EUAN wp Oe Nine 80am PAE AS . sereetetnsaatel srumtaen he ; + eee ese “ TN My yo Bal 172 Sonatenstedense net ate Fe Ml te ee ae Arak te al od Ah meine At ea oat ota an im OE gS Bai tg atrats ne ope en eee et cttialinf ele te Eanes seas - < a . Spe AL AURORE tia —_ on aro Mehy Smear 4 4 : ~ oa te tate te Sane tat an Manse armokafarty etin Sea e eth hare te . " Modem’ ee " oe Ty Re eae RENNER Ms tet Nerd eA oy Mark atta tvtiew MANA RC STatHe Mitel NAS ee eras “7 2 Ne ep teat ge eis fie eaten NiRhe ee oe Nasi Aee A wRN PPA NTN wie a eaten Cte erm att. ee ed we ete at Sener seals ed Pipt ih ve e HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology od Uf BREVIORA Museum of Comparative Zoology HARVARD UNIVERSITY "US. COMP, ZOOL LIBRARY FEB ~ 3 1983 HARVARD UNIVERSITY NUMBERS 437-463 1976-1980 CAMBRIDGE, MASSACHUSETTS 02138 U.S.A. 1980 BREVIORA MUSEUM OF COMPARATIVE ZOOLOGY CONTENTS NUMBERS 437-463 1976 No. 437. Anolis Alumina, New Species of Grass Anole from the Barahona Peninsula of Hispaniola. By Paul E. Hertz. 19 pp. July 30. No. 438. Crocodilians from the Late Tertiary of Northwestern Venezuela: Melanosuchus Fisheri Sp. Nov. By Carmen Julia Medina. 14 pp. July 30. No. 439. Ona New Specimen of the Lower Cretaceous Ther- opod Dinosaur Deinonychus Antirrhopus. By John H. Ostrom. 21 pp. July 30. No. 440. West Indian Anoles: A Taxonomic and Evolutionary Summary. |. Introduction and a Species List. By Ernest E. Williams. 21 pp. July 30. No. 441. Lizard Karotypes from the Galapagos Islands: Chromosomes in Phylogeny and Evolution. By D. Paull, E. E. Williams, and W. P. Hall. 31 pp. December 27. SWF No. 442. A Laboratory Study of the Turkish Hamster Meso- cricetus Brandti. By Charles P. Lyman and Regina C. O’Brien. 27 pp. May 27. No. 443. Epigonus Trewavasae Poll, A Junior Synonym of Epigonus Constanciae (Giglioli) (Perciformes, Apogonidae). By Garry F. Mayer and Enrico Tor- tonese. 13 pp. May 27. No. No. No. No. No. 444. . 445. . 446. . 447. . 448. 449. 450. 451. 452. Stations of the Thayer Expedition to Brazil 1865- 1866. By Myvanwy M. Dick. 37 pp. May 27. Natural History of Cerion. VII. Geographic Varia- tion of Cerion (Mollusca: Pulmonata) from the Eastern End of Its Range (Hispaniola to the Virgin Islands): Coherent Patterns and Taxonomic Sim- plification. By Stephen Jay Gould and Charles Paull. 24 pp. November 30. 1978 A New Interpretation of the Mammalian Teeth of Tribosphenic Pattern from the Albian of Texas. By Percy M. Butler. 27 pp. December 20. Food Selection by Beavers: Sampling Behavior. By Stephen H. Jenkins. 6 pp. December 20. Systematic Notes on the Loons (Gaviidae: Aves). By Robert W. Storer. 8 pp. December 20. 1979 South American Anoles: The Species Groups. 2. The Proboscis Anoles (Anolis Laevis Group). By Ernest E. Williams. 19 pp. February 21. A New Species of Moenkhausia from the Mato Grosso Region of Brazil (Pisces: Characidae). By William L. Fink. 12 pp. February 21. A New Species of Cybotoid Anole (Sauria, Iguani- dae) from Hispaniola. By Albert Schwartz. 27 pp. February 21. Lizards of the Sceloporus Orcutti Complex of the Cape Region of Baja California. By William P. Hall and Hobart M. Smith. 26 pp. February 21. No. No. No. No. No. No. No. 453. The Large Palaeotragine Giraffid, Palaeotragus 454. 455. 456. 5 457. 458. 459. 460. . 461. Germaini, from Late Miocene Deposits of Lotha- gam Hill, Kenya. By C. S. Churcher. 8 pp. Febru- ary 21. Evolution of Life Histories: A Comparison of Anolis - Lizards from Matched Island and Mainland Habi- tats. By Robin M. Andrews. 51 pp. July 31. A New Paleocene Palaeanodont and the Origin of the Metacheiromyidae (Mammalia). By Kenneth D. Rose. 14 pp. July 31. Description of a New Hawaiian Gobid Fish of the Genus Trimma. By Phillip S. Lobel. 15 pp. July 31. 1980 Two new Species of Eleutherodactylus (Amphibia: Leptodactylidae) from the Lowlands and Lower Cloud Forests of Western Ecuador. By John D. Lynch and Kenneth Miyata. 12 pp. January 31. A New Species of Ate/opus (Anura: Bufonidae) from the Cloud Forests of Northwestern Ecuador. By Kenneth Miyata. 10 pp. January 31. A New Species of Dendrobates (Anura: Dendrobati- dae) from the Lowland Rain Forests of Western Ecuador. By Gregory O. Vigle and Kenneth Miyata. 7 pp. January 31. On the Evolution of the Jaw Adductor Musculature in Primitive Gnathostome Fishes. By George V. Lauder, Jr: 10) pp: June 30: Geographic Variation in Anolis Brevirostris (Sauria: Iguanidae) in Hispaniola. By Douglas L. Arnold. 31 pp. June 30. No. 462. Eleutherodactylus Eremitus, a New Trans-Andean Species of the Lacrimosus Assembly from Ecuador (Amphibia: Leptodactylidae). By John D. Lynch. 7 pp. June 30. No. 463. Jaw Musculature of the West Indian Snake A/sophis Cantherigerus Brooksi (Colubridae, Reptilia). By Kenneth V. Kardong. 26 pp. August 15. BREVIORA MUSEUM OF COMPARATIVE ZOOLOGY INDEX OF AUTHORS e NUMBERS 437-463 1976-1980 ENN DRIEW Sse OBUNG Wiis c0.505 ocean guna scvamsciebciacesseebidctoseocncesees 454 EMR NOLED se 1) OU GAS IE was caccouos secsevesbacsecseosstosccovseereuteesenas 461 PAE RASECERGNee Mc, «cecistwas stcbesmseanosnbetomastsrdestadecsessoenamens 446 GOR GHERS2 Cie pactactecs stn cceccenonnccbeice wccsustansencueeees tunes 453 PDK VALVAVIAINIIYo= VAL Acesea ce ssleciecanek cousumonnicttee dcates cane neue ene 444 SINIKGaaWW TIGIETAIM Io seetaacnancceecesasasscacseces * i > ~ we © -_ ‘! MUS. COMP. ZOOL LIRRARY MAR_1 8 498 BREVI®. Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, MaAss. 30) fomya 1976 NuMBER 440 WEST INDIAN ANOLES: A TAXONOMIC ) AND EVOLUTIONARY SUMMARY 1. INTRODUCTION AND A SPECIES LIST Ernest E. WILuiAMs!? Asstracr. Accumulation of morphological, karyological and _ ecological data on West Indian anoline lizards permits and requires a taxonomic analysis more elaborate than usual and employing both formal and informal taxonomic categories. The categories are defined in this, the first paper of a series, and a species list of West Indian anolines displays the new arrange- ment. INTRODUCTION Since the pioneer study by Etheridge (1960) there has been a remarkable growth both in our factual knowledge and in the sophistication of our knowledge of one group of lizards — the anolines. This has been especially true in the islands of the West Indies. The mainland members have at the same time received less attention and have proved more refractory. (Cer- tainly the latter fact has influenced the first. ) There is, however, nowhere any gathering together of the new knowledge. Partly this results from the continuing activity. Not even species lists have remained constant. A statement that Cuba or Hispaniola has x species is outdated before it is pub- lished. In the same way ecological information, ideas and theory have expanded far past the published record. It is not easy to keep on top of the field. Just because of this it is necessary that beginnings of a sum- mary be made. Both for old hands and for newcomers an exposition of how far we’ve come, where we are, and where 1Museum of Comparative Zoology, Harvard University, Cambridge, Massa- chusetts 02138 2 BREVIORA No. 440 we might go will be a useful thing. I propose as a first step a taxonomic-evolutionary summary. The summary is inevitably provisional and, more than that, intended to promote research, provoke criticism and encourage the search for further evidence and the endeavor for greater understanding. My objective has been to illuminate ecology and evolution. The taxonomic study I will here provide is one means to that end. The West Indian anoles are a group of enormous diversity, but the interest of this diversity is not in its bare existence but in its structure and origin — its balance, the interlocking of its parts and the historical paths by which this was achieved. This is no simple radiation — not just the checkerboard sub- division of some original widespread unit. Here the severed units have doubled back upon one another and are completely layered, juxtaposed and interdigitated. The fitting together of so many species is the problem. Such ccmplexity involves several levels. One such level is generic. In the West Indies I recognize three very distinct genera, two genera autochthonous and old, species-poor and obviously relict, one genus species-rich beyond ordinary imagin- ing, a colonizing and expanding group, a newer invader from the mainlands that adjoin the West Indies to the west and south. The latter is, of course, the group, as full of problems as of interest, that must receive the maximal attention it deserves. In point of fact, there are few problems for the two old species-poor genera, but also little information. One of these two genera, Chamaeleolis, is represented by two giant casque-headed arboreal species on Cuba which much re- semble the tree-crown giants of the Anolis equestris group with which they share the island (Garrido and Schwartz, 1968). The species of Chamaeleolis, however, are more primitive osteo- logically, more chameleon-like in movement and appearance and apparently rarer than giant Anolis. The other of the genera, Chamaelinorops, initially erroneously reported from the tiny mile-square island of Navassa, west of southwestern Hispaniola, is, in fact, from the south island of Hispaniola, dwarf, ground-dwelling and extremely peculiar osteo- logically. Richard Thomas (1966) recognizes only a single species. Chamaeleolis and Chamaelinorops, except that they are true anoles sharing the characteristic adhesive pads of Anolis and the typical Anolis dewlap, are not close to each other nor to 1976 WEST INDIAN ANOLES lI. 3 Anolis. Their greatest interest lies in the possibility that they may represent an early (pre-Miocene ?) invasion of the Greater Antilles and may be relicts of an earlier island radiation of which we otherwise know nothing. The three species just mentioned apart, the remaining anolines of the West Indies are here regarded as belonging to the genus Anolis. Fortunately this overlarge taxon divides naturally, as Etheridge showed in 1960, into two sections called by Etheridge alpha and beta. Though this is a dichotomy based on an ap- parently trivial character, it makes excellent geographic sense. Savage (1973) has suggested that, instead of two sections, two genera be recognized — Anolis Daudin (type Anolis carolinensis Voigt) and Norops Wagler (type Anolis auratus Daudin). This would substitute formal designations for the currently in- formal ones but it would leave no formal (or informal) term for the two sections (or genera) taken together. Savage’s action is well taken if alpha Anolis are closer to Chamaeleolis and Phenacosaurus than to beta Anolis or the betas are closer to Chamaelinorops than to the alphas. This is a point I regard as at least doubtful, preferring to leave it in decent obscurity until there is more and better evidence. My own suggested phylegeny for anolines would have the alpha anoles the more primitive (as they certainly are in many respects), and the beta type of caudal transverse process (which does not resemble those of other iguanids or of other lizard groups) arising secondarily, but only once, from the alpha condition in which the transverse process is absent. The transverse processes of Chamaelinorops have only a verbal similarity to those of beta anoles; I question the closeness of the relationship. Richard Etheridge would dis- agree with this scheme fundamentally. Very recent immuno- logical data (Dessauer et al.) reported at the 1974 meetings of the American Society of Ichthyologists and Herpetologists ques- tion the fundamental distinction. I remain convinced of the reality of the two groups but, while so much remains contro- versial, I do not see the value of the formal designation; it 1s not even useful mnemonically. MEANS TO ANALYSIS OF A RADIATION: THE Group TrerMs UTILIZED The two sections of Anolis have, according to my interpre- tation, provided three and only three invasions of the West -} BREVIORA No. 440 Indies — one by betas into Jamaica, one by alphas into His- paniola, and a third by alphas into St. Lucia. (But see Yang, Soulé and Gorman [1974] for the evidence for a landfall for the third invasion in Grenada instead.) All the extraordinary proliferation, diversity and complexity of Anolis in the West Indies has arisen out of these three stocks by intra-island radia- tion and inter-island interchange. There is therefore a formidable problem in analysis. Fortunately, part of the basic information is already available. A just published checklist of West Indian Amphibians and Rep- tiles by Schwartz and Thomas (1975) provides an informed and very careful list of Antillean taxa (including Anolis) with original citations and synonymies, as well as the distributions as known to the date of publication. The species are, however, listed alphabetically; no taxonomic arrangement or indication of rela- tionship is attempted. The taxonomic ordering presented below, in remedving this, endeavors to synthesize a great deal of bio- logical information. I have myself seen 72 species or members of superspecies of West Indian anoles in the field, six additional alive in captivity, and, 33 more as preserved specimens. Only three species, all very recently described from Cuba, are known to me only from descriptions (A. pygmaequestris, A. juangundlachi and A. fugi- tivus). I have collected and studied anoles on all four of the Greater Antilles, on several of the Lesser Antilles and on one ‘sland in the Bahamas. This field knowledge I regard as basic to an undestanding of the group. I have also participated in, encouraged or aided ‘studies at many other levels — osteology, karyotypes, electrophoresis, aut- and synecological studies. All of this information is utilized in the classification below. No classification can mirror perfectly the complexity of the evolutionary events that have produced the more than 100 species of West Indian Anolis. Nor, indeed, are the minuter details of relationship and evolutionary sequence so well under- stood (or likely to be) that we should attempt so perfect a system. Nonetheless the wealth of species to be allocated and the amount and variety of detail known about these same species seem to me to afford at once the possibility and the justification for an arrangement elaborate much beyond the usual. I there- fore utilize a number of informal terms, partly based on those employed by Etheridge in 1960, but descending into greater detail. I define these below. 