959. However, specimens from Saltillo, Coahuila, Mexico, are on loan to me from the American Museum of Natural History, and ap- pear to be this dark-headed form. Sphaeropthal- nui carinata is simply an individual which has a scutellar carina formed from the united mar- gins of the punctures on each side of the mid- line. The expression of punctures and their separating ridges is so variable in a series of a dozen specimens from any given locality, that the variation in the sf>ecies must exceed even that found in carinata. The genitalia probably contain the best taxonomic characters that indi- cate the unity of this species, and clearly distin- guish it from all other known species of noc- turnal North American Mutillidae. The para- meres are arcuate dorsad, not arcuate mesad, acuminate, glabrous on apical one-half, basal one-half of mesal surface with dense, mesally- directed hairs; cuspis one-half the length of the parameres, rodlike, slightly arcuate, with sparse, long hairs attached to the ventral surface and directed ventromesad. When the genitalia have been extruded from the body, the dense, basal hairs of the parameres, described above, provide instant recognition of the species. In discussing Schuster's subspecies concepts with him in 19.59, he indicated that he studied only one or two specimens which represented each of the names that he used for the taxa listed in synonymy above. Certainly this is not an adequate basis for recognition of subspecies and closely related species in the nocturnal sphaeropthalmine Mutillidae. Schuster ( 1958: 43) stated of the group alhicincta, in which he placed helicaon, "This is one of the very diffi- cult and polymorphic complexes; the following key is frankly tentative." I consider this to be a good indication that the characters of colora- tion, eye and ocellar size, puncturation, clypeal convexity, and petiole proportions are so variable and difficult to describe precisely that identifi- cation according to Schuster's key is very diffi- cult. Considerable variation is to be expected in a widespread species, but Schuster did not pre- sent distributional evidence in support of his taxonomic concepts. I believe it is better to con- sider that all of the above names are synony- mous until enough specimens and distributional data have been massed to make intelligent de- cisions about possible resurrection of names. Sphaeropthalma (Photopsis) unicolor ( Cresson ) Mutilla unicolor Cresson, 1865. Proc. Entomol. Soc. Phila., 4:389. Agama mendica Blake, 1871. Trans. Amer. Ento- mol. Soc., 3:259. New synonymy. Mutilla auraria Blake, 1879. Trans. Amer. Ento- mol. Soc, 7:248. New synonymy. MUTILLIDS OF THE NEVADA TeST SiTE 19 Mutilla Aspasia Blake [not SpliaeropJitlialmu (sic) aspasia Cameron 1895, Biol. Centrali- Amer., Insecta, Hyinenoptera, 2:370.], 1879. Trans. Amer. Entomol. Soc, 7:2.50. New synonym\'. Mutilla Phaedra Blake, 1879. Trans. Amer. En- tomol. Soc., 7:251. New synonymy. Agania rustica Blake [not Sphacrophthahna (sic) rustica Cameron, 1895, Biol. Centrali- Amer., Insecta, Hymenoptera, 2:342.], 1879. Trans. Amer. Entomol. Soc, 7:252. Schuster, 1958, Entomol. Amer., 37(n.s.):32. New synonymy. Plwtopsis nehulosus Blake, 1886. Trans. Amer. Entomol. Soc, 13:275. New synonymy. Sphaerophthalma anthophora (sic) Ashmead, 1897. Proc Southern California Acad. Sci., 1(3): 5. New synonymy. Mutilla monochroa Dalla Torre, 1897. Catalogus Hymenopterorum, 8:63. New name for Mu- tilla unicolor Cresson 1865, not Mi/rmosa uni- color Say, 1824, in Keating, Narr. Long's Second Expedition, 2:331. Dasymutilla sumneriella Cockerell, 1915. Ento- mol., 48:249. New synonymy. Sphaeropthalma (Photopsis) rustica ocellaria Schuster, 1958. Entomol. Amer., 37(n.s.):32. New synonymy. Type data. Mutilla unicolor, lectotyj>e d California (ANSP type no. 1887). Agama men- dica, holotype (?) d' Nevada (ANSP type no. 4.551). Mutilla uura/ia, holotype (?) 9 Nevada (ANSP type no. 4573). Mutilla Aspasia holo- type (?) 9 Nevada (ANSP type no, 4.574). Mutilla Phaedra, holotype (?) 