Pianka (1991)-Phynosoma platyrhinos.pdf

7/28/2019 Pianka (1991)-Phynosoma platyrhinos.pdf

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RE-: SQUAMATA: SAURIA: IGUANIDAE PHRYNOSOMA PLATYRHINOS

platyrhinos 8 platyrhinos Q@alidiarum 3 %alidiarum Q@oodei d' goodei Q

DeBnLtion. A medium to large horned lizard (adults 72-95rnrn SVL) characterized by a single row of relatively small lateralabdominal fringed e s ong the side of the body. Othercharacter-istics include: relatively short occipital horns; the longest temporalspine nearly a s long as the occipital horns; small throat scalessometimeswith a single row of slightly largerscaleson each ide ofthroat; enlarged chin shields; nostril inside a line connecting thesupraorbital ridge with the tip of the snout; snout very blunt. Thebelly is usually spotted. The dorsal color is variable,and canbe buff,yellowish, reddish, or grayish. Two prominent dark patches arepresent immediately behindtheneck, grading into a series of severalmore down the back and ontothe ail.

Catalogue ofAmerican Amphibians andRept i les .Pianka,E.R 1991. Pbynosomaphryrbinar.

Pbrynosoma platyrhinos GirardDesertHorned UzardPbynosomaphryrbinosGinrd,nBairdandGinrd, 1852 (1854):69.Type-locality, "from [the vicinity of the] Great Salt Lake, [Salt

Lake County, Utahl". Syntypes, National Museum of NaturalHistory (USNM) 189 3 specimens,all adults, one male and twounsexed) andMuseumof Comparative Zoology, Harvard(MCZ)5948, subadult female, collected "by Capt. Stansbury's party",date of collection unknown (not examined by author).Phrynosomaphryrbynas: Hallowell, 1853107.Doliasaumplatyrbinos: Ginrd, 1858:409.P h ~ m a p h r y r b i n u n r .ope, 1875:49.Ph ynosoma maccallii:Yarrow and Henshaw, 1878:1617 (part).Phrynosoma phryrbinus: Boulenger, 1885:247.Anotaphryrbimz Cope , 1900:443.Pbynosomaphthynbinar: Biihrne, 1988:38. Lupsus.

Content. Three subspecies are recognized: phtyhinos,

Diagnosis. P. DlatMbinoscanbe dis tiwished fromall con-

culidianrm,and goodei. See Remarks.

geners by the followi6: G o moderately elon Ge d occipital horns atthe back ofthe headusuallvnot n c o m aat their base: oosteriorhead

Map. The solid circle isthe estricted ype-locality forP.p.goodei; heremaining type-localiiies are too imprecise to plot. Open circlesrepresent other records; those outsidethe range boundary are extra-limitd (see text). Stvs indicate fossil localities.

margin betweenoccipital hornsnot indented; nostril & nin gs insidethe unthus rostralis;eardrumeither exposed or covered with scales;gular d e s mall, granular, uniform in size or with a single marginalrow of slightly enlargedscaleson each ide of the threat; side of bodybetween limbs with a single row of elongate, pointed fringe scales;smooth ventral scales; tail broadening gradually rather than abruptlyat base.Descriptions. Gimrd (1852), Cope (1900), Van Denburgh(1922), ~uesta-Terron(l932), mith(lW6), ~eeve(1952), herbrooke(1981),andMontulucu(1981)provided~ed~aiptiom.iihme(1988) provided a brief -1ption of the hemipenes. Go- et 11.(1969) described the karyotype (2N = 34).

Figure. Computer-aided camera luu& drawings of the heads ofrepresentativesof thethree ubspeciesofP.plaryrbinos: plutyrhinosn A c M 716% [male] and 716% [female]. 8.3 km north of Lovelock.PershingCounty,Nevada); calidimum(Lk~87&1 male]and6876[female]. 16.7km northwest of Casa Grande.P i oun~r. rizona):i o o d e i h c ~4302 M e ] nd 59948 [fekle l, 41.7km southeastofPueao Libextad and 6.7 km northwest of Puerto Penasco, respec-tively, Sonora, M6xico).