1976 WEST INDIAN ANOLES 1. 5 FORMAL AND INFORMAL CATEGORIES USED Section. The primary dichotomy, a group osteologically de- fined at the highest level below the genus. Proposed by Etheridge (1960) for his alpha-beta division with Anolis osteologically defined on presence or absence of transverse processes on pos- terior caudal vertebrae. Subsection. A division setting off a major portion of a sec- tion, again osteologically defined. This term, not used by Etheridge, distinguishes punctetus and carolinensis subunits within the alpha section, basing them on the relationship of interclavicle and clavicle. In the shorthand terms used below, the relationship is described in terms of an arrow-shaped or T- shaped interclavicle. Series. A phyletic unit under the subsection definable on multiple characters. Osteological, chromosomal and even scale characters are utilized. Ordinarily this includes more than one species group and displays substantial morphological and even chromosomal diversity, i.e., products of a complex radiation that inferentially included several intermediate or annectent forms now extinct. I have found series to be the unit in terms of which evolution is most conveniently discussed. I shall, therefore, in succeeding papers mention more characters under this unit than under taxa at a higher or lower level. This permits higher compara- bility between series and puts on display also many of the features and conditions the evolution of which I will later trace. This is not to say that the greater number of characters makes the definition of series sharper or more rigorous. On the con- trary, precisely because series are the units within which evolu- tion is most readily seen, recognition of series is a matter of some subtlety— as Tables 1-4 show. Morphological and karyological characters may broadly overlap (Table 1). Ecology and geography are major clues, but convergence in ecological adaptations is rampant (Table 2), and geography must be used with discretion. A balancing of all the evidence — not all of which is on record in Tables 1-4 — is the basis for the recognition of series. subseries: a category utilized when a series has several recog- nizable subunits, ordinarily including more than one species group. 6 BREVIORA No. 440 species group: the products of a simple radiation but often in- cluding species now widely sympatric (they may exhibit chro- mosomal diversity). I have sometimes used species group for a single species when that species is very distinct and may well be the last remnant of a radiation. species subgroup: employed when readily dennavley and sym- patric subgroups can be determined. superspecies: the products of a radiation, the representatives of which are still completely or mostly allopatric and usually chromosomally uniform. Species status often uncertain. (See below. ) species: the most recent evidence indicates that the recognition of valid species is much harder than was formally assumed. It cannot be routinely assumed that allopatry or parapatry imply subspecific status. As I indicate above, I have used the superspecies category for a number of equivocal or doubt- ful cases. subspecies: intraspecies differentiation differs so much in dif- ferent species that this unit is of very different value from one case to another. I do not report currently recognized subspecies in the species list below. I will do so in the de- tailed species group and species definitions to come, but I will do so without passing judgment on their validity. I imply only that the differentiation so indicated —in contrast with that implied by the use of superspecies— is clearly- intra- specific. WEST INDIAN ANOLES 1. 1976 ‘saqias sty) ul siaded Surpaacons ul UOrssnosIp Ja[[NJ ay UT UIATS aq [[IM Inq JIaY pai! Jou are saSuEI ULIPUT IsoA\-UONz ‘say[NUY JassaT usayinos ayi Jo dnoss saiads zanbox ay) Jo staquiaut snoraqo aie Aayi asnesaq (snunjjnbuvjq pur stsuasvuog) eyinbuejg pue aiteuog jo sajoue ay) apnypout J ‘saanefat uvaqqueg OU aABY saAtasMIayI sajouR ay) asnedeq ‘samTUye UeIpUT 89M PAY Spurs! asayi UO sayndar Jao Ysnoyre (Ajaatadsa1 40j02U02 pure roOYyouid) saIpuy Ue pure vIDUaPIAOIg JO safoue ay) Ist] sty} Wor yWWO J ‘oeIDUASOIPT ‘saIpUy IS9AA 9YI JO suONTUYap Iso dyI][ ‘SI SaIpUy IsaAA Fy) JO UONTUYIp AWr WUIIUTA 3S L8Q. UBewUeEy snasus oseqoy, ‘epruair) LEQ] Uoiqig puv [sung Ipupyri saroadsiadns ipavy ou qureuog S76 UIAYINY sisuaspuoqg eymbuelg OFG Yourowumnyx, snuvjypinbuvjq RINT] 3S LQQ] URrULIeD api9n] satoadsuadns apian} dnois satoads ani9n) sates janbo uonoasqns snyojound uonoes vydiy NIGAVGesieuy © .sngen) ejoruedsiyy 6161 LGIWHOS 1noqsvg — satsads LAUINHDS sdesounavwpy;) snuasy eqny F981 AdOD snIs0d eqn) L681 NOUPIE GNV TIMANAC sapiwoajavwmvys — satoads QVALOIOD Sijoajavuonyy) snus) zed IO pueysy UOX® |, t'SHIGNI LSAM FHL AC SANIIONV FHL No. 440 BREVIORA ‘oyedurks Ajapim aq 0} Way) syiwied jeyy UONeNUaIayIp 709190j029 yo apezd ayy paddtyoe you sey somadssodns xv jO Sloquau JO UONeNUDIZyIpP ay) yey? WJ 9qeNsuousp sy OF ATafos UONUDNe s[feo IY “spraqdy apMasyur Jo ouoz |[eus ew ATUO YIIM 30 SUIPsaigiajJUI ynoyVIM yoBIUOD UT oq ©} UMOUY aie AURYY ‘sateds [edTS0forq Jou ate so;adsiadns Jo siaquiow 94} Jo Jsour 10 Aueur yeyy ATduit jou Soop 2UdIIIJIP sty} UO siseydwa sy, “uastIe U9j0 aavY snyeIs sadeds yo suonsanb yoryM noqe sarvads o1yedoyye Apsou 10 onnedoye pajpjas Mjasoj2 asoyy YUM Jseguod sapads onneduds youNstp Ajdaeys ay} Buryeu yo asodind ay) s07 Ajaaua st sapodsiodns jo sioquiaw ayy Jo aso ayy ut idaoxa suaquosap savads jo saumeu yi soy s[ertdeo Sulsn JO dIAap IY ¢ Ory oVong eporuedsipy vjoruedsipy vporuvdstpy BAqg[ny onbrunue yy epnullag UO psonpoiUr ‘sopeqieg vURANL) ‘peplulyy, uo psonpowur ‘epeusiy pepluly, uo paonposur ‘us.UuTA 39 yurg 10 pueysy Salas snpvjnovuig Oc8l WHY Maeno dnois sarads twaiang FOBT edop snjzvajvq C96] SWIM avuoyving LE8t wouqig pue found upsooiu soroodssodns 2psooi IS6l LNVYO 4]92asoos dnois sarads upsoou SOLIS alan 88 2epodaoey janhou OFS ueULIeED snwasxa Avy) snauav sorsadsaadns janbou dnois satsads janbou eNHMLAT GNV LGOUYVHNIDY Sipps UOXe T WEST INDIAN ANOLES l. 1976 ‘WT ouieu JOU pIp Inq ‘UOXe) sy) JO ssauJIUNSIP 9YyI palejsuoWap (fF/G[) SUING puUe JaISqaA\r yurg vneig eqesg yueg eypinsuy XIN) "IG ETNA NS vporuedsipy vjoruedsipyT ‘aavuoy vpolurdsipy epluoyy ul paonponut ‘eforurdst PHO, ul psonp [ “ejor 'H vplo[y Ul psonpour ‘seureyeg oor oVong oory oLong PBL] uurusedg snynjnovwuig sapadsiadns snyojnovuig L88T NVINGVD Snungps P98 AdOD snwawus dnoisqns snjpjnovwg dnois satods snyvjnonuig 9S8t TTaMOTIVE s2nop dnois sarads snynov saliasqns snypjnavuiig UdISSVH GNV WIGON ssue[saeje suing pue I9Isqa\, YW satoadg GEO) ULIOO)! SU pry? OL8I Whnosog sujsomaasq satadssadns s14jsou1aaqg COB UexIN'T pur ipreyUray seuasrurmop 1981 odor snysusip saroodsiadns snyousip dnoisqns snyoysip [98 HdOD snjnivas dnoisqns snjnjoays dnois sarads snjynqo.js F061 WAOANLALS tuunusaaa dnois saieds tuunwaaaa Saliasqns snjnjo1s No. 440 BREVIORA 10 vjoruedsipy vjoruedsipy vjoruedsipy vporuedsipy vjoruedsipy vporuedsipy yurg eyMsuy eed eHelsS epnqieg ensynuy BOTUTLUO(T qqueyer) suey yurg odnojapens P1198] UO JY epuopoy yurg ensnuy yuegq 10 puPysy oO) ACOH TIL SN CARTE S661 PIGON Sypiqinsuo] 6S6[ UeIyIOD laaasys FSG Uelyoon) wnowsv /8Q] URULIVy snuDyany CIB edon sazogdo sotoodsiodns sajyoqgta dnoas satseds sajoqto soliasqns sajoqda SIIIS §N]J9]VISILI CLOL [PZe'] sndod CLE M92] t2j4pay 9s C76] dnoqieg wysassof $68 Jasuatnog zs7jpaM satoadsaadns 25770 dnois satoads isyjom 0881 AdOD snjvjnI0 dnoisqns snjvjnzo0 $981 odor snassaf LEQ] Uolqig pue [own snypsowsvul LOS Ce as rier L8Q, UeWUey snpqnu sopadsiadns snjpsouwvu LE8[ Uoruqiq pue [MounN¢y ay IVI] UOXe [, 11 WEST INDIAN ANOLES Il. 1976 vporuedstpy OORY OWN vyorurdsipy OorYy OWN PUuO! euRnoleyy ‘yuRg soore) ‘enseuy Oo OWN. Oo OWong ONY OWN OY OWN OOTY O}JLONG vporurdsipy CE61 NVUHOOD 2osujvp dnois sawads iuopsurpuvp SITIIS 1U0]SU1]LDP 6961 OUGATY GNV SINVITTIM, $77"290 _ Satiasqns snjjn290 bL6L ZLUVMHDS 2uyjdays sotiasqns 2upjdays Solas §N]JNIIO WONIISYNS SISUIUOLDI SGD 12 22 FOG] tosoulayg sisuauou saroodsaiadns sisuauou L8Q[ UeuLIEy snjdisys LEQ] Uoiqig pue [lawn snpjjaypjsi49 sotadsiadns $19j]9)D)S149 dnoas saiads snjjaqpps149 FOG] WADANLALS sesuasuod L68l NOWIA ANV TYAN Od sigjay ang 9/81 SUALAd ny dnoasqns snpjayojnd 9/81 SUALAd y2jpuns dnoisqns 1y9njpuns dnois satoads snyayajnd SOIISQNS $7)]9]VISILI CLG] SWRI loupssvU No. 440 BREVIORA eqny eqny eqnr) vqny eqn’) eqn vpotuedstzy vjorurdstzy vjorurdsipy vporuedsizy vjoruedsipy vporuedsipy vporuedsipy vjoruedsipy yueg 10 purysy dnois saieds snuptzos0;y9 GL61 OCGIYAVD Slsanbavusig F9G[ ZUVMYIG avoovinq F9GI ZWIeMYIG tpooayjvus CE6I AeIYg pue mnoqreg tajqou CEG Jesse pue s[GoN swvpnsoajny OZ8l Wwearayy stugsanba saroadsaadns stujsanba dnois saroods stajsanba SOLIS SISUaUyosDI 1961 GNVY tvudooy bL6l UALSHAM CANV SWVITIIM 20uedns 9661 AAUAHS 270917u0UW dnoisqns pjooyuow L961 ZLUVMHOS AUNV SVWOHL “néowis C961 SWVITIIM ?25puayza dnoisqns 2aspiayja dnois sorads vjyooyuowm 0961 SWVITTIM 22ydojstuys dnois sorads raydojsiyo E461 ZLAVMHDS /ajmof dnoas saisads wajznof SOIIS DjOINUOW 6961 GNVY ANV SWVITIIM s2p2j0su2 UOXe |, 13 WEST INDIAN. ANOLES l. 1976 ‘Atuo dnois ay) jo satoads add} ay) se parts orqnday uvoTUWOG ay) JO BIeD UNIeP VAVIg IYI UT stuvjnsuts *p Sur[quissat wioy podde[map-an[q peqiiosapun ue st d19Y4 Lz *paddoip aq prnoys 8urdnos8 sapodsiadns ay) ‘payeajsuowap sty) WIA “SAAD LIAL Isasopa JOU ‘sjuapeAInba [eorsojooa Aju aq AvU asoyLt VSSVAR NT uewaArr) IVT les Avy yurg vuiryrg worn eqnr) vporuedsipy vpoluvdsipy vporuvdsipy vjorurdsipy vporurdsipy vporurdsipy vporurdsiyy GIGI wrlwuyosg sda2suo) 8g. URULIeD IpspudvU GEG] PAeTyS puv anoqieg Ipjy7.uuv{ GE6l eAwYg pur Moqieg snupsoivwus OFS Avg snyvoL0d clLEBI WIOA sesuauiosy9 | sarodsaodns sisuauro1v9 dnoisqns siuauyosv9 dnois satoads sasuauyosva COST SWIM Snyvydar0yru0p ECG da[sseP{ pure s[GoN sisusoonsoyng C76 UWeIyoor) 2uossapuay satoadsuodns ruossapuay dnois saieds tuossapuay G96 SWIRTTTLM Sluvjnsuas 6S6[ SUIMIP, 4adzuyD saradsuadns sadiuyp Z98I edory snuaysajao9 LEB] Uuomqig pue [own snupd2040]49 ssavedsiodns snupd2010]49 No. 440 BREVIORA 14 eqny eqn) eas) ean) eqn) eqny eqn’) aul) eqny eqny Aer uooyw Jey ‘spueysy Avg ‘eqn SUrPICN. yuerg 10 pureysy 198[ edo snasnjnyo GEG] Vo1yg pu IMOqIeg v7091049 satoodsiadns snaavjnjo dnois satoads snaspvynjv SOIIS sNaID]NJD LE8l NOU ANV THN snepnotusaa 8661 NVYHOOD 14951404 dnois sateds snypjnoimsaa 198][ don snjoajuasuv LEQ Uosqig pue [uouNd sn] saisadsaodns snion) dnois satsads snianj SOLIS SNIIN| Le8l NOUWIE AGNV TWYANN Yuvisdo7 OL61 SUa.Lad $247099 6981 AdOD Starvjjis4v dnois saivads snasnppiauv 996] Aplex, snusazod 9681 TTAMOTIVH S4asusnsup sarsodsiadns sdaausndup dnoisqns sdasysndup 1981 AdOD sigajosz 8661 WNOCAVEA /vosT]D $681 Woy snauunsg UOXe J, ae ‘snyvautiuas OVUT A[[eUT]D 981OUT 0} pa}eysUOWap Useq sey ‘SUIRIT[IM 9DUDLYIOI *p ‘paztuSooa1 Aysnoraaid saiads paryy oy, *(uonesedeaid ur ‘Z319H) e[Nsulusg PUOYLIeY 9} Woy savods poqmosapun uyt voreuref 6C6l SNVITIIM 8&8 COOMUHGND Shipuosas vOIVUe [ OF8L AVUD sndojauy Borer [ OS8l ASSOD snuyodo uewAer puri LQQ[ UvUIeEL snsuadsuos epnuliag oyUT poonpo.nut ‘vorewel CHg] Avaig lupynss a sapodsisdns mupyps3 n vorvure [ 6681 WHOUNTALS /upuss Sy dnois soweds wupynas o SOMIIS MUDYDAG = uoKoIs vIIg Z - eporuedstpy satoods Mou ‘ vjoruvdsiy GIOT Gains 4405572 A vporuvdsipy F9BL AdOD snppaurpuas S dnoas saweds snjpauyiuas eqnry) ZL6[ ZAeMYSG puR Opes snsipruva eqn) Slajyag wnajoaqs satoadsuodns wnajoads ASU) G/6) SVINOBIL GNY ZLUVMHOS Seis CS) GL61 OGINAVD fy2vjpunsuonl eqny) GL61 OGINAVD snayon{ i eqn) OL61 SUALAd sisuoppdadng eV) eqnr) 1981 AdOD) snanajdouvds eqny eqn eqn) eqny (Asta) No. 440 eqn’) eqny eqn’) eqn) pues] ueMms eqny ovlg uvuIdeT BpMOo,y OJUL psonpowut ‘aZztpagq ‘OIIXaPY JO svare ]e\soo ‘UBWART) ap] ‘seueyeg ‘eqn’ BREVIORA vorewef 16 yurg Jo puryjsy 1981 AdOO stdajorydo 1961 SINVITIIM CNV JIVEINe Svimea 6l6l NHGSWVY GNV WNOdAUVEA SNqevquqns 616] UspsueyY pue anoqieg snsoz7p Cool INoqivg yp sstoadsuadns snsoyp 9T6l NHGSWVY ANV YUNOGAVA 2347sau 8961 ZueMYyoSg snqnl 616] Uepsuvy pue mmoqieg 4afyjaro.upynb FOBT odor) sey rajowoy satoadsiodns Sty Ia]OWLOY FIG, Imoqieg iuosjau FIG] Ioqivg mwawaurg 888I UreUIey safiusisoajn) LE8I Uorqig pue [own] 2acs'ps sotoodsaadns 2assns dnoas satoads iausns Le8l NOU ANV TMAWAG uaiuappe dnois soweds muaruazpa SOLOS 19408 UOXe | ™~ ‘aiay aaetd e& darasap Ary) ‘poojs ™ = -zapun Ap10od axe yey) satias purpureui Jo aayeUOseIdar UeIpuy Isayq ATUO dy) Se Inq ‘satias-qns eB st sty BuNsty AW Uys ‘SqLI datj JO Jaquinu dy} puOdaS ‘sqlI paxy Jo Joquinu ay? ysiy saarts yotyM ‘ (aqqei Juasaid oy) ur se) epNUTIOJ [eoaWNU ev Aq paitodat Ja}DeIVY ITUIOUOXE)] & ST squ [eUOTdLDsut Suneroy 10 poxy Jo Jaquinu oy, ‘eIVUTWODOAU JY) UT daIJ JOH JO sqit [es1op Surpuodsai109 ay} 0} payee JsyI sie AZUL ‘(ggg ‘e8pli9yIy) spreziy pruensr ur vyeurwosoku ay) ut poppaquwa sjuamaa o8e[N4ed payioyed aie sqi jeuondrosuyz ‘ueWIOD 9310929 Wo st uoneuozyut odAjohiey ‘asplioyiq = preypry Aq paptaoid Ajsnoisuas 19M YOIYM ‘sIsay) SIY IOJ sayqei pue saj0U ay) Woy 91e BJep [eITBO[OI}SOr G ¢/§ r S/6P = = 82/0 33 — + raudos ane +/§ | OP — = LE-08/0§ 18 - + se ed a Ih G ES p — — 9§ I:¢ — snarnjnpv <| 9 p be euoulo P — = £6/96 «178 = + snign] 5 89 +/§ vole ee = — 9§ 3:2/1:¢ (—) + —— sasuauno.4va Ss 01/6 g re a + + 9g 1:6 + + siuysanba Z g/L G We 2 = 8F-0F/9E = d qr Dj0I1j WOU sre g wm 8 — (+4) pr/e 6/0 — —— uojSunanp 4 8/Ile g te Of = (+) 9g 1:6/0:9 ¢ =e $N}]NIIO a [uonvasqns siswau201v9 = dpIIAK[II9}UI — J — *] lee t/§ €/¥g © + = 6G-L3/9§ BS + + $j] av 48149 = 9/5 v/§ Vee a = 66-96 I'S ae 4p snyongvULrg 8/L (3) § 1G OS +: + 9§ 1:6/6:§ = + 1a1ana g/L +/¢ Wee = = £8/9§ 0h +: + ¢ Ganbou) [uonvasqns snzoja2UNd = apItAvpoI9IUT MOIIe — P| sjepned (apo) sTeides 1so19 $9.19 (uz) zSqul uado SAIUaAS awidasy srequiny -d1g depmoq [epney jeiqawiaAj adj jeuon juasaid saueyd -oAIey = -dosuy peruaydg Auojoiny SILIIS SOUP ULIPUT ISAAA JO S19IIVILUD paqIa]I¢S D tl DIAVL *(payesedas Ayjeostydes80a8) o1yedoyye Jo (paieredas Aqjeoneurtys) stdojojye