9 Nevada (ANSP type no. 4575). Agama rustica, holotype (?) cT California (ANSP type no. 4550). Pho- topsis nehulosus, holotype (?) d" Nevada (ANSP type no. 4549). Sphaerophthalma antho- phora, holotype cT, allotype 9 Los Angeles, California ( USNM types no. 6113). Dasymutilla sumneriella, holotype 9 La Jolla, California (USNVI type no. 20409). Sphaeropthalma rus- tica ocellaria, holotype d" Berkeley, California (UM). Characteristics of the named forms. Mutilla unicolor is typical of males with the integu- ment entirely light ferruginous; pubescence aureous dorsally and ventrally on the abdomen; plumose fringes prominent only on segment two; eyes, ocelli, and wings normal. Females of this form have been identified as aspasia or phaedra. Photopsis nebtdosus is the form with the integument light ferruginous, except the femora slightly infuscated; pubescence hyaline except slightly aureous dorsally; eyes and ocelli nor- mal. This was synonymized with auraria females on the basis of rearing the two sexes from cells of Diadasia (Linsley and MacSwain, 1952; Fer- guson, 1962). Agama rustica represents males with the in- tegument ferruginous, except the thorax lateral- ly; antennae, legs, and petiole castaneous to black; pubescence aureous; eyes, ocelli, and wings normal. Females usually were identified as auraria. Schuster (1958) used the name in- correctly for males with the coloration of rustica, but which have the abnormal condition of brachyptery and small eyes and ocelli. The type specimen is normal in these respects. Sphaeropthalma rustica ocellaria (Schuster, 1958:32) is the name proposed in error for nor- mal specimens like the type of rustica. Agama mendica has the integument entirely castaneous to black, and pubescence entirely hyaline. Sphaerophthalma anthophora males have the integument ferruginous, except the flagellum, legs, and thorax laterally black, with pubescence reddish aureous. Females have the integument translucent ferruginous, except the thorax later- ally and the legs black. Pubescence is reddish. Dasymutilla sumneriella is the same as antho- phora females, and was synonymized by Krom- bein (1951). Discussion. The long synonymy above is the result of study of well over one thousand speci- mens of this species, including all of the type specimens. Sphaeropthalma unicolor occurs from sea level on the Pacific Coast, eastward into the Mojave and Great Basin Deserts to Utah and New Mexico. In the north-south direction, it ex- tends from Washington and Idaho into the coastal areas and middle altitudes of the north- ern mountains of Baja California. I have not seen specimens from the Sonoran Desert. The synonymy well represents the variability in this widespread species which occupies so many ecologically different yet intergrading environ- ments. At the test site, unicolor apparently is one of the most abundant nocturnal Mutillidae, rank- ing equally with Odontophotopsis microdonta Ferguson. It occurs from the valley floors to the Pinyon-Juniper and Artemisia communities at well over 7000 ft. At approximately the lower 20 Brigham Young Univehsity Science Bulletin limit of the junipers the ferruginous-colored val- ley form is replaced by the melanistic one, which is found only at the higher altitudes, above .5.500 ft. Since the two forms apparently are not sym- patric any place within their distribution, yet ap- pear to have continuous distributions, it appears to me that one fonn may be gradually replacing the other. The more favorable conditions of the desert valleys and the Central Valley of Cali- fornia, as well as the Columbia and Snake River Valleys, certainly must permit more generations per year. Hence, if the ancestral form of unicolor were melanistic like the Great Basin form, and mutations for lighter coloration occurred in the lower altitude populations, the more rapid genetic turn-over there would have permitted rapid spread of lighter coloration in the areas of favorable climate. I have traced the limits of the two major color forms from southern Idaho, along the east flank of the Sierra Nevada, and along the basins and ranges where the Mojave and Great Basin Deserts interdigitate. These rec- ords show the same altitudinal relationship of the iwo male color forms, without the distribu- tional overlaps which are characteristic of dif- ferent species. Collections at the test site include 76 females which were found in can pit-traps, while only