IUustntions. Behler and King (1979) and Pianka (1985)provided color photographs. Sherbrooke (1981) provided a series ofcolor photographs, includingrepresentative habitat anda copulatingpair. Switak (1979) provided a color photograph of a lizard and ablack andwhite photograph of representative hab it . Other blackand white photographs or line drawings are included in G i d(1852), Ditmars (1903, Bryant (19111, Van Denburgh (19221, Smith(1946), Stebbii(1954,1985),Collins(1988), andMontanucci(l989b).


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Montanucci (1989a) provided a black and white photograph of amandible, and Tanner and Avery (1982) provided several line draw-ings illustrating buccal anatomy. Wood (1917) illustrated thevascularized ocular fundus.Distribution. PbrynosomaplafyrSJinos s a widespread liz-ard in western North America, occurring in lowland deserts fromsoutheastern Oregon and southwestern Idaho through the GreatBasin of Nevada and western Utah into the Mojave, Colorado, andSonoran deserts of southeastern California, western Arizona in theUnited States and noaheastern Baja California and northwestern

Sonom in Mtzico. It can be found more frequently on sandy thanrocky substrates, and has been reported to elevations of 2000 m(Smith21946). Ste bb ii (19%) suggested hat several disjunct extra-l i d ocalities represented escaped pets .

Fossil Record Brattstrom (1954) reported fossil materialindistinguishable fromextant specimens, dated &10,000 ybp, fromGypsum Cave in the Frenchman Mountains near Lake Mead, ClarkCounty, Nevada. Other Pleistocene fossils have been reported fromSmith Creek Cave, White P i e County, Nevada (Brattstrom, 1976;Mead et al., 1982),and rom nearby CrystalBallCave, Millard County,Utah (Mead et al., 1989).Pertinent Utecature. The most comprehensive reatment isby Reeve (1952), who examinedvirtudly dl extant material and gavelocalities, range maps, descriptions, and keys. Other systematictreatments are provided by Presch (1969) and Montanucci (1987).

Jenkii and Tanner (1968), Presch (1969), and Montanucci (1987,1989a) reponed o n osteology; the latter along with Hotton (1955)examined anatomicalrelationships to diet. Other anatomical aspectshave beenreported by ~ever(19?3),~iUer(1981), a n n e rn d ~ v e r ~(1982). Schwenk (1985). and Schwenk and Throdunorton (1989).control of c& G g e s by temperature and light wasstudiedby Atsatt (1939), and Lowe (1947) noted the variability in and highdegreeof background color matching. Dumas 1 9 6 4 ) c o ~ e dnhome range, thermal requirements, and dop at ry with P. douglassi(these two species are sympatric near Grantsville, Utah; P i a ndParker, 1975). Themnoregulation has been studied by Cowles andBogert (1944), Branstrom(1965), Heath(1965), Pianka (1966), Porter(l%7), and Pianka and Parker (1975). Bennett and Licht (1972)studied anaerobic metabolism during activity.Behavior has been studied by Lynn (1965), Tollestrup (1981),andCollins (1988). Harris (1958) has suggested nocturnal habits, butboth Wiiiarns (1959) and Mays and Nickerson (1 968) argued againstthis nterpretation. Homingmovements were reported by Pianka andParker (1975). Reproductionhas been studied by Mayhew (1%8),Howard (1974), Vitt (1 977), andVittand Congdon (1978). Diets wereexamined by Knowlton and Janes (1932), Knowlton (1934), Fautin(1946, Pianka (1966), Pianka and Parker (1975), Rissing (1981), andWhitaker andMaser (1981). Pietruszka(l981) usedthis species in anevaluation of stomach flushing for dietary analyses.Pianka and Parker (1975) provided a comprehensive eview ofthe ecology of P. platyhinos. Important autecological studies in-dudedMedica etal. (1973) andT-r andKrogh(1973). Additionalecological aspects havebeen eported by Mosauer (1932), Duellman(1955), Gates (1953, AUred et al. (1%3), Tanner and Banta (1%6),Baur (1973), Busack and Bury (1974), Tomko (1975), Switak (1979),Rissing (1981), Werschkul(1982), Pietruszka (1986), Collins (1988),and Gonzllez-Romero and Alvarez-CSirdenas (1989). Pianka andPianka (1970) compared the species with its Australian ecologicalcounterpart, the agunid Mobcb bowidus. Baur (1986) reponed onlongevity.