Ayfensn ase ydiowosa auies ay) jo sxaquiaut ynq ‘o1doiuds uaqjo aie sydiowio9a yUua1ayIG ‘sauieu tay) Aq paisassns se ‘puno18 ay) UO 10 saaq) Ur suOTeISs oNStAIDeIeYD YIM pue ASojoydiow way) jo sywadse sayi0 ut pue 1ojoo pue adeys ‘azis ur aanounsip sadAi aandepe (Z/61) suet pue (6961) sumer pue pury jo asues ay) uy, ie ats ele 19LODS + i + ap + a d is snaonjnjv a8 snionj + + + + SISUIUIJOADI 4 stajsanba ab a. aS pjOI1jUoU a ae 1U07)0U1)4Dp + §N]]NIIO0 I[IIALPIIIIUL J —” No. 440 BREVIORA + + §N]]9]DISIAI + as + ae snypynavUg + awn Be (janbo.) IPIARPIIOIUL MOIIE — % — ee eee ee gayjo 8=9yund =6oysnq.) ss punors —sjaemp UMOID =jIeMp -queIo. = queIs ~—s-»Istyeaauas SALUAS pue sseis. = yunt) = uMoID yunt} = SIM) SIM} UMOIO yor yuny tSHdYONOOA 6 HIAV.L 18 19 WEST INDIAN ANOLES l. 1976 say[QUY Jassa’y +++ oory OWN a 1940 DS +1 1ULDY DLS J + snaavjnjv sni9n} SISUIUL]OLDI siujsanba DJ0I1]UOUL 1U0]8 U1] LDP $n]jnII0 ack aie [uonsasqns srsuauiyosv9 = apotaepai9}u1 — J, — | §N]]910]8149 snqvjnavuiig 1a1ana (ganbo.) [uonsasqns suosfi70) = apIAvpIIOIUL MOLIE — 7} eporueds 1Fy salias JO UONNGIsIG § TIAVL vorewe[ eqny SAHTYas No. 440 $1]]9VDISIAD S$1]]9]DISIAI DjOI x -4Uuou +- D102 = snqpjnd —--2qU0UL + a pjonjuow “DUG = snazDvIN}D —_§n}Ja4} 5119 ~Q taudvs +- 1uDpYyvid 1949DS +- sisuauy 1a49D8 -- snion} -OLD2 —- snaavynjv 1919S ayo yuniy ysnq punoi3 sseis yunay snjpjno snjpjno -putiq +- -DULIg $nq]NII0 1u0j3ut] sisuauy -ipp + -OLDI $n})NII0 qupypois = (up yvas) sisuauty sisuauy -OLDI -OLDI uMO.D jiemp yuna 31my qganbou snjvjno -DULIG 1uauang 1404 -Sutjivp 14910NI 1a40D$ 1uDYy DAD nuvyvid stujsanba yueis juers IST] 31M} uMO.9 -B19Ua3 puvysr Aq sydiourosa yo (satias) aoInos d1WoOUOXe J, 20 v ATaVL saynuy IassaT u1ayINOG say[nuy Jassa’] LRELIBRCOING OTN 01nd pjorued “STH porewe eqny SS a ee 1976 WEST INDIAN ANOLES l. DA ACKNOWLEDGMENTS The new data on West Indian anolines is the work of many hands. The singularities of the present taxonomic arrangement are the author’s own. Financial support for the work result- ing in the present paper has been provided by the National Science Foundation under grant GB—37731X and prior grants to E. E. Williams. = REFERENCES CITED ETHERIDGE, R. 1960. The relationships of the anoles (Reptilia: Sauria, Iguanidae): an interpretation based on skeletal morphology. Ann Arbor, Michigan: University Microfilms. xiv + 236 pp. 1965. The abdominal skeleton of lizards in the family Igua- nidae. Herpetologica 21: 161-168. Garripo, O. H. AND A. ScHwarrz. 1968. Cuban lizards of the genus Cha- maeleolis Quart. Jour. Florida Acad. Sci. 30: 197-220. GorMAN, G. C. 1965. Interspecific karyotypic variation as a systematic character in the genus Anolis (Sauria, Iguanidae). Nature 208: 95-97. 1974. The chromosomes of the Reptilia, a cytotaxonomic interpretation. Jn A. B. Chiarelli and E. Capanna eds., Cytotaxonomy and Vertebrate Evolution, pp. 349-424. Academic Press, New York. GorMAN, G. C. AND BraAp StAmMM. 1975. The Anolis lizards of Mona, Redonda, and La Blanquilla: chromosomes, relationships, and natural history notes. Jour. Herp. 9: 197-205. Ranp, A. S. AND E. E. WitiiAms. 1969. The anoles of La Palma: aspects of their ecological relationships. Breviora Mus. Comp. Zool. No. 327: Eal7e SAVAGE, J. 1973. A preliminary handlist of the herpetofauna of Costa Rica. Los Angles, 17 pp. ScHWARTZ, A. AND R. Tuomas. 1975. A checklist of West Indian am- phibians and reptiles. Special Publication Carnegie Museum No. 1: 1-216. THomas, R. 1966. A reassessment of the fauna of Navassa Island. Jour. Ohio Herp. Soc. 5: 73-89. WitutaMs, E. E. 1972. The origin of faunas. Evolution of lizard congeners in a complex island fauna: a trial analysis. Evolutionary Biology 6: 47-89. Yanc, S. Y., M. Sourf AND G. C. Gorman. 1974. Anolis lizards of the eastern Caribbean: a case study in evolution. 1. Genetic relationships, phylogeny and colonization sequence of the roquet group. Syst. Zool. 23s SO -o9o. a —<— | ¢ od < ~ Ba if =_ ‘ =. - ~ - SS is ae - - ial ° : ae = 7 MUS, COMP. ZOOL LIBRARY MAR 1 8 1985 BRE V 1IG2R A Museum of Co mparative Loology US ISSN 0006-9698 CamsripcE, Mass. 27 DECEMBER 1976 NuMBER 441 - LIZARD KARYOTYPES FROM THE GALAPAGOS ISLANDS: CHROMOSOMES IN PHYLOGENY AND EVOLUTION D. Pau,’ E. E. WituiaMs’ anp W. P. HALL? Asstract. The iguanid lizards, Conolophus subscristatus, Tropidurus albe- marlensis, T. delanonis, and T. duncanensis have similar 2n — 36 karyotypes. C, subcristatus has a 12 metacentric macrochromosome and 24 microchromo- some karyotype that is here shown to be primitive for the Iguanidae and probably for all lizards, while the three Galapagos Tropidurus have iden- tical patterns to Tropidurus species from eastern South America and differ from the primitive karyotype of C. subcristatus by non-Robertsonian modi- fications of three pairs of macrochromosomes. All available karyotypic data for the Iguanidae are summarized and used to discuss hew one may determine which karyotypes are “primitive” within radiations and what possible roles Robertsonian karyotypic variation may piay in the process of evolution. Analysis of karyotypic and systematic in- formation suggests a causal relationship between karyotypic differentiation and the rapid proliferation of new species, such that the need for geographic isolation seems to be minimized by the chromosomal differentiation. INTRODUCTION Among the karyotypically well-studied families of squamate reptiles, the iguanid lizards are known to show great chromo- somal diversity, with most of the variation apparently resulting from Robertsonian mutations (centric fusions and/or fissions) (Gorman, 1973; Hall, 1973). To more fully understand the biological significance and evolution of this diversity, workers 1Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138. 2Department of EPO Biology, University of Colorado, Boulder, Colo. 80309. 2 BREVIORA No. 441 at the Museum of Comparative Zoology have been accumu- lating karyotypes for as many and diverse iguanid species as possible. As part of this program, during the winter of 1969-70 Paull was able to karyotype four iguanid species fromthe Galapagos Islands. Two of the four principal phyletic branches of the family, the “iguanines” and the “tropidurines” (Savage, 1958; Etheridge, 1964) are represented in these islands, and both were sampled in the study. All four species showed a 2n = 36, 12 metacentric macrochromosome, 24 microchromosome pat- tern which is believed by some to be primitive for the Iguanidae or, indeed, for all lizards (Gorman et al., 1967, 1969; Webster et al. 1972: Gorman, 1973). Addition of karyotypes for these four species to our data base provides the occasion to discuss the evidence of ‘‘primitiveness” for karyotypes within a radiation and the role Robertsonian karyotypic variation may play in the process of evolution. How- ever, we must first describe the karyotypes of the Galapagos lizards sampled. KARYOTYPES OF GALAPAGOS IGUANIDS (D. PauLtLt AND W. P. Hatt) Specimens examined: Representatives of two distantly re- lated branches of the Iguanidae have reached the Galapagos, presumably by overwater colonization. ‘The iguanine species in the Galapagos belong to the endemic genera Amblyrhynchus (one species, the marine iguana) and Conolophus (two species of land iguanas). Of these, Conolophus subcristatus was karyo- typed. The tropidurine radiation is represented in the archi- pelago by eight endemic species of Tropidurus (lava lizards). Additionally, Tropidurus has a South American continental radiation of 12 species (Etheridge in Peters and Donoso-Barros, 1970). Island species karyotyped were Tropidurus albemarlen- sis, T. delanonis and T. duncanensis. Table 1 lists the species karyotyped and their collection localities. Methods: All chromosome preparations were made in the Galapagos Islands using laboratory facilities kindly supplied by the Charles Darwin Research Station on Santa Cruz Island. Cells were spread for karyotyping by air drying smears of metha- nol:acetic acid (3:1) fixed suspensions of testis, bone marrow or spleen tissues prepared directly from colchicine pretreated animals. The techniques used were similar to those of Evans et 1976 CHROMOSOME EVOLUTION 3 Table 1 Galapagos specimens karyotyped individuals species locality karyotyped Conolophus subcristatus South Plazas Id., Las Plazas 1 Tropidurus albemarlensis South Plazas Id., : Las Plazas 6 Tropidurus albemarlensis Bartolome Id., Sullivan Bay 4 Tropidurus albemarlensis Santa Cruz Id., between Galapagos Hotel and Darwin Station 6 Tropidurus delanonis Hood Id., behind beach, Gardiner’s Bay 5 Tropidurus duncanensis Duncan Id., small cove on NE end 2 al. (1964), Bianchi and Contreras (1967) and Patton and Hsu (1969). Results: All species had 36 chromosomes, with 12 biarmed macrochromosomes and 24 microchromosomes (Fig. 1). No cytologically distinct sex chromosomes or intrageneric variation of any kind was seen. However, conspicuous differences in arm ratios and relative sizes of the macrochromosomes were noted between the genera (Fig. 1). In Conolophus subcristatus (Fig. 1, lower), taking the macro- chromosomes in order of size, beginning with the largest, pair one is very slightly submetacentric; pair two is distinctly sub- metacentric, with the long arm slightly less than twice as long as the short; pairs three and four are almost exactly metacentric and, in many spreads, indistinguishable in size; pair five is nearly metacentric; and pair six is submetacentric, with the long arm about 1.5 to 2.0 times the length of the short arm. Pairs one and two are similar in length, three and four are slightly but distinguishably shorter than two, five is distinguishably shorter than four, and six is conspicuously shorter than five. Some of the microchromosomes seem to be metacentric or sub- metacentric, but our preparations do not resolve their structures well enough to allow them to be unequivocally paired. No. 441 BREVIORA ‘uoneoytuseur owes ay} 0} pajutid oie somn8y YIog *(Molaq) snzn]s209 -qns snydojouoy pue (aaoqe) siuwounjap sninpidosy wory saddyodrey afew yo uostaedwog *] ainsry ae? ©*¢ @# @= £@ @®@BRweeaee © Be @ & usa E a “-*9 © ® s+ -# @P@ poke @r geoeged O22 F «£ a 1976 CHROMOSOME EVOLUTION 5 etrgpsaeeets O@ Cd Oe -*** see 0O eg BO ‘see see eden @ of Pree" Figure 2. Comparison of male Tropidurus karyotypes. ‘The species are T. delanonis (left), T. duncanensis (middle) , and T. albemarlensis (right) . The karyotypes are printed to the same magnification. 6 BREVIORA No. 441 In Tropidurus (Fig. 1, upper; Fig. 2) pairs one through five show a fairly even gradation in length, with pair six being conspicuously smaller than five. Comparing the arm ratios to those of Conolophus, Tropidurus pair two is more submetacen- tric, with the long arm being slightly more than twice the length of the short; and Tropidurus pair five, rather than being meta- centric, is almost subacrocentric, with the long arm about 2.5 times the length of the short. Again, some of the microchromo- somes appear to be metacentric or submetacentric, but they are not adequately resolved to allow accurate pairing. PRIMITIVE KARYOTYPES CE. Eo WILLIAMS AND W,. PS Harr’) Comparisons with other iguanid genera and other families of lizards suggest that the Tropidurus pattern is derived with re- spect to the Conolophus pattern, which may be primitive for several families, including the Iguanidae. As Gorman (1973) in the latest review of reptilian chromo- somes has emphasized, our knowledge of lizard karyotypes and especially of iguanid karyotypes has increased immensely since the early work of Matthey (1931, 1933). In the iguanids even such very speciose genera as Sceloporus and Anolis have now been sampled very extensively and at least one or two species have been examined in all major subgroups of the family (Table 2). The more species studied, the more widely one chromosomal arrangement is demonstrated: that with a 2n of 36, with 12 metacentric macrochromosomes and 24 micro- chromosomes. The kind of 2n = 36 karyotype characteristic of the Galapa- gos Tropidurus (i.e. 2n, arm ratios, etc.) has also been reported for representatives of this genus in eastern South America (Gor- man et al., 1967; Peccinini, 1969; and Becak et al., 1972); however, we are not aware of its occurrence in any other genera. On the other hand, the 2n = 36 Conolophus karyotype, or at least the details of its macrochromosomal pattern, is found in many different lizard groups. In the Iguanidae (Table 2) pre- cisely this macrochromosomal pattern is found in such diverse groups as the anolines (Gorman, 1973), sceloporines (Cole, 1970; Pennock et al., 1969; Gorman, 1973; Hall, 1973), Crotaphytus (Montanucci, 1970), iguanines (Cohen et al., 1967; Gorman et al., 1967; Robinson, 1974), oplurines (Gor- 1976 CHROMOSOME EVOLUTION 7 table 2 Genera are those in Peters and Donoso-Barros, 1970 unless otherwise noted. Reference for number of species karyotyped and diploid number is Gorman 1973 or this paper unless other- wise noted. Number of Number of species species Range of Genus in genus karyotyped known 2n Amblyrhynchus le Anisole pis 2 1 230 Anolis carZ007 80s 2n=25-48 A peropristis* 1 — Aptycholaemus 1 ————— Basiliscus 4} 1 2n—36 Brachylophus os oe Callisaurus ee 2 2n—34 Chalarodon 1 ——. Chamaeleolis Oe 1 2n=3 Chamaelinorops le 1 2030" Conolophus 2 1 2130 Corytophanes 3 —— Crotaphytus oe 3 2n—30 Ctenoblepharis or == Ctenosaura™” 7 1 2n=—36 Cyclura a 1 2n—36 Diplolaemus ss) ——. Dipsosaurus oan 1 2n=—36* Enyalioides i 1 2n=—36° Enyalius BF” —- Holbrookia one 2 In=34* Hoplocercus 1 — Iguana 2 1 DNS Ss Laemanctus 2 —— Leiocephalus™ 20 5) 2n=32-36° Leiosaurus 4 — Liolaemus i Oisiea 8 2n=30-40 Morunasaurus 2 — O phryoessoides a —— O plurus 6! 1 2u—30 Petrosaurus 2 2 2n= 34" Phenacosaurus See 1 2n—36 8 BREVIORA No. 441 Table 2 (Continued ) Number of Number of species species Range of Genus in genus karyotyped known 2n Phrynosaura ) Phrynosoma 147% 9 2n=34 Phymaturus ne — Plica 2 1 2n—40 Platynotus”* 1 —. Polychrus GF 4 2n—20=3 0 Pristidactylus”® 4 1 20 Proctotretus 3 — Sator 2 1 2n—34*" Sauromalus qe 478 2n=—36 Sceloporus 64729 A Date 202 On Stenocercus’® 29 — Strobilurus*® 1 ——. Tropidurus 20 4 VAN 15) Uma Her g 2n—34 Urocentron 4.30 —— Uranoscodon +} ———— Urosaurus LOZ 5 2n—34° Urostrophus 3 —= Uta 62 6 2n— 34 Because they do not appear on the Etheridge dendrogram, two recently described genera have been omitted from the table: Vilcunia Donoso-Barros and Cei, J. Herp. 5: 90. 