four males were found in the same traps. During a brief period of intensive light-trapping in August, 1964, .36 of the lighter-colored males and 13 of the melanistic males were collected. Sphaeropthalma (Pholopsioides) amphion (Fox) Mutilla amphion Fox, 1899. Trans. Entomol. Soc. Amer., 25:263. Photopsis abstrusa Baker, 1905. Invert. Pacifica, 1:113. New synonymy. Photopsis nudatus Baker, 1905. Invert. Pacifica, 1:114. Type data. Mutilla amphion, holotype (?) c^ Nevada (ANSP type no. 46.54). Photopsis abstrusa, holotype (?) cT Clareniont, California (CU). Photopsis nudatus, holotype (?) cf Clare- mont, California (CU). Discussion. Ten males of amphion were col- lected at the test site. In all of the collections made from autumn, 19.59, to August 1, 1964, only one specimen was taken. This was found on the ridge which separates Frenchman and Yucca Flats, within a box-type live-catch trap used for mammals. Through intensive light-trapping in August, 1964, eight additional specimens were collected from the hillsides around Mercury, study area .5A on the west side of Frenchman Flat, and the area of Quercus gamhelii and Pinus monopIitjUa at the base of Rainier Mesa. One additional specimen was taken at ultraviolet light in the same general area on August 5, 1965. The new synonymy listed above is based upon study of all of the type specimens and a series of more than sixty others. Schuster ( 1958: 38) synonymized nudatus with amphion, but considered abstrusa as a subspecies of amphion. This is absurd because the type specimens of nudatus and abstrusa were collected at the same locality. Schuster separated the two subspecies on the basis of proportions of marginal cell length and stigma length, depression of the clypeus, and body size. These factors are vari- able even within given localities, and size is in- dej:>endent of the other two variables. Figure 8 shows the variability in the proportions of stigma length compared with marginal cell length on J^i- 1.5 1.4 _ 1.5 _ _ _[ajiipl Fig. 8. Sphaeropthalma (Pholopsioides) amphion (Fox) stigma length foinpared with marginal cell length ( unity ) . Cohinuis represent measurement.s of all individuals in population samples from: A, Nevada Test Site; B, New Almaden, Santa Clara Co., Calif.; C, Coleville, Mono Co., Calif. Arrows indicate limits of subspecies proposed by Schuster (1958). MuriLLlDS OF THE NEVADA TeST SlTE 21 the costa (expressed as unity) in a series of eight, ten, and twelve specimens, the total num- bers taken from three widely separated localities. I believe that this graphic evidence establishes the former arbitrary and invalid division of this species into two subspecies. Odontophotopsis (Odontophotoj)sis) (irmata Schuster Odontophotopsis (Odontophotopsis) armata Schuster, 1958. Entomol. Amer., 37(n.s.):60. Discussion. Fifty-two specimens were col- lected at incandescent light on the hillsides near Mercury and at Cane Springs. Apparently this is a Mojave Desert species and occurs on the val- ley slopes, but not in the bottoms of the valleys or as high as the Pinyon-Juniper community. This is the first locality recorded for this species since Schuster validated the name in a key, without designating a type specimen or type locality. The identification of this species, there- fore, is based entirely upon Schuster's key. Odontophotopsis (Odontophotopsis) chjpeata Schuster Odontophotopsis (Odontophotopsis) chjpeatus Schuster, 19.58. Entomol. Amer., 37(n.s.):59. Type data. Holotype d" Tucson, Arizona (UM). Paratype d Arizona (UM). Discussion. Of fourteen specimens captured at the test site, seven were found in can pit- traps in study areas 5A and 5E on the west side of Frenchman Flat in July, August, and Septem- ber, 1961. In the same general area, three speci- mens were collected at ultraviolet light and two at a gasoline lanteni during August, 1964. Al- though much light-trapping was done at Mer- cury and the nearby hillside, only one specimen was taken there. Only one specimen (ref. no. 73) is from Yucca Flat, where it came to light. Specimens from can pit-traps were preserved in alcohol before being pinned; therefore, the lighter parts of the integument are darkened more than in specimens which were pinned while fresh and