Parasites were reponed by G v n b io and Heynernan (1%0),Babero and Kav (1967). Telford (1970). and Mankau and Widmer(1977). ~u t&(l971 ) provided an~;learophoretic datum, andYousef et al. (1977) reported bide acid composition. Alberts (1991)employed this species in a study of lizard femoral gland secretions.Nomenclatural History. Cope (1896, 1900) erected thegenusAnota,defied bytheconcealment of thetympanumbya scalyintegument, to include the three subspecies of platyrbinos, thenconsidered distinct species. The defining characteristic proved to bevariable in two of the three taxa and they were returned to the genusP b p o m a by subsequent workers. h t a 1971) argued p& ua-sively hat the original desaiption of P. phtynbinos by Charles Ginrdapp&ed in a sfiort paper the pr&ee&n8s of the Academy ofNatural Sciences in Philadelphia in 1852, prior to those contained in

the complete repom to Congress of the western railroad surveys aterthat same year.Remarks.Acursorvenminationof soecimens romthrouph-out the range of P.plaiyrb>ttarsuggests hatthe putative diagno&characteristics of the subsoecies are not reliable. The subsoeciesgrade more-or-less con&ously into one another. ~ e o g r a ~ G caceis currently best determined by the localiiy of collection.Etymology.Thenamephtynbinosis a combinationfromtheGreek phtqP(broad, or flat)and' nbinS(nose). Thenameculidimum

(L., 'Roman tnthhouse", fromthe words 'calid', meaning 'hot", and'ariumw,meaning'? placewhere something s keptm)probablyefersto the type-locality. The name goo& is a patronym honoring thenAssistant Secretary of the Smithsonian Institution, Dr. George BrownGoode.Comment. Various features of the anatomy, behavior, diet,temporal patterns of activity, thermoregulation, and reproduaivetactics of Pbrynaro map htyrbim a n e profaably interrelatedandinterpretedtoprovideanintegratedviewof itsecologyQ i i a , %6;Pianka andParker, 1975). These hornedlizardsueant specialists andusually eat nothing else.Ants are s d nd contain muchundigestablechitin, so hat large numbers of them must be consumed. Thus theselizards possess ai r~ po rt io na te l~arger stomach(about 13percent ofbody mass) thandoallothersympatricdexn lizardspecies, ndudingthe herb60rous Desert 1 g u G Dipsosaunrs dor&lis (herbivorestypically have lower digestive assimilation rates and larger stomachs

than do carnivores). Possession of such a large gut necessitates atanklike body form, reducing speed and decreasing a lizards abilityto escape from predators by flight. Natural selectionhas avored theevolutibn of a spiny archit&t


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~b& oso maphty rbinur : Stephens, 1921 62.Pb?ynosoma pkztirbinus: Cuesta-Tenon. 1932:116.~b&osoma>htyrbinospkz&rbims:~ L u b e r , 935179 (part).Pbrynosomupltyrbinac calidiancm.Reeve, 1952:856. First use ofcombination.Diagnosis. Occipital horns are heavy and long (45% or moreof head length); interoccipital space one-half the basal diameter ofoccipital spine; 5 or 6 temporal horns, each longer than the on eanterior; tail often somewhat flattened posteriorly.

3. Pbrynosoma phtyrblnos gooclel StejnegerPbrvnosoma bemandezi,. Streets, 1877336 (part).~b&osoma~oodeiStejneger,893:191. TG- loc ali ty, coast desertsof the state of Sonora, Mexico", resvia ed to Puerto Libenad bySmith and Taylor (195Va, b). ~olotype , ational Museum dfNatural History (USNM) 85671, age andsexunknown, collectedby T. H. Streets, M.D., date of collection unknown (not exam-ined by author).Anota good& Cope, 90042.Pbrynosomapkz~inacoodefi Klauber, 1935:179. F i t use of tri-nomial.Pbrynosomaphtyrbinargoodi: St eb bi i, 1954:314.

Diagnosis. Three temporal horns are enlarged, the oneposterior equal in size to occipital horns; the 3 posteriorchinshieldsare greatly enlarged and pointed.