1971. Pelusaurus Donoso-Barros, Neotropica 19; 132. 1973. Both are said to be allied to Liolaemus and Proctotretus. Etheridge (personal communication) infers from their descrip- tions that they are allied to the Liolaemus-Phrynosaura-Cteno- blepharis complex and not to Proctotretus. 1Eibl-Eibesfeldt, I. 1962. Neue Unterarten der Meerechse, Amblyrhynchus cristatus, nebst weiteren Angaben zur Biologie der Art. Senckenbergiana, Biol. 43: 177-199: 2This is only a rough approximation. Continental members of the genus are in general poorly understood, and species are still being discovered in Hispaniola and Cuba. *We include unpublished data from material in the Museum of Com- parative Zoology. 1976 CHROMOSOME EVOLUTION 9 4Etheridge (personal communication) after examination of specimens prefers to recognize the genus as distinct from Leiosaurus. Cf Miller, L., 1922: Uber Aperopristis paronae Peracca und die Genera Aperopristis Peracca und Leiosaurus Dumeril and Bibron. Senckenbergiana 4: 153-159. 5Avery, D. and W. W. Tanner, 1970. Speciation in the Fijian and Tongan iguana Brachylophus (Sauria, Iguanidae) with description of a new species. Great Basin Nat. 30: 166-172. 6Smith, H. and E. H. Taylor, 1950. An annotated checklist and key to the reptiles of Mexico, exclusive of the snakes. Bull. U. S. Nat. Mus. 199: 1-253. Cophosaurus (Holbrookia) texana is placed with Callisawrus here. See Norris, K. S., 1958. The evolution and systematics of the iguanid genus Uma and its relation to the evolution of other North American desert reptiles. Bull. Amer. Mus. Nat. Hist. 114: 247-326, and Axtell, quoted by Norris. The only important character separating the two genera isthe condition of the car opening, which is evaluated as less important than characters separating other sceloporine genera. In fact, it would not be unreasonable to lump the three sand-swimming genera, Callisaurus, Hol- brookia and Uma. This larger genus would still have only 10 species. TAngel, F. 1942. Les Lezards de Madagascar. Mem. L’Acad. Malgache 34: 1-193. SGarrido, O. H. and A. Schwartz, 1968. Cuban lizards of the genus Chamaeleolis. Quart. J. Fla. Acad. Sci. 30: 197-220. Thomas, R. 1966. A reassessment of the herpetofauna of Navassa Island. J. Ohio Herp. Soc. 5: 73-89. 10Montanucci, R. 1969. Remarks on the Crotaphytus-Gambelia contro- versy (Sauria: Iguanidae). Herpetologica 25: 308-314. Montanucci, R. 1970. Analysis of hybridization between Crotaphytus wislizenii and Crotaphytus stlus (Sauria, Iguanidae) in California. Copeia 1970: 104-123. 11Cei, J. 1974. Two new species of Ctenoblepharis (Reptilia, Iguanidae) from the arid environments of Central Argentina (Mendoza Province) . J. Herp. 8: 71. In contrast to Cei, we continue to recognize Phrynosaura as distinct from Ctenoblepharis. On the key character of juxtaposed versus imbricate dorsals, Cei’s two new species are Ctenoblepharis. 12Etheridge follows an unpublished MS by Clayton E. Ray in uniting Enyaliosaurus and Cienosaura. 13fide Albert Schwartz and Michael Carey, who are preparing a revision of the genus. 14Smith and Taylor 1950, as in footnote 6 above. 15Etheridge, R. 1969. A review of the iguanid genus Enyalius. Bull. Brit. Mus. (N.H.), Zool. 18: 233-260. 16Williams, K. L. and H. M. Smith, 1958. Herpetologica 13: 265-267. 17Etheridge, R. 1966 (The systematic relationships of West Indian and South American lizards referred to the iguanid genus Leiocephalus. Copeia 1966: 79-91) listed 16 species in his restricted genus Leiocephalus. Subse- quent to 1966, Schwartz, by reinterpretation of already described forms, has raised the number to 20. O. H. Garrido, 1973 (Nueva especies de Leio- 10 BREVIORA No. 441 cephalus (Lacertilia, Iguanidae) para Cuba. Poeyana No. 116: 1-19) has added a 2lst species. 18Richard Sage (personal communication) believes the taxonomy of Lio- laemus to be still in a very primitive state. He suggests that the genus may contain as many as 100 biological species. loFritts, T. H. 1974. A multivariate evolutionary analysis of the Andean iguanid lizards of the genus Stenocercus. Mem. San Diego Soc. Nat. Hist. 7: 1-89. Fritts reallocates to Stenocercus a number of the species formerly placed in Ophryoessoides, leaving only six described and two undescribed species in the genus. *oLazell, J. D. 1969. The genus Phenacosaurus (Sauria: Iguanidae) . Breviora No. 325: 1-24. 21Presch, W. 1969. Evolutionary osteology and relationships of the horned lizard genus Phrynosoma (family Iguanidae). Copeia 1969: 250-275. 22Cei, J. M. and L. P. Castro, 1973. Taxonomic and serological researches on the Phymaturus patagonicus complex. J. Herp. 7: 237-247. 23Etheridge (in a letter): “I know of no reference that presents evidence for or against recognition of this genus. Most, but not all, recent authors include the species semitaeniatus in Tropidurus. *4Gorman, G. C., R. B. Huey and E. E. Williams, 1969. Cytotaxonomic studies on some unusual iguanid lizards assigned to the genera Chamae- leolis, Polychrus, Polychroides and Phenacosaurus, with behavioral notes. Breviora No. 316: 1-17. Polychroides synonymized with Polychrus. “5Peccinini, D. 1969. WVariacao nos cromossomos do lagarto Polychrus marmoratus (Sauria, Iguanidae) de diferentes localidodes (Nota preliminar) . Rev. Brasil. Biol. 30: 1-4. (The Polychrus from Sao Paulo with 2n—20 is actually P. acutirostris [P. Vanzolini, pers. comm.]. See also Peccinini, D., Becak, O. Frota-Pessoa and Iris Ferrari, 1971. Sex determination of the “pseudo-XO/XX” type in the Brazilian lizard Polychrus sp. (Sauria, Iguanidae). Caryologia 24: 129-139, and Becak, M. L., W. Becak and L. Denaro, 1972. Chromosome polymorphism, geographical variation and karyo- types in Sauria. Caryologia 25: 313-326. 26“Barrio (1969) has demonstrated Cupriguanus araucanus Gallardo 1964 is conspecific with Leiosaurus fasciatus Dorbigny in Dumeril and Bibron 1837. Barrio used the combination Cupriguanus fasciatus. However, Leiosaurus fasciatus was in 1843 designated the type of Pristidactylus by Vitzinger. If fasciatus is congencric with achalensis, the type of Cupriguanus, the latter becomes a junior synonym of Pristidactylus which then would be regarded as having four species, fasciatus Dorbigny 1837, scapulatus Burmeister 1861, achalensis Gallardo 1964 and casuhatiensts Gallardo 1968.” Etheridge (personal communication) . 27TLowe, C. H. and M. D. Robinson, 1971. The chromosome pattern in Sator grandaevus (Reptilia: Iguanidae) , Baja California, Mexico. J. Arizona Acad. Sci. 6: 282. 28Robinson, M. D. 1974. Chromosomes of the insular species of the 1976 CHROMOSOME EVOLUTION 11 chuckwalla lizards (genus Sauromalus) in the Gulf of California, Mexico. Herpetologica 30: 162-167. 29S8ixty-four Sceloporus species are currently recognized. When taxonomic revisions suggested by Hall, 1973 are incorporated, this number will be raised to about 72, with considerable uncertainty remaining concerning the number of biological species to be recognized within the torquatus and formosus species groups. 30Etheridge, R. 1968. A review of the iguanid lizard genera Uracentron and Strobilurus. Bull. Brit. Mus. (N.H.), Zool. 17: 48-64. 31Williams, K. L.. P. §. Chrapliwy and H. M. Smith, 1959. A new fringe- footed lizard from Mexico. Trans. Kans. Acad. Sci. 62: 166-162: Mayhew, W. W. 1964. Taxonomic status of California populations of the lizard genus Uma. Herpetologica 20: 170-183. 32Ballinger, R. E. and D. W. Tinkle, 1972. Systematics and evolution of the genus Uta (Sauria: Iguanidae). Misc. Publ. Mus. Zool. Univ. Michigan No. 141: 1-83. man et al., 1967) and the tronidurines (Gorman et al., 1967). In other families (Table 3) this pattern has been demonstrated in the Agamidae (Arronet, 1965; Gorman and Schochat, 1972; Hall. 1970; Sokolovsky, 1972), in the Teiidae (Gorman, 1970), in the Gerrhosauridae (Matthey, 1933; Hall, unpub.) and in the Amnphisbaenidae (Huang et al., 1967). This most widely distributed Conolophus-like karyotype is a source of controversy. On the one hand, it has been called “primitive” (Gorman and others). On the other hand, it has been interpreted, as by Cole (1970. 1971b), as derived in at least some iguanids, or, as by M. J. D. White, as a possible example of an exceptionally stable configuration that has been repeatedly evolved within a group. (White at one time called this “the principle of homologous change,” but he now prefers to call it “karyotypic orthoselection” [see White, 1973 for dis- cussion]. ) Those who deny the primitiveness of the 2n = 36 pattern hold very firmly to the concept that primitive karyotypes in lizards consist entirely of acrocentrics with karyotypic evolution then occurring by centric fusions of them. This view, that acrocentrics are prima facie primitive, has rested on the belief that fusion is cytologically much easier than fission and hence much more common (Matthey, 1949; White, 1954, 1959; Reiger et al., 1968). In particular, the generation of a new centromere, which supposedly occurs in fission, has seemed to lack any mechanism that would readily permit the 12 BREVIORA No. 441 event, while fusion has been interpreted as the result of the (conceptually) less difficult process of reciprocal translocation followed by loss of a centromere carrying small segments of one or both chromosomes. However, there are now many cases for which fission is an obligatory explanation of the origin of the karyotypes of highly derived groups and species — too many cases to allow any doubt of the reality of fission as one possible path of karyotypic evolu- tion — whatever its mechanism. Even White (1973) now ad- mits its existence under the name “‘centric dissociation” in cer- tain cases. Morescalchi (1973) finds fission the hypothesis of choice for the origin of the karyotypes of certain species of Hyla and. Eleutherodactylus. In reptiles, Webster et al. (1972) Table 3 Karyotypes in Non-Iguanid Lizard Families 12V -- 24m or direct derivatives Only karyotypes not readily occur in some member related to 12V + 24m known Agamidae Gekkonidae Chamaeleontidae Pygopodidae Xantusiidae Lacertidae GOV rtrd 2 a 24m 1a ( ) Anniellidae Gerrhosauridae ‘Teiidae (see Gorman, 1970) Anguidae Helodermatidae (10V + 4I + 24m) Varanidae Seincidae (see Greer et al., in prep. ) Based on karyotypic data in Gorman (1973), unless other- wise noted. 1976 CHROMOSOME EVOLUTION 13 presented evidence for the highly derived phyletic position of Anolis monticola, the one species within that very large genus that has a 2n as high as 48. It is similarly inescapable that fissioning has occurred several times in Sceloporus (Hall and Selander, 1973; Hall, 1973). A recent discussion in the journal Evolution has summarized some of the mammalian evidence for fission (Lawlor, 1974; Baker et al., 1975). It is no longer reasonable to peremptorily reject fission as a plausible mode of Robertsonian karyotypic evolution. The argument for the primitiveness of the 12 macro- and 24 microchromosome pattern for iguanids and for lizards, how- ever, does not depend on the supposed plausibility or implausi- bility of fission. The argument becomes easier to accept if fission is admitted in certain cases, but primitiveness for a karyotype, as for any other character state, can be determined on its own merits, independent of any theoretical mechanism for the evolu- tion of that character state. A large literature now exists dealing with objective recogni- tion of primitiveness. Kluge and Farris (1969) may stand as an example. They would use the following criteria (1969:5), listed in order of reliability: (1) The primitive state for any particular group is likely to be present in many representatives of closely related groups. (2) A primitive state is more likely to be wide- spread within a group than is any one more advanced State. (3) The primitive state is likely to be associated with states of other characters known from other evidence to be primitive. They add that “closely related groups can be selected through estimates of overall similarity that make no assumptions about primitive conditions.” “Widespread” they define not by counting taxa but as occurring in several taxa that otherwise would have little in common. They would also use “available fossil material.” At least in intention these criteria have merit, but, in general, such criteria are especially difficult to use in our present stage of knowledge of karyotypes. Fossils are clearly unavailable. It is still rare for karyotypes to be known for even a substantial number of any group and, on the contrary, those at hand may be a very biased sample.. The problem of real similarity may 14 BREVIORA No. 441 be serious; diploid number by itself is meaningless; there must be near identity in chromosome morphology paralleling taxo- nomic relationships inferred on other grounds. When a group has been as well sampled as the Iguanidae now are, however, the Kluge and Farris criteria begin to be applicable and the comparative method can lead to sound re- sults when appropriately applied. Cole misrepresents, indeed caricatures, the comparative approach when, in opposing the concept of the 12 + 24 karyotype as ancestral in iguanids, he says (1970231): “These conclusions are based on the assumption that the general karyotypic condition found in the majority of species that were available for sampling, at whatever level of the taxonomic hierarchy one happened to be working with, was, therefore, the most primi- tive,” and, again (1970231): “If I were to simply employ the principal [sic] on which these authors’ arguments are based, I would reach a rather different con- clusion for the spinosus group, for in this group the karyotype of the lundelli subgroup (12. biarmed macrochromosomes plus 10 smaller chromosomes, most of which are clearly biarmed; a 12 + 10 karyotype) would then be considered ancestral because it occurs in five of the nine species in the species group and none of the three remaining general karyotypes of the group is represented in more than two species.” Our own reasoning has a quite different base than that por- trayed by Cole. We start from a base that is broad and quite independent of the ideas of Cole or ourselves. Figure 3 provides a dendrogram of the phylogenetic relation- ships of the genera of the Iguanidae which represents the present Figure 3. Phylogeny and karyotype distribution in the family Iguanidae. The base dendrogram summarizes phylogenetic conclusions of Richard Etheridge from his osteological studies and is reproduced here with his permission from a privately circulated figure. The available karyotypic information for the family (references cited in Table 3) is superimposed on this dendrogram and indicates (1) the basic diploid karyotype for each genus chromosomed or (2) in karyotypically variable genera, the most primitive karyotype in the genus plus the range of 2n known for it. “V” chromosomes are metacentric macrochromosomes, “I? are acrocentric macro- chromosomes, and ‘“m” are microchromosomes. Thus, an 8V,81+24m karyotype has a 2n of 40 with eight metacentric macrochromosomes, eight acrocentric macrochremosomes, and 24 microchromosomes. 