allowed to dry. Little can be said about the distribution of chjpeata at the test site except that it occupies the Mojave Desert portion, although one speci- men was taken in Yucca Flat. It seems to be much less abundant than O. microdonta Fer- guson, which is its closest relative (see discus- sion of microdonta). Additional material was examined from the following localities: St. George, Washington Co., Utah; 8 mi NE Mesa, Maricopa Co., Arizona; Organ Pipe National Monument, Pima Co., Arizona; and 6 mi S Palm Springs, Riverside Co., California. Odontophotopsis (Odontophotopsis) cookii Baker Odontophotopsis cookii Baker, 1905. Invert. Pa- cifica, 1:93. Type data. Holotype cT Claremont, Cali- fornia (CU). Discussion. Twelve specimens were collected from the test site. One of these came to light near Cane Springs ( 4000 ft ) , but all others were collected at incandescent or ultraviolet light along the base of Rainier Mesa. Those three areas contained Coleogyne ramosissima; Juniperus osteosperma and Coh'ogync ramosissima; and Quercus gamhelii and Finns monophyUa. Identi- fication of this species was based on examina- tion of the type specimen and comparison of the test site specimens with a large series from California. The test site sjjecimens are different from all others in having the mesosternal tubercles reduced considerably, so that in some specimens they are inconspicuous. Otherwise, the key characteristics, form of the mandibles, genitalia, and sculpture apj>ear to be identical in specimens from the test site and from Cali- fornia. When collections in intermediate locali- ties can be made, it should be possible to deter- mine whether or not the test site specimens should be given a different name. Odontophotopsis (Odontophotopsis) inconspicua inconspicua ( Blake ) Photopsis inconspicinis Blake, 1886. Trans. Amer. Entomol. Soc, 13:272. Type data. Holotype (?) cf California (ANSP type no. 4610). Discussion. Twenty-seven specimens were collected from the hills surrounding Mercury, the Atriplex area near the southwest edge of the playa in Frenchman Flat, Cane Springs, and in the lowest edge of the juniper belt at the base of Rainier Mesa. Although O. inconspicua incon- spicua occurs in both the VIojave and Great Ba- sin Desert portions of the test site, it apparent- ly does not occur as high as the Artemisia tri- dentata association on the top of Rainier Mesa. This species occurs commonly in the Central Valley of California, and south into tlie Colorado 22 Brigham Young University Science Bulletin Desert of California, at least as far as Indio in Riverside County. Odontopliotopsis (Odontophotopsis) microdonta, new species Diagnosis. Male: Length 9 mm. Integument of head, thorax, and petiole ferruginous; gaster castaneous; legs and antennae testaceous. Man- dibles with posterior margin deeply excised, with subtending tooth large, protuberant, rounded. Clypeal disc longitudinally concave, transverse- ly straight, with a pair of blunt, apical teeth, basally not protuberant or carinate. Mesostemal tubercles scarcely differentiated from puncture margins. Coxae and trochanters unarmed. Stern- al felt lines approximately one-third length of those on second tergum. Pygidium polished, im- punctate, apical fringe absent. Description. Head: Integument polished, with outline dorsad of eyes forming an arc of a circle in frontal view; punctures fine (diameter 2 units), sparse. Measurements: eye length 47 units, cx-ellocular distance 16 units, ocellar length 16 units. Interocellar area darkened. Man- dibles overlap dorsoventrally; anterior margin of distal one-half slightly twisted dorsad, and bearing a distinct carina from base to tertiary apical tooth. Apical mandibular teeth with 1° tooth basal width Ix and length 2x the 3° tooth; 2° tooth basal width Ix and length Ix the 3" tooth. Clypeal surface polished, with sparse micropunctures bearing long or short, simple hairs; apex not impressed below anterior margin of mandibles. Antenna with first flagellar seg- ment 0.77x the length of second segment. Front- al line and preocellar pit distinct. Thorax: Pro- notum dorsally and laterally coarsely, shallowly punctate except rugose on posterolateral mar- gin. Mesonotum moderately, shallowly