LiteratureCitedAlberts, A.C. 1991. Phylogenetic and daptivevariation inluard fem-oral gland secretions. Copek 1991:69-79.Allred, D.M., D.E. Beck, nd C.D. Jorgensen. 1963. Biotic cornmuni-ties of theNevada Test Site. Brigham Young Univ. Sci. Bull.,Biol. Ser., 2


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Mayhew, W.W. 1968. Biology of desertamphibians andreptiles, p.195-356. InG.W. Brown (ad.), Desert biology. Academic Press,New York.Mays, C.E. andM.A. Nickerson. 1968. Nocturnal behavior in iguanidlizards and possible correlation with roadkills. Herpetologica24:258-259.Mead, J.I., T.H. Heaton, and E.M. Mead. 1989. Late Quaternaryrep-tiles from two caves in the eastcentral Great Basin.J. Herpetol.23:186-189.-R.S.Thompson, andT.R.VanDevender. 1982. Late W1x:onsinanand Holocene fauna from Smith Creek Canyon. Snake Range.- .Nevada. Trans. San Diego Soc. Nat. Hit. 20:l-26.Medica, P.A., F.B. Tumer, and D.D. Smith. 1973. Effects ofradiationon a fenced population of homed lizards (Pbymsoma phf y-nbinos) in southern Nevada.J. Herpetol. 7:79-85.Miller. M.R. 1981. electron microscope studies of the audi-torypapillae of somiiguanid lizards. G e r . . Anat. 162:55-72.Montanucci. R.R. 1987. A phylo~enetictudy of the homed lizards.genus Phrynosoma, b d - o n skeletal -and external morphol-ogy. Contrib. Scii, Nat. Hit. Mus. LosAngeles Co. (390):l-36.- 1989a. The relationship of morphology to diet in the homedlizard genus Pbrynosoma Herpetologica 45:208216.- 989b.Maintenanceandpropagationofhomedlizards Pbym-soma) in captivity. Bull. Chicago Herpetol.Soc. 24229-238.Mosauer, W. 1932. Adaptive convergence in the sand reptiles of theSaharaand California: a study ins trumre and behavior. Copeia1932:72-78.Pianka, E.R. 1966. Convexity,desenlizards, and spatial heterogene-ity. Ecology 47:1055-1059.- 985. Some intercontinentalcomparisonsof d e s e a l i . Natl.Geogr. Res. 1:490-504.- nd W.S. Parker. 1975. Ecology of homed lizards: a review withspecial eference to Pbrynosomaphty&inos. Copeia 1975141-162.-andH.D. P i a . 970.TheecologyofMobcb o n i d u c ( l a e :Agamidae) in western Au st nl i. Copeia 1970:[email protected], RD. 1981.An evaluation of stomach flushing for desertlizard diet analysis. Southwest. Nat. 26:lOl-105.- 1986.Searchtactics of desert lizards: how polarized are hey?Anim.Behav. 341742-1758.Porter, W.P. 1967. Solar radiationthroughthe living body walls of ver-tebrateswithemphasis ondesettreptiles. Ecol. Monogr. 37273-2%.Presch, W. 1969. Evolutionary osteology and relationships of thehorned lizard genus Pbrynaroma (family Iguanidae). Copeia1%9:25@275.

Reeve,W.L. 1952. Taxonomy and d isviut ion of the homed lizardsgenus P b w niv. Kansas Sci. Bull. 34817-960.Wing,S.W. 1981 Prey preferences inthe deserthornedlizard: influ-ence of prey foraging method and aggressivebehavior. Ecology62:1031-1040.Schwenk, K. 1985. Occurrence, W ~ b u t i o nnd functional signifi-canceoftaste buds inlizards. Copeia 198591-101.- nd G.S. Throdvnorton 1989. Functional and evolutionarymorphology of l i d eeding in squama.te reptiles: phyloge-net& and J. Zod. (London) 219:153-175.Sherbrooke,W.C. 1981. Horned lizards, unique reptiles of westernNonh America. Southwest Parks and Mon.Assoc.,Globe,Ari-zona.