1976 CHROMOSOME EVOLUTION 15 Uracentron Ophryoessoides Phrynosaura Sceloporus Stenocercus Ctenoblepharis 2n: 22-46 Uma (inc. 12V+22m) l2V +22m Strobilurus Liolaemus Uta Holbrookia Platynotus Proctotretus 2n=30-40 I2V+22m I2V+22m (inc. l2V+ 22m) Plica BV,81*24m Galapagos Phymaturus Urosaurus Calica us Tropidurus 12V+22m y |2V+24m Leiocephalus eastern Tropidurus 2V+24m Rie ad i hrynosoma 12V+24m aw 12V+20m 2m l2v+22m western Tropidurus p, etrosaurus Uranoscodon Crotaphytus 12V +22m 12V+24m a Anolis Phenacosaurus coir, paleo 18. Nee AMERICA Sauromalus inc. l2V+24m Iv +24 SOUTH AMERICA, | SOUTH AMERICA ee AMERICA un CENTRAL AMERICA, ; © % : WEST INDIES BAnolis |= & Dipsosaurus 2n: 29-40 3 12V+24m MEXICO TO q Aptycholaemus SOUTH AMERICA, | © Iguana a Anisolepis CUBA, JAMAICA Buseon enosaura 12V+24m l2V+24m Chamaelinorops Cyclura l2V+24m i2V+24m Urostrophus HISPANIOLA Ambiveiunch NORTH AMERICA, Chamaeleolis MOIYrNyNCNUS — CENTRAL AMERICA, NORTHERN SOUTH lev+24 aiekeee eoreelts q AMERICA, WEST INDIES nt m Enyalius GALAPAGOS / Corythophanes Laemanctus 5 Brae lophus Basiliscus : \2V+24m SOUTH AMERICA Diplolaemus A CENTRAL AMERICA, CENTRAL AMERICA Leiosaurus NOUTHERNISGUTH Aperopristis AMERICA Pristidactylus Polychrus steal 2n: o&=29 % =30 Oplurus 9 =28 l2V+24m 9 =26 Hoplocercus 9=20 Morunasaurus | oplurines Chalarodon Enyalioides MADAGASCAR 12V+24m SOUTH AMERICA 16 BREVIORA No. 441 views of Richard Etheridge and Richard Estes, extending those of Savage (1958), Etheridge (1964) and Presch (1969). This has been based wholly on osteology and thus it gives us a picture of relationships constructed without knowledge of or reference to karyotypes. We have then superimposed on this dendrogram the range of karyotypes for each iguanid genus for which these are known. We use both the published reports summarized by Gorman, 1973 (most references cited by Gorman are not re- peated) and our still unpublished data. We use the latter despite the lack of formal (and especially pictorial) documenta- tion because it fills out much of the picture of karyotypic varia- tion in the Iguanidae without significantly altering it. The Etheridge scheme recognizes a basal stock (called “morunasaurines” by Estes and Price, 1973) from which seven major lineages arise (Polychrus, the Anolis-Enyalius-Diplolaemus lineage, the “tropidurines,”’ the “sceloporines” plus Crotaphytus, the “iguanines,” the “basiliscines,’ and the “oplurines’’). Karyotypes are known for one genus of morunasaurines (one species of Enyalioides — our unpublished data); in this and in six of the seven derived lineages (all except Polychrus) the “primitive” 12 + 24 karyotype is known to occur or is the only karyotype known. Furthermore, in every one of the derived lineages (except Polychrus) those genera closest to the base on Etheridge’s diagram — i.e. those believed for osteological reasons to be the more primitive members of each lineage (and, in fact, each sublineage) -—have either the 12 + 24 karyotype or a 12 + 22 karyotype that differs from the primitive condition by the absence (by loss or fusion) of a pair of microchromosomes. In the iguanine line, for example, all genera in the sublineage containing Iguana, Ctenosaura, Dipsosaurus and Sauromalus have a 12 + 24 or (Iguana) 12:+ 22 karyotype. The sceloporine line is especially instructive in this and other respects. Crotaphytus, an early offshoot of this line, has 12 + 24 (Cohen et al., 1967; Montanucci, 1970). Every primitive sceloporine and some Sceloporus have 12 + 22 (several authors, summarized in Hall, 1973). Within Sceloporus (a morphologi- cally more derived genus) numbers range from 12 + 10 to 24 + 22; if the 12 + 22 pattern is accepted as primitive for the genus, it is clear that both fusions and fissions must have been involved in the evolution of its karyotypic diversity. The tropidurine line is not well sampled, but as we have shown above, Tropidurus from eastern South America and the 1976 CHROMOSOME EVOLUTION ey Galapagoan Tropidurus (derived from western South American stock) have the basic 12 + 24 pattern, although their karyo- types are slightly derived in non-Robertsonian ways. The only West Indian Leiocephalus published (Gorman et al., 1967) also has the basic pattern. Besides confirming the 12 + 24 pattern in Hispaniolan species, Hall (unpub.) has found representatives of the Cuban branch of the genus to have 12 + 20 patterns. In the speciose genus Liolaemus, the single species so far re- ported (Gorman et al., 1967) has 12 + 22, although slides made by Richard Sage and examined by Hall indicate the presence of considerable karyotypic variation in this genus Cnt 30403) *. In the line leading to Anolis, the primitive genera Chamaeleo- lis, Phenacosaurus and Chamaelinorops (Hall and Williams, in preparation) have 12 + 24 karyotypes, as do many of the alpha section of the genus Anolis itself. The 12 + 24 pattern is also found in the related lineage including the genera Pristidactylus and Anisole pis. Given this evidence, it is difficult to contest the hypothesis that the 12 + 24 pattern is primitive in the Iguanidae. It is possible to go to greater detail: arguments similar to those above support the idea that even the detailed chromosome size and arm ratios found in Conolophus must be primitive for iguanids. Additionally, when we notice (1) that quite similar 12 + 24 karyotypes are found in other lizard families (Table 3), both in families that everyone agrees are closely related to the Iguani- dae, i.e. the Agamidae and the Chamaeleontidae, and in fam- ies that are just as universally regarded as not closely related (Gerrhosauridae, Anguidae, Amphisbaenidae); (2) that. still other families (Scincidae, Helodermatidae, Varanidae) have karyotypes easily derivable from 12V + 24m (Gorman, 1973), it then becomes clear that, using the most neutral descriptive terms, the 12 + 24 karyotype is an extraordinarily stable and conservative pattern. To us it is evident that the most careful and skeptical use of the Kluge and Farris criteria of primitive- tAlthough variation in chromosome number was clearly demonstrated in this material, the preparations were not of good quality and the data were complicated by the inclusion of unnamed taxa (clearly Liolaemus, however) . Further work will be required before publication is warranted. However, we think that the existence of substantial karyotypic variation in this genus should be noted. 18 BREVIORA No. 441 ness points unequivocally to the ancestral position of the 12 + 24 karyotype not only for iguanids, but for all lizards. On the other hand, there is no doubt that some lizard families and at least one group within the Iguanidae have karyotypes that are very difficult to reconcile with derivation from a 12 + 24 pattern (Gorman, 1973). The Gekkonidae will serve as an example of the first case; Polychrus is, of course, the second. We are impressed that these deviant families and groups are in general isolated cases, neither closely related to one another, nor arguable as ancestral to forms with the 12 + 24 karyotype. In all of these cases, morphological and other evidence suggests long separation from plausible basal stocks, and hence leads all the more strongly to the conclusion that the Conolophus-like 12 + 24 pattern is the one primitive for lizards. EVOLUTIONARY ROLE OF ROBERTSONIAN CHANGE (W. P. Hatt anno E. E. Witiiams) Let us point out immediately that there is no antithesis be- tween primitiveness and stability — the two explanations that have been proposed for the iteration of one chromosome pat- tern throughout a large number of species. On the contrary, the genera which show only the widespread chromosome pat- terns which we believe to be primitive seem not to be the ones that have radiated widely. This is a point which we want to stress, especially for the Iguanidae. Karyotypically conservative groups, so far as we can see, have produced only few species. Of the more than 50 genera of iguanids, only three — Anolis, Sceloporus and Liolaemus — are very large, each including more than 50 species (several times 50 in Anolis), or have produced high levels of sympatry (5+ syntopic species in several areas of the ranges of Anolis or Sceloporus). Of the others, only Steno- cerus (as revised by Fritts, 1974), Tropidurus (if the Galapagos species are included), and Leiocephalus (a purely insular radi- ation in the West Indies) have as many as 20 species, and no others have as many as 15. Excepting Polychrus, whose several karyotypes bear little obvious relationship to one another and none to any other iguanid, the other small, conservatively spe- clating genera show on current evidence little or no intrageneric variation in karyotypes, and indeed very little variation among genera. Of the 14 non-sceloporine small iguanid genera (in- 1976 CHROMOSOME EVOLUTION 19 cluding Crotaphytus) sampled, 11 have the 12 + 24 pattern. All of the primitive sceloporines (eight genera) and J/gwana among the iguanines have 12 + 22. Only Plica, aside from Polychrus, stands out in showing a notably different karyotype (16 + 24), and its modifications seem relatively simple (presumably fissions of four of the primitive metacentric macrochromosomes). Of the genera of middle size (20-29 species), the four sampled species of Tropidurus have again the 12 + 24 pattern but differ somewhat in arm ratios from the usual condition, and while some Leiocephalus have the 12 + 24 pattern, others have 12 + 20 (reduction in two pairs of microchromosomes — Hall, un- published). Stenocercus has not yet been sampled. Contrasting strongly with this picture of conservative specia- tion and karyotypic evolution in the small iguanid genera is a picture showing extensive, usually Robertsonian karyotypic vari- ation in each of the three prolifically speciose genera. In Anolis 2n’s range from 25 to 48 (Gorman, 1973; Hall, unpub.), in Sceloporus they range from 22 to 46 (Gorman, 1973; Hall, 1973), and in a few Liolaemus they range from 30 to 40 (Sage and Hall, unpublished). The apparent association of conservative speciation with con- servative karyotypic evolution, and prolific speciation with re- markable karyotypic diversity suggests the possibility of an evolutionarily important causal relationship between karyotypic differentiation and speciation. Though there are undoubtedly other possibilities and explanations that might be raised, it is this possibility of causal relationship that we here want to evalu- ate. We offer the following arguments to demonstrate that the relationship between speciation and karyotypic diversity is genuine. (1) Intrageneric variation in karyotypes. Since few small genera from six of the seven major iguanid lineages are repre- sented by karyotypes from more than one species, we must agree that we cannot safely compare the amounts of intrageneric vari- ation between small and large genera in these lineages. This defect, however, most certainly does not apply to the sceloporine lineage: all nine sceloporine genera and the related Crotaphytus are cytologically well known. Half or more of the species from each of these 10 genera have been karyotyped: 3/5 from Crota- phytus, 2/2 from Petrosaurus, 9/14 from Phrynosoma, 2/2 from Callisaurus, 3/3 from Uma, 2/3 from Holbrookia, 1/2 from Sator, 6/6 from Uta, 5/10 from Urosaurus, and 45+ / 20 BREVIORA No. 441 Table 4 Distribution of karyotypic variation in Iguanidae: interspecies comparisons (large genera except Sceloporus omitted). Other Other small All small scelo- iguanid iguanid 2n Sceloporus’ porines’ — genera genera 48 0) 0 0 0 46 2 0 0 0 44 2 0 0 0 42 I 0 0 0 40 3 0 1 Plica plica i 38 0 0 0) 0 36 0 13 13 34 18 33 I Sue a2 eS) 0 0 0 30 2 0 1 Polychrus 1 marmoratus 28 0 0 1 Polychrus 1 peruvianus 26 1 0) 1 Polychrus 1 femoralis 24 3 0 0 0 22 13 0) ) 0 20 0 0 1 Polychrus acutirostris species karyotyped 45,7 33 19 DZ species known 64 + a9 122 171 % karyotyped species with 2n—J010ro4) 42% 100% 74% 90% includes polymorphisms within currently recognized “species.” “here including Crotaphytus. 