punctate (puncture widths 4 units) with flattened, narrow interspaces polished. Parapsidal lines entirely impressed. Mesopleuron with anterior impressed area moderately to finely punctate, sparsely plumose-hairy; posterior raised area mtxlerately punctate to reticulate. Propodeum coarsely reticulate. Pterostigmal cell testaceous, 1.05x length of marginal cell on costa. Abdomen: Plumose fringes dense on segments two and tliree apically. First tergum moderately, closely punctate, apically 0.5x greatest width of second tergum. First sternum arcuate in cross section, with a mediolongitudinal carina on the anterior two-thirds, terminating anteriorly in a prominent blunt tooth. Second tergum [wlished, finely, sparsely punctate, the punctures two to three diameters apart, except anterolaterally more closely, coarsely punctate, with brachy- plumose, recumbent hairs and erect simple hairs; felt lines one-half length of tergum lateral- ly. Second sternum mcxlerately, shallowly, con- tiguously punctate, with brachy|olumose, recum- bent hairs and erect simple hairs; anteromedially tumid. Genitalia: Parameres arcuate dorsad, lat- erally straight, glabrous. Cuspis length one-half free length of paramere, rodlike, flattened, with approximately twenty mesally-directed short hairs on mesal surface. Female. Unknown. In spite of can pit-trap and light-trap records, it has not been possible to correlate the sexes of this species. Discussion. Odontophotopsis microdonta is most closely related to O. chjpeata Schuster ex- cept for the reduction of the mesostemal tu- bercles, but should key out to the latter species. The holotype was selected because its tubercles are well developed for this species. Even when the tubercles are absent, microdonta can be recognized as belonging to Odontopliotopsis be- cause of the typical genitalia (described above), with the cuspis about one-half the free length of the parameres, and the abundance of brachy- plumose body hairs. The mesostemal tubercles, which are key generic characteristics, normally are so small that they can be easily overlooked. Hence, with the normal condition minimal, slight variation results in absence or in a relative prominence of the tubercles. Since this species verv closely resembles clypcata, specimens with relatively distinct mesostemal tubercles may be mistaken for that species. However, the tubercles of chjpeata are larger and are in a different place. In microdonta the tubercles are approxi- mately two puncture widths anterior to the mesocoxae, but in chjpeata they are approxi- mately two puncture widths farther anterior. Punctures used as the basis for measurement lie lateral to the tubercles. An additional complica- tion in identification involves variation in pig- mentation of the gaster. Dark specimens are not a problem, but some specimens have the dark integumental pigment reduced to the vicinity of the felt lines, and occasionally specimens lack dark pigment entirely. In this case identification may be difficult, depending on the prominence of the mesostemal tubercles. Only a few speci- mens of microdonta have been observed with some pygidial fringe hairs, whereas specimens of chjpeata characteristically have the fringe well- developed. In chjpeata this character is variable, probably owing to abrasion and heredity, but MuriLUDs OF THE Nevada Test Site 23 there is nearly always evidence of at least a few fringe hairs. Odontoplwtopsis microdonta differs further from chjpcata by the slightly twisted mandibles mentioned in the description. Tlie dif- ference between twisted and untwisted is very subtle in this case, and difficult to describe and ascertain in individuals, flowever, in series of the two species separated on the basis of other characteristics, the distinction is more apparent. Furthermore, clypeata tends to have the medial base of the clypeus slightly more produced at its juncture with the frons than in microdonta. Type data. Holotyix; cf (USNM): 5 mi NNW Mercury, Nye Co., Nevada (USGS Specter Range Quadrangle, 1 mi from Mercury High- way, at .3400 ft), ultraviolet light, W. E. Fer- guson. Actual label data: Mercury, Nevada, N. T.S.; BYU-AEC Code 5M(TB); VIII-2.5-64; ref. no. 1298. Seven male paratopotypes same data. S