-. 1985. A fieldguide to westernreptiles an da mp hi bi i. Znded.,rev. Houghton Mifflin Co., Boston.Stejneger, L. 1893. Annotated l i t of the reptiles and batrachians col-lected by the DeathValley expeditionin 1891,with descriptionsof new species. North American Fauna 7(Pt. II):159-228.Stephens,F. 1921.Anannotatedlistof theamphibiidreptiles ofSan Diego County, California. Tram. San Diego Soc.Nat. Hist.357-69.Streets,T.H. 1877. Contributions o the na tud history of the Hawai-ian and Fanning islands and lower California, made in connec-tion with the United States North Pacific surveying expedition,1873-75. Bull. U.S. Natl. Mus. 01 -172 .Switak, K.H. 1979. Leben in derWiiste, Krotenechsen der GarmngPbrynosoma Wiegmann, 1828. I. Teil: Beobachtungen in WierWildbahn. Das Aquar. 124:470475.Tanner, W.W. and D.F. Avery. 1982. Buccal floor of reptiles, a sum-mary. Great Basin Nat. 42:273-%9.-and B.H. Banta. 1966. A systematic review of the Great Basinreptiles in the collection of Brigham Young University and theUniversity of Utah.Great Basii Nat. 26:87-135.-and J.E. Krogh. 1973. Ecology of Pbrynaromaphtynbinaclt theNevada Test Site, Nye County, Nevada. Herpetologica 29:327-342.Telford, S.R., Jr. 1970.A comparative study of endoparasitisrnamongsome southern California lizard populations. Amer. Midl. Nat.85516-554.Tollestrup,K. 1981. The social behavior and displays of two speciesof homed lizards, Pbr yna rom aph tyn bi~ s nd Pbrynaromawmnahrm Herpetologica 37:13@141.

Tornko, D.S. 1975.The eptiles andamphibians of theGrandCanyon.Plateau 47161-166.Van Denburgh,J. 1922. The reptiles of western North-~a. Vol.I. Lizards. Occas. Pap. California Acad. Sci. (10):l-611.Vitt, L.J. 1977. observatiok on du tch andegg size and evidence formultiple clutches nsome lizards of southwestern United States.~e rpktolo~ica5333-338.- ndJ.D. Congdon. 1978.Body shape, reproductive effort,andrelative dutc hmass in l i d s : esolution of a paradox. hmer.Nat. 112:595-608.Werschkul, D.F. 1982. Species-habit relationships in an Oregoncold desert lizard community. Great BasiNat. 42:380-384.Wever, E.G. 1973. The function of the middle ear inlizards: divergenttypes. . Exp. Zool. 18497-126.Whitaker,J.O., Jr. andC.Maser.1981. Foodhabi of seven species oflizards fromMalheur County, southeastern Oregon. NoIthwest.Sci. 55:202-208.

W i i , .L. 1959. Nocturnal activity of some species of homedlizards, genus P b w erpetologica 1543.Wood, CA. 1917. The fundusoculiofbids.Lakeside Press, Chicago.Yarrow, H.C. andH.W. Henshaw. 1878. Report upon the reptiles andbatnchians collectedduring the years of 1875,1876, and 1877,in Cdidornia,Arizona, and Nevada, p. 1628-1648. In Ir.G.M.Wheeler, Geographical surveys of the territory of the UnitedStates west o fthemehundredthmeridian... Ann.Rept. Chiefof Engineus 1878, Pt.3, Appendix NN. U.S. 45th Congr., 3rdSess., House of Representatives Exec. Doc. 1.Yousef, I.M., W.G. Bradley,andM.K. Yousef. 1977. Bile acid compo-sition of some lizardsfrom southwestem United States. Proc.Soc. Em . Biol. Med. 15422-26.Smith, H.M. 1946. Handbook of lizards: lizvds of theUnired Statesand m d a . cornstock h b I . ~ o . ,~c., t h a , New York. & ~innlyepvtmentf ~ o g y ,niversity of e ex as ata n d . H.Taylor. 1950a. Type localities of Mexican reptiles and Austin, Austin, 78712-1061.amphibians. Univ. Kansas Scii.Bull. 33,h. I:313-380.a n d . 1950b.An annotated c hedrl it and key to the reptiles Primvy editor forthis account,mw. rice.of Mexico exclusive of the snakes. Bull. U.S. Natl. Mus. (199):~

+ 253 p. Published 15 October 1991andCopyrightQ 1991 by the Society forStebbins,RC.1954.Amphb'~andreptilesofwesternNoahAmerica. the Study ofAmphibiuls and Reptiles.McGraw-Hill Book Co., New York.

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Pianka (1991)-Phynosoma platyrhinos.pdf