1976 CHROMOSOME EVOLUTION 21 64+ from Sceloporus (data summarized from ‘Table 2). None of these genera (except Sceloporus) shows any intrageneric vari- ation, and the only intergeneric difference is between the 12 + 24 Crotaphytus and the 12 + 22 sceloporines. Within Scelo- porus only 13 species (15 after taxonomic revisions by Hall) are known to have the primitive sceloporine condition (2n = 34), while the remaining 32 (40 or 41 after revisions) karyo- typed species have derived patterns — and most of these belong to the phylogenetically more advanced large-scaled branch as defined by Smith (1939). In the sceloporine lineage (Table 4), the correlation between chromosomal diversity and prolific spe- ciation is clear cut and does indeed appear to be fundamental. And even with our poor sampling of the small non-sceloporine genera, the association between chromosomal conservation and few species per genus is, at the least, suggestive. (2) Intergeneric diversity in karyotypes. Although compara- tively few species of the small, non-sceloporine genera have been karyotyped, still there is less intergeneric diversity observed than we would expect if variation were randomly distributed in the family. Phylogenetic relationships inferred from morphology (Fig. 3) show that many of these genera must have been evolv- ing as independent lineages for comparatively long times, possi- bly since the Cretaceous (Estes and Price, 1973). Given so long a period of evolution, they show remarkably little evidence of the acquisition or accumulation of chromosomal differentia- tion. As we have said, there are very few known differences among genera, and, in fact, few departures from the 12 + 24 pattern. In the 25 small genera sampled (Table 5), the few observed cases of intergeneric variation are slight indeed com- pared to the known intrageneric variation of the phylogenetically more recent large genera. Unless the sampling of the small genera has been biased in some unknown way, this should be quite significant. The deviations from the 12 + 24 pattern among the small genera are again: Plica (16 + 24 in no more than four muta- tional events, and possibly in only one, fide Todd, 1970), Iguana (12 + 22 in one event), all of the “primitive” scelo- porines (12 + 22 in one event in the common ancestry for all species), and Polychrus (2n’s = 20-30 resulting from an un- determined number of events producing karyotypes derived in relation both to one another and the 12 + 24 pattern). Con- trasted to the limited intergeneric variation in the family as a 22 BREVIORA No. 441 whole is the remarkable interspecific diversity involving many mutational events found within each of the three especially speciose genera (cf. Fig. 3). Again, this relationship is clearest in the well-investigated sceloporine lineage (Hall, 1973, in prep.). Table 5 Distribution of intergeneric comparisons in the Iguanidae small genera large genera NN NI N N ss a ac} tr ee ris iS by aS & s 2 6.8 % uF Pax 2 ob Pes oO CS MLN oO Sit GN ee pe a pee er lineage = eu 5 see basal stock 3 1 0) 0 = = oplurines 2 1 0 0 — = basiliscines 3 1] 0 0 = = iguanines 8 6 0) 0) = = sceloporines & Crotaphytus g) 9 0 1 1 1 tropidurines® 11 1 1 4 3 2 anolines 3 2 0 1 1 ] anoline relatives 8 3 0) 0 = = Polychrus 1 1 l 0 — = Totals eae) 8) 2 6 5 4 'The number of genera in the lineage for which karyotypic data exist. “The number of genera in the lineage containing one or more species with a karyotype deviating from the 12 + 24 or 12 + 22 pattern. 3Stenocercus, Tropidurus, Leiocephalus, and Liolaemus, each with 20 or more species, are treated as large genera. 1976 CHROMOSOME EVOLUTION eS (3) Relative recency of species and of karyotypic diversity. If rates of fixation of Robertsonian mutations were independent of the process of speciation, one expectation might be that many of the older genera would accumulate karyotypic variants while phylogenetically recent groups might show little variation, even though they include many species. Hall (1973) would adduce Sceloporus as a counter-example, since he believes it to be a phylogenetically quite recent genus. We summarize Hall’s views and arguments here. (A) While the Iguanidae may have originated in the Cre- taceous (Estes and Price, 1973), the differentiation of the present sceloporine genera probably did not antedate the de- velopment of the xeric habitats in North America during the Miocene (Axelrod, 1950, 1958). This conclusion is supported by the facts: (1) that Crotaphytus and all sceloporine genera are largely or entirely restricted to the North American deserts (except Sceloporus itself, which has extensively radiated in mesic habitats as well as in deserts); (2) that all sceloporine genera except for the osteologically primitive Crotaphytus and Petrosaurus (Etheridge, 1964; Presch, 1969) and Sator (which simply has not been tested) will ‘shimmy bury” (Axtell, 1956) in loose soil or sand for escape or sleeping cover (Stebbins, 1943, 1948; Axtell, 1956; Norris, 1958), a behavior not seen in any of the other North American or West Indian iguanids (we know nothing about South American iguanids in this re- spect); and (3) that all sceloporines but no other North Ameri- can iguanid genera (including Crotaphytus) show a “‘sink-trap” type of nasal passage and almost always an associated valvular nostril’ (Stebbins, 1943, 1948; Savage, 1958; Hall, unpub.), which clearly seems to have been evolved in relationship to the use of shimmy burial for cover in xeric environments where loose soil is likely to be more readily available for cover than that provided by plants or permanent burrows in firm soil. (B) Within the sceloporines, Sceloporus seems to be one of the most recently differentiated genera. Osteological data clearly show four groups within the sceloporines (Savage, 1958; Ethe- ridge, 1964; Presch, 1969): the primitive Petrosaurus; the specialized Phrynosoma; the group of ‘“‘sand-swimming” (Norris, 1958) genera, Callisaurus, Holbrookia and Uma; and the group 1Petrosaurus, which lives in xeric habitats but which does not shimmy bury under experimental conditions, has the nasal sink trap but lacks the nasal valve (Hall, personal observation) . 24 BREVIORA No. 441 of four genera, Sceloporus, Urosaurus, Sator and Uta. Accord- ing to Etheridge (1964), these last four genera cannot be dis- tinguished osteologically. However, comparisons suggest that Sceloporus, with its mucronate imbricate scales developed to a degree not found in any of the other North American iguanids (where granular scales seem to be the primitive condition) and with its loss of the gular fold found in all other sceloporines, is the phylogenetically most recent of these four genera. (C) Within Sceloporus, again based on characteristics of the squamation, it seems clear that Smith’s (1939) small-sized, small-scaled species are more closely related to the other sceloporines than is the large-sized, large-scaled branch. To summarize the cytosystematics of these two divisions, the phylo- genetically more primitive small-scaled branch contains 20 spe- cies by present taxonomy: 12 of these are karyotypically con- servative,' five have not been karyotyped, and the three that are karyotypically derived are also highly derived ecologically (merriami, 2n = 46, is a specialized cliff-face dweller and scalaris and aeneus, 2n = 24, are specialized montane bunch grass dwellers). Within the morphologically more advanced large-scale branch, according to present taxonomy, only the single species, S$. orcutti, has the primitive 12 + 22 pattern, while all other karyotyped species are derived. At present (very conservative taxonomy) this branch contains a minimum of 43 species, of which all but 13 (all 13 are in chromosomally highly derived species groups) have been karyotyped. This phylogenetically most recent radiation of the sceloporines (the large-scaled Sceloporus) has covered the entire ecological and geographical range of lizards in North America (Smith, 1939) and shows simultaneously a truly remarkable karyotypic diversity (2n’s from 22 to 46). It is also notable that the most ecologically differentiated small-scaled species in the genus (the scalaris group species [2n = 24] and the merriami [2n = 46]) are also among the karyotypically most highly derived forms. The one egregious example of chromosomal diversity in a small genus, Polychrus, seems in the very fact of its uniqueness equally a counter-example to the generality of the proposition that deviant karyotypes tend to accumulate in all genera with time. 1Karyotypically at least to a first approximation: Cole (1971) notes that maculosus has a 2n —= 31, X,X,Y4 (based on three specimens, only one a male) and that in pyrocephalus chromosome | shows a pericentric inversion. 1976 CHROMOSOME EVOLUTION D5 Polychrus is seen on Etheridge’s diagram as an isolated basal twig, truly very old and very distinct, entirely suitable as a group in which deviant karyotypes might accumulate. But each of the other six major groups is as old in Etheridgean terms. If karyotypic diversity is a product only of time, even a random and superficial sampling of the other small genera should, so it seems to us, have resulted in more cases of highly derived karyo- types than are in fact in front of us. Our own surmise regarding Polychrus is that the six forms currently recognized, all highly arboreal, may represent only the few survivors of an old and formerly more prolific lineage of tree dwellers that, perhaps, has been largely replaced by the radiation of Anolis in the arboreal habitat. CHROMOSOMAL EVOLUTION IN THE IGUANIDAE: TWO MAJOR PATTERNS We do not deny that the history of karyotypic change is not now and never will be known from direct evidence, that the real and unique historical process must be inferred from its products, nor that the survey of iguanid karyotypes, though it is already impressive, is incomplete. We insist, however, that the present sample is large enough to justify conjecture and to point to the kinds of evidence that will verify or negate postu- lated sequences. Our picture of chromosomal evolution in lizards, and _ per- haps also in many other groups, is that there are both periods of chromosome conservatism with usually slow geographic modes of speciation (Mayr, 1963) and episodes of karyotypic instability associated with rapid proliferations of new species (Hall, 1973). We believe that both Anolis and Sceloporus exhibit these phe- nomena (cf. especially Webster et al., 1972; Williams and Web- ster, 1973; and Hall and Selander, 1973), and presumably Liolaemus also does. Todd (1970) suggests a similar relation- ship between karyotypic diversification ‘and prolific speciation in the Canidae. To us the comparative data strongly suggest that karyotypic diversification and speciation are in many cases functionally related, such that the temporal and/or geographic requirement for the separation of populations is somehow minimized (not eliminated but very greatly reduced) when chromosomal differ- ences become fixed between them. White’s model of “stasi- 26 BREVIORA No. 441 patric” speciation (White et al., 1967; White, 1968; Key, 1968) offers one mechanism, and others are possible (Hall, 19733 im prep.). Here we wish to emphasize only that Robertsonian mutations frequently are found fixed between species of rapidly proliferating groups but only rarely are found as intrapopula- tion poly morphisms. (Wallace, 1959, provided an early note of this phenomenon in Drosophila.) Given this distribution of Robertsonian mutations, we think it especially significant that, among the varieties of chromosomal rearrangements, the Robert- sonian ones probably have the least impact on the meiotic as- sortment or recombination of balanced genomes; but, on the other hand, at least in mammals where breeding and cytological studies have been made, these mutations are increasingly impli- cated as a significant source of chromosomal malassortment in meiosis serving to reduce the effective fertility of chromosomally heterozygous individuals ( Polani et al., 1965; Gustavsson, 1971a, b; Cattanach and Mosely, 1973). Once a ‘chromosennl differ- ence is established, reduced heterozygote fertility could then serve in appropriate circumstances as an intrinsic partial barrier to gene flow between karyotypically differentiated homozygous populations, thereby reducing the requirement for extrinsic bar- riers to gene flow before speciation could ensue (Hall, 1973; in Preps). Then, assuming some model of chromosomal speciation based on cytogenetically reduced fertility in heterozygotes, the proba- bility or frequency of such speciation in given lineages should be highly dependent on parameters of their genetic systems such as: mutation rates, malassortment rates, population structures, mating systems, ets. Chromosomal speciation might then be precluded in some lineages because of unfavorable genetic sys- tems that would allow speciation only by conservative geographic modes; on the other hand, genetic systems of other lineages may especially favor chromocomal speciation, and thus allow great proliferations of species, even in the absence of strong extrinsic barriers to gene flow. Such a chromosomal speciation theory can easily account for the associations of karyotypic diversity and prolific speciation found in Sceloporus, Anolis, and apparently in Liolaemus. The test of the chromosomal speciation model of karyotypic evolution as it pertains to the iguanids will be found in the still unsampled or inadequately sampled iguanid radiations, par- ticularly those of South America. Stenocercus, now with 29 1976 CHROMOSOME EVOLUTION Zi recognized species and with notable sympatry, is certainly cru- cial. The karyotypic variation in Liolaemus and Leiocephalus must be confirmed, and the karyotypic patterns in these two genera adequately documented. We suggest that the species of mainland Tropidurus, which seem to have rather complicated distributions (Peters and Donoso-Barros, 1970), may also repay careful attention. Only such a wider survey of the karyotypes of the Iguanidae can provide either a verification of the evolu- tionary patterns we have suggested here, or, by demonstrating new patterns, require alternative models. RESUMEN Los iguanidos: Conolophus subcristatus, Tropidurus albemar- lensis, T. delanonis, y T. duncanensis tienen cariotipos similares de 36 cromosomas. C. subcristatus, con 12 macrocromosomas metacéntricos y 24 microcromosomas, tiene un cariotipo que se demuenstra ser “primitivo” dentro la familia Iguanidae, y que probablemente es también primitivo entre todos de los lagartijos. Los cariotipos 2n = 36 de los tres Tropidurus son iguales y también al Tropidurus del este de Suramérica, pero ellos son diferentes del cariotipo primitivo porque hay modificaciones “no-Robertsonianas” de tres pares de los macrocromosomas. La filogénia y todos de los datos cromosémicos de la Iguani- dae estan resumidos para una discusion sobre la determinacion de que cariotipos son “primitivos” dentro radiaciones de espe- cies, y también sobre los funciones que sirven las mutaciones de Robertson en el proceso de evolucién. Analisis de la informacion sobre los cariotipos y sistematica demuenstra una conexion causal y cerca entre la diferenciacién cariotipica y la prolifera- cion rapida de especies nuevas, donde el requisito para ais!lami- ento geografico se minimiza a causa de la diferenciacidn cromo- somica. ACKNOWLEDGMENTS We are grateful to Captain Sir Thomas Barlow and Mr. Roger Perry for permission to collect on the Galapagos Islands and for the opportunity to use the facilities of the Charles Dar- win Research Station, and to Professor James J. Hoff and Richard C. Paull for assistance in the field. We thank R. B. Stamm for his assistance in the lab and T. P. Webster, Richard Etheridge, Richard Estes and James J. Jackson for comments 28 BREVIORA No. 441 and criticisms. Richard Etheridge generously provided the dendrogram of iguanid relationships and has amplified or an- notated our statements on generic size in Table 2. The research was partially supported by NSF grants GB 19801X and GB 37731 to E. E. Williams and GB 27911 (1969-70) to R. Rol- lins of the Committee on Evolutionary Biology at Harvard and NIH grant RR—8102 administered by the Division of Research Resources. REFERENCES ARPONET, V. N. 1965. Description of the karyotypes of Agama caucasica and Phrynocephalus helioscapus (Agamidae, Reptilia) [in Russian, English summary]. Tsitologiya 1: 237-239. AXELRoD, D. I. 1950. Evolution of desert vegetation in western North America. Carnegie Inst. Wash., Publ. 590: 215-360. 1958. Evolution of the Madro-Tertiary geoflora. Bot. Rev. 24: 433-509. AXTELL, R. W. 1956. A solution to the long neglected Holbrookia lacerata problem and the description of two new subspecies of Holbrookia. Bull. Chicago Acad. Sci. 10: 163-179. Baker, R. J., J. H. Bowrrs, AND M. H. Situ. 1975. Reply to comment on “Chromosomal evolution in Peromyscus.’ Evolution 29: 189. Barrio, A. 1969. Sobre la real ubicacion generica de Leiosaurus fasciatus DOrbigny (Lacertilia, Iguanidae). Physis 29: 268-270. Becak, M. L., W. Brecak, AND L. Denaro. 1972. Chromosome polymorphism, geographical variation and karyotypes in Sauria. Caryologia 25: 313-326, Brancut, N. O. AND J. R. Contreras. 1967. The chromosomes of the field mouse Akodon azarae (Cricctidae, Rodentia) with special reference to sex chromosome anomalies. Cytogenetics 6: 306-313. Bury, R. B., G. C. GorMAN, AND J. F. LyNow. 1969. Karyotypic data for five species of anguid lizards. Experentia 25: 314-316. CATTANACH, B. M. AND H. Mosery. 1973. Nondisjunction and reduced fertility caused by the tobacco mouse metacentric chromosomes. Cyto- genet. Cell Genet. 12: 264-287. CoHEN, M. M., C. C. HuAnc, AND H. F. Crark. 1967. The somatic chromo- somes of three lizard species, Gekko gecko, Iguana iguana, and Cro- taphytus collaris. Experentia 23: 769-771. Coir, C. J. 1970. Karyotypes and evolution of the spinosus group of lizards in the genus Sceloporus. Amer. Mus. Novitates No, 2431: 1-47. 1971a. Karyotypes of the five monotypic species groups of lizards in the genus Sceloporus. Amer. Mus. Novitates No. 2450: 1-17. 1971b. Karyotypes and relationships of the pyrocephalus group of lizards in the genus Sceloporus. Herpetologica 27: 1-8. Estrs, R. AND L. I. Price. 1973. Iguanid lizard from the Upper Cretaceous of Brazil. Science 180: 748-751. 1976 CHROMOSOME EVOLUTION 29 ETHERIDGE, R. 1964. The skeletal morphology and systematic relationships of sceloporine lizards. Copeia 1964: 610-631. GorMaNn, G. C. 1970. Chromosomes and the systematics of the family Teiidae (Sauria, Reptilia). Copeia 1970: 230-245. 1973. The chromosomes of the Reptilia, a cytotaxonomic interpretation. Jn A. B. Chiarelli and E. Capanna eds., Cytotaxonomy and Vertebrate Evolution, pp. 349-424. Academic Press, New York. ——— ——., L. ATKINs, AND T. Horzincer. 1967. New karyotypic data on 15 genera of lizards in the family Iguanidae with a discussion of cytological and taxonomic information. Cytogenetics 6: 286-299. AND D. SHOCHAT. 1972. A taxonomic interpretation of chromo- somal and electrophoretic data on the agamid lizards of Israel with notes on some East African species. Herpetologica 28: 106-112. Gustavsson, I. 197la. Chromosomes of repeat-breeder heifers. Hereditas 68: 331-332. 1971b. Distribution of the 1/29 translocation in the A. I. bull population of Swedish Red and White cattle. Hereditas 69: 101- 106. Hartt, W. P. 1970. Three probable cases of parthenogenesis in lizards (Agamidae, Chamaeleontidae, Gekkonidae). Experentia 26: 1271-1273. Comparative population cytogenetics, speciation, and _ evolu- tion of the iguanid genus Sceloporus. Unpublished Ph.D. thesis, Har- vard University. AND R. K. SELANDER. 1973. Hybridization of karyotypically differentiated populations in the Sceloporus grammicus complex (Iguani- dae). Evolution 26: 226-242. Hsu, T. C. ano J. L. Patton. 1969. Bone marrow preparations for chromo- some studies. Jn K. Benirschke ed., Comparative Mammalian Cyto- genetics. pp. 454-466. Springer-Verlag, New York. Huane, C. C., H. F. Crark, AND C. Gans. 1967. Karyological studies on fifteen forms of amphisbaenians (Amphisbaenia-Reptilia). Chromo- soma 22: I-15. Kry, K. H. L. 1968. The concept of stasipatric speciation. Syst. Zool. 17: aS 22. KiucE, A. G. AND J. S. Farris. 1969. Quantitative phyletics and the evolu- tion of anurans. Syst. Zool. 18: 1-32. Lawtor, T. E. 1974. Chromosomal evolution in Peromyscus. Evolution 28: 688-691. MaATTHEY, R. 1931. Chromosomes des Reptiles, Sauriens, Ophidiens, Che- loniens. La evolution de la formule chromosomiale chez les Sauriens. Rev. Suisse Zool. 38: 117-186. 1933. Nouvelle contribution a l’etude des chromosomes chez les Sauriens. Rev. Suisse Zool. 40: 281-316. 1949. Les Chromosomes des Vertebres. 353 pp. F. Rouge, Lausanne. 30 BREVIORA No. 441 Mayr, k, 1963, Animal Species and Evolution. 797 pp. Harvard Belknap Press, Cambridge, Mass. Montanuccl, R. R. 1970. Analysis of hybridization between Crotaphytus wislizenii and Crotaphytus silus (Sauria, Iguanidae) in California. Copeia 1970: 104-123. MorescatcH!, A. 1973. Amphibia. Jn A. B. Chiarelli and E. Capanna, eds., Cytotaxonomy and Vertebrate Evolution, pp. 233-248. Academic Press, New York. Norris, K. S. 1958. The evolution and systematics of the iguanid genus Uma and its relation to the evolution of other North American desert reptiles. Bull. Amer. Mus. Nat. Hist. 114: 247-326. PeccinIni, D. M. V. M. 1969. Caridtipo e mecanismo de determinacao de sexo em algumas espécies de lacertilios Brasileros (Iguanidae e Teiidae) . Master’s ‘Thesis, Universidad de Sao Paulo, 46 pp. PENNOCK, L. A., D. W. TINKLE, AND M. W. SHAW. 1969. Minute Y chromo- some in the lizard genus Uta (Family Iguanidae). Cytogenetics 8: 9-19. PETERS, J. A. AND R. Donoso-Barros. 1970. Catalogue of the neotropical Squamata. Part II. Lizards and amphisbaenians. 293 pp. Smithsonian Institution, Washington, D. C. PoLani, P. E., J. L. HAMERTON, F. GIANNELLI, AND C. O. CARTER. 1965. Cytogenetics of Down’s Syndrome (Mongolism). II. Frequency of interchange trisomics and mutation rate of chromosome interchanges. Cytogenetics 4: 193-206. PrescH, W. 1969. Evolutionary osteology and relationships of the horned lizard genus Phrynosoma (Family Iguanidae). Copeia 1969: 250-275. RiEGER, R., A. MICHAELIS AND M. M. GREEN. 1969. A Glossary of Genetics and Cytogenetics: Classical and Molecular. 3rd ed., Revised. Springer- Verlag, New York. 507 pp. Ropinson, M. D. 1974. Chromosomes of the insular species of the chuck- walla lizards (genus Sauromalus) in the Gulf of California, Mexico. Herpetologica 30: 162-167. Savace, J. M. 1958. The iguanid lizard genera Urosaurus and Uta with remarks on related groups. Zoologica 43: 41-54. SmMITH, H. M, 1939. The Mexican and Central American lizards of the genus Sceloporus. Field Mus. Nat. Hist., Zool. Ser. 26: 1-397. SoKoLovsky, V. V. 1972. Comparative karyology of the reptiles [7m Rus- sian]. Viniti— All Union Institute of Scientific and Technical Informa- tion, Moscow. 51 pp. Strppins, R. C. 1943. Adaptations in the nasal passages for sand burrow- ing in the saurian genus Uma. Amer. Nat. 77: 38-52. --———_—._ 1948. Nasal structure in lizards with reference to olfaction and conditioning of the inspired air. Amer. J. Anat. 82: 183-222. Topp, N. B. 1970. Karyotypic fissioning and canid phylogeny. J. Theor. Biol. 26: 445-480. WaALLAce, B. 1959. Influence of genetic systems on geographical distribu- tion Cold Spring Harbor Symp. Quant. Biol. 24: 193-204. 1976 CHROMOSOME EVOLUTION 31 WEsstTER, T. P.. W. P. HALL, aND E. E. WILLIAMS. 1972. Fission in the evolution of a lizard karyotype. Science 177: 611-613. Waite, M. Jj. D. 1954. Animal Cytology and Evolution, 2nd Ed. Cam- bridge Univ. Press. —. 1959. Speciation in Animals. Australian J. Sci. 22: 32-39. 1968. Models of speciation. Science 159: 1065-1070. 1973. Animal Cytology and Evolution, 3rd Ed. Cambridge Univ. Press. 961 pp. » R. E. BLAckiTH, R. M. BLACKITH, AND J. CHENEY. 1967. Cytogenetics of the viatica group of morabine grasshoppers. I. The “coastal” species. Australian J. Zool. 15: 263-302. WituiAMs, E. E. and T. P. Wesster. 1974. Anolis rupinae, new species, a syntopic sibling of A. monticola Shreve. Breviora Mus. Comp. Zool. 429 N22. NOTE ADDED IN PRESS Since we wrote the above, Vegni Talluri et al. (1975) pub- lished karyotypes identical to those Tropidurus karyotypes given here for the additional two species, T. jacobi (James Bay, James Id.) and T. indefatigabilis (Academy Bay, Indefatigable Id.) ; and for two additional populations of T. albemarlensis (Vil- lamil, Albemarle Id. and Punta Espinoza, Narborough Id.). These data further support our conclusion that chromosomal differentiation plays no functional role in classical geographic speciation and add contrast to the situations of frequent asso- ciation between the fixation of chromosomal differences and speciation which does not involve obvious geographic separa- tion. ADDITIONAL REFERENCE VecNI Tatturi, M., R. DALLAI, AND B. Lanza. 1975. The karyotype of some Tropidurus (Reptilia, Iguanidae) from the Galapagos Islands. [published reprint from: Galapagos, Studi e Ricerche — Spedizione “L. Mares—G.R.S.T.S.”].. 9pp. Gruppo Ricerche Scientifiche e Tecniche Subacquee, Firenze (Italy) . MUS. COMP ZOOL LIBRARY BREV TOUR A t} Ni! VERSITY Museum of Comparative Zoology US_ISSN_0006-9698 CAMBRIDGE, MASS. 27 MAY 1977 NUMBER 442 A LABORATORY STUDY OF THE TURKISH HAMSTER 4 MESOCRICETUS BRANDTI CHARLES P. LYMAN! AND REGINA C. O’BRIEN! Apstract. Ten years of laboratory observations of a breeding colony of Meso- cricetus from central Turkey are summarized. From morphological and karyological evidence, as well as cross-breeding experiments, it is concluded that Mesocricetus brandti is distributed in specific areas throughout Asiatic Turkey, south into Pales- tine, Syria, the northern part of Iraq, northwest Iran, and over the Caucasus into Daghestan. The chromosomal polymorphism noted in this species may be due to the isolation of groups to these specific types of terrain. The reproductive cycle, growth, and care and behavior in the laboratory are de- scribed. Records of hibernation are detailed and compared with those of Mesocri- cetus auratus. It is emphasized that M. brandti offers a unique opportunity to study the factors which influence hibernation using matched animals of known age and lineage. INTRODUCTION The study of hibernation in mammals has long been hampered by the lack of an animal which could be bred and raised readily under laboratory conditions. Ground squirrels, marmots, dormice, Euro- pean hamsters, hedgehogs and microchiropteran bats have been used extensively as experimental animals, but all of these species are usually obtained by collecting them in the wild. The Syrian hamster is easily raised in the laboratory and has been used as an experimental animal for more than forty years. How- ever, hibernation in this species is extremely unpredictable, and many individuals fail to hibernate even when exposed to cold for many months (Lyman, 1948, 1954). ‘Department of Anatomy, Harvard Medical School Boston, Massachusetts, 02115 and Museum of Comparative Zoology Harvard University, Cambridge Massachusetts 02138 2. BREVIORA No. 442 In August, 1965, we obtained 13 hamsters trapped in central Turkey.! These animals hibernated readily when exposed to cold (5 + 2°C) the following winter and bred in the laboratory during the next spring. Since that time, the colony has been continued, detailed notes have been maintained, and various experiments have been undertaken. In 1971, we received an additional 29 animals. We maintained and bred these animals separately from the original group for comparative data. The two groups resembled each other closely, and clearly differed from laboratory specimens of the Syrian or golden hamster, Meso- cricetus auratus. As information concerning the Turkish hamsters accumulated, it became apparent that this species is the best experi- mental animal for the study of certain aspects of hibernation, par- ticularly those which require animals of known history and lineage. Offspring of the two original colonies have been distributed to sev- eral investigators and we hope that the Turkish hamster is now established as an experimental animal. For this reason, our knowl- edge concerning this hamster is detailed below. Identification is, of course, of critical importance in animals used for experimental studies. However, because of the limited amount of comparative material available, comparisons of skins and skulls have led to conflicting opinions on the taxonomy of Mesocricetus (Ognev and Heptner, 1927; Aharoni, 1932; Ellerman, 1941, 1948; Vinogradov and Argiropulo, 1941; Ellerman and Morrison-Scott, 1951; Vereshchagin, 1959; Hamar and Schutowa, 1966; Ivanov, 1969; Raicu et al., 1969; Vorontsov and Krjukova, 1969; Todd et al., 1972). The genus exhibits chromosomal polymorphism while show- ing relatively few morphological differences over a wide geographic area. Emphasis by some investigators on the analysis of karyotypes has led to divergent taxonomic views which can not be fully recon- ciled until more definitive data such as chromosomal banding studies and cross-breeding experiments in the laboratory have accumulated. For this reason we have limited ourselves to a consideration of the identity of the Turkish animals and have attempted to delineate their distribution in their natural habitat. !'We are grateful to Dr. Bahtiye Mursaloglu of the University of Ankara, Ankara, Turkey for making arrangements to obtain the hamsters, and to Mr. Haluk Anat of the same university for trapping the hamsters and supplying us with information con- cerning their natural history. 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 3 TAXONOMIC MATERIALS The animals which formed our original colony in 1965 were cap- tured by Mr. Haluk Anat in Malya, near Kirsehir (39°N, 34°E)’ about 125 kilometers (km) southeast of Ankara, Turkey (Fig. 1). Mr. Anat trapped the second group six years later near the city of Ankara. We refer to these two groups collectively as “Turkish” hamsters in this text. Skins and skulls of hamsters from Malya which had been trapped alive and kept in captivity for varying periods of time were used for comparison with the specimens of Mesocricetus in the British Mu- seum. As far as we have been able to determine, the British Mu- seum collection is the only representative collection of this genus in the United Kingdom or the United States. It contains specimens from a now defunct colony in the Wistar Institute which were ob- tained from Pirbadan [m] (35°N, 48°E), Iranian Kurdistan, a small village 125 km north-northwest of Hamadan, Iran (Fig. 1). Hamsters from this colony have been the subject of karyological and breeding studies and have been referred to as “Kurdistan” or “K” hamsters* (Lehman and Macpherson, 1967; Palm, Silvers and Billingham, 1967; Todd et al., 1972; Raicu et al., 1972). We observed four hamsters originating from a colony of Ru- manian hamsters maintained by Dr. Petre Raicu, University of Budapest’, alive in our laboratory. We used two skins and skulls from the Rumanian colony to compare with the specimens in the British Museum. In addition, hamsters trapped for Dr. Michael Murphy’ in Aleppo, Syria were maintained in our laboratory for comparative studies and breeding experiments. Breeding crosses were usually attempted by placing the female in the cage of the male, though occasionally the reverse process was used. In many instances we used vaginal smears to determine the estrous cycle, and exposed the female to the male in the afternoon *The report that these animals came from Aksehir, Turkey (Todd et al., 1972) is an error derived from faulty original information. 3We thank Drs. J. E. Hilland G. B. Corbet of the British Museum for making these specimens available *All place names in this paper are those given in the original articles. ‘We are grateful to Dr. William Nixon of Randolph, Mass. for supplying these animals. ‘Dr. Murphy made a trip to Aleppo to obtain this colony and observe their be- havior. We acknowledge his gift with thanks. 4 BREVIORA No. 442 Figure 1. Map showing distribution of Mesocricetus. prior to estrus. If the animals fought, they were separated tem- porarily, and pairing was attempted at later dates. Otherwise, the animals were left together for three or four days and then separated. The animals were observed for copulatory behavior and vaginal smears were often made if copulation was suspected but not ob- served. We prepared karyotypes from metaphase spreads obtained from normally dividing primary skin cultures. The cells were harvested on the seventh to ninth day of culture. After exposure to colchicine (final dilution 36 micrograms per milliliter of medium) for five hours, and hypotonic solution for 10-15 minutes, cells were fixed in 3:1 methanol:glacial acetic acid and stained with Wright-Giemsa stain. Chromosomes were counted in at least 100 cells per animal. Se- lected metaphase spreads were photographed and homologous chromosomes were paired in the conventional manner according to their length and position of the centromere. DISCUSSION OF TAXONOMY It is generally agreed that the distribution of the genus Meso- cricetus includes the eastern parts of Rumania and Bulgaria, across Anatolian Turkey into the Caucasus, and south into Syria, the northern tip of Iraq and the northwest part of Iran. Hamar and Schutowa (1966) have presented the most complete taxonomic investigation of the genus in recent time. They had 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 5 available for comparison 102 specimens from the Moscow and Leningrad Zoology Museums as well as the Humbolt Museum in Berlin. Based on morphological data, they recognized three species of one group. They proposed that Mesocricetus newtoni was found in eastern Rumania and Bulgaria and that Mesocricetus auratus was confined to the type locality of Aleppo, Syria. Mesocricetus brandti appeared to occupy the rest of the range of Mesocricetus except for an area north of the Caucasus which was occupied by Mesocricetus raddei and M. r. nigriculus. Based on cranial measurements, they found little difference be- tween newroni, auratus and brandti, but the two forms of raddei were distinctly different, with relatively massive skulls and larger molar teeth. The bodies of animals of the raddei group were also larger in size, and the black color of their bellies contrasted with the grey or white of the smaller newroni-auratus-brandti group. Our examination of specimens of raddei from the Caucasus in the British Museum and the Museum of Comparative Zoology con- firmed the observations of Hamar and Schutowa (1966). These authors summarized their findings by suggesting that raddei should be separated from the other group at the subgeneric level, and the evidence presented for this view is convincing. When compared to specimens from Rumania or Turkey, the dor- sal pelage of the Syrian or golden hamster is notably different. The color of the latter is a rich Sanford’s brown (Ridgway, 1912) while our animals from Turkey are tawny-olive to Saccardo’s umber dor- sally, with an overlay of black-tipped guard and pile hair which darkens the pelage. Although Hamar and Schutowa do not empha- size this, our animals from Rumania were darker dorsally than our animals from central Turkey, as were several of the seven Rumanian specimens in the British Museum. The subauricular stripe in the Syrian animals is poorly defined, being a mixture of brown and black hairs, while in the Turkish and Rumanian animals it is pure black. There is a difference in the coloring of the chest, for in the Syrian hamster it is brownish with a narrow white mid-stripe, while in the Turkish and Rumanian animals there is a pronounced bar of black, as illustrated by Hamar and Schutowa (1966). The Kurdistan or “K” specimens were paler on the dorsal surface but otherwise closely re- sembled our animals, which were trapped approximately 1350 km to the west-northwest. It is of incidental interest that the pelage of the type specimen of M. auratus in the British Museum is indistinguishable from speci- 6 BREVIORA No. 442 mens which Dr. Murphy collected in Syria in 1971 and these in turn match the common laboratory Syrian hamster. Thus, laboratory breeding since 1930, when the first animals were captured (Adler, 1948), has not altered the natura! coat color of this animal, though many mutations in coat color have occurred. More than one thousand Turkish hamsters have been raised in our laboratory, and slight differences in the pelage have been observed. Some individuals have darker dorsal pelage than others, and the ventral surface may vary from grey-white to grey, particularly on the abdomen. There are no consistent differences in color between the colony which originated from Malya and the colony from An- kara. The four living animals from Rumania were about the same size as laboratory strains of Syrian hamsters, but our Turkish hamsters were bigger as full-grown adults. Twenty-one two-year-old Turkish hamsters chosen at random averaged 163 grams (g) with a range of 137 to 258 g. In comparison, 111 two-year-old Syrian hamsters averaged 105 g with a range of 97 to 113 g (Altman and Dittmer, 1964). The heads of the living animals from Rumania differed from the others, for the face was more pointed and ratlike, though the nasal portion of the skulls of Rumanian animals is not narrower. Laboratory interbreeding experiments have been detailed by Todd et al. (1972) and can be briefly summarized. No offspring have re- sulted from atings between our animals from Malya or Ankara in Turkey. | laboratory animals originating in Syria (Todd et.al., 1972; presen study), though copulation has been observed (Mur- phy, persc al communication; present study). Attempts to cross Kurdistan 21imals with laboratory Syrian hamsters have been simi- larly unsuccessful (Palm et al., 1967). The cross between Rumanian female and Malya male hamsters has produced viable offspring, but the reciprocal cross has not been attempted. Todd et al. (1972) report the survival of five females, with a male living only two weeks. In our laboratory, a female killed her first litter and two females survived in the second litter, both of which had developed diabetes mellitus by the age of nine weeks. Diabetes was not the invariable result of this cross, however, for we tested a female raised by Todd et al. and found the urine negative for glucose. The hybrids were larger than either parent, with one weighing 293 gasan. ult. 7 d eial. report copulation between a hybrid Rumanian-Turkis! female and 2 Malya male, but presented no evidence that the hybrid was fertile. 1977 LABORATORY STUDY OF THE TURKISH HAMSTER I Viable hybrids have been produced in the cross between female hamsters from Kurdistan and males from Rumania, but the recip- rocal cross was unsuccessful (Raicu et al., 1972). (Dr. M. Bah- manyar of the Pasteur Institute of Iran collected both Dr. Raicu’s and the Wistar Institute hamsters. He writes that the former ani- mals were trapped 25 km west of Pirbadan.) The number of young per litter was small and averaged about 50 per cent of the number expected in non-hybrid litters. Histological examination revealed that both fhe male and female reproductive organs of the adult hybrids were atrophic, and on this basis both sexes were consid- ered to be totally sterile. Raicu and Bratosin (1968) and Raicu et al. (1969) have obtained reciprocal crosses between Syrian and Rumanian hamsters. Though the litters were small, the individual animals were large and the gonads of the hybrids appeared histologically atrophic, leading to the conclusion that the animals were sterile. Todd et al. (1972) found that the hybrid females resulting from crosses between Rumanian males and Syrian females would mate with Syrian males. No viable young were produced, though reabsorbing embryos were found at autopsy in three cases. In contrast, there was no indication of lack of fertility between the two groups of animals from Turkey. Reciprocal crosses in our laboratory between hamsters originating in the Ankara and Malya areas result in normal healthy litters and the F; generations are fer- tile, producing normal young. The karyotypes of the various groups of hamsters, including hamsters in our laboratory which onginated in Malya, have been reported by Todd et al. (1972). Since that time we have established the karyotype of hamsters from the area of Ankara. The latter animals have a diploid number (2N) of 44 chromosomes with a fundamental number of autosomal arms (FN) of 80 (Fig. 2), while in the hamsters from Malya 2N = 42 and FN = 78 (Fig. 3). Hybrids of both sexes were examined and in these 2N = 43 and FN = 79. In the animals from Ankara, the autosomal complement is com- posed of two pairs of metacentric chromosomes (#3 and #13), two pairs of acrocentric chromosomes (#19 and #21) and the remaining 16 pairs, which are a graduated series of submetacentric and subte- locentric chromosomes. Because of the similarity and gradation in size of the majority of the chromosomes, it was not possible to identify with complete certainty many individual chromosomes, including X and Y. ia 8 BREVIORA No. 442 bh Gh 8 BR Bt X Y Sf Sh KS Ba BA ux 6 i K 8 AR ae na Figure 2. Representative karyotype of Mesocricetus brandti from the area of Ankara, Turkey. Autosomes are arranged sequentially from upper left to lower right. In comparing the karyotype of the Ankara hamsters with that ot the Malya group the same difficulty obtained, yet it is obvious that the difference in diploid number between the two groups is the result of the absence of the acrocentric pair #19 in the Malya hamsters leaving only one pair of acrocentric chromosomes. The hybrid ani- mals have three acrocentric chromosomes, two small ones and one slightly larger. The X chromosome of the Malya hamster is unmis- takably larger than that of the hamster from Ankara. Comparison of the karyotype of the Ankara hamster with that of the Kurdistan hamster (Lehman and Macpherson, 1967) reveals certain differences. In the Kurdistan hamster 2N = 42 and FN = 80. The smallest autosomal pair appears similar to #20 in the Ankara karyotype and there are no acrocentric elements. The Y chromo- 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 9 s AB AL ER OUR AE WA O&K ORR OGRA OMA nh OAR BR ORK OBA AA 8X 83) be a8 Figure 3. Representative karyotype of Mesocricetus brandti from Malya, Turkey. Autosomes are arranged sequentially from upper left to lower right. some of the Kurdistan hamster is obviously much larger than that of the Ankara animals. From the evidence which has accumulated to date, a few conclu- sions can be drawn. It is apparent that the hamsters of the genus Mesocricetus from Anatolia to at least as far east as Iran are not subspecies of the Syrian hamster, Mesocricetus auratus, as some authors have suggested (Ellerman and Morrison-Scott, 1951; Vereshchagin, 1959). While wild-caught Syrian, Turkish, or Ru- manian hamsters breed readily in captivity with animals from the Same area, all attempts to cross the Syrian with the Turkish or Kurdistan hamster have failed. If the animals will not breed in the laboratory under a wide series of conditions, it is unlikely that they would breed in the field. The bright golden brown color of the back, 10 BREVIORA No. 442 the absence of black subauricular and chest patches and the creamy white belly plus the smaller size of the Syrian hamster reinforce the concept that M. auratus is a separate species. When Syrian and Rumanian hamsters are compared, the differ- ence in pelage is equally striking. These animals are similar in size, but the Rumanian animals are much darker dorsally and on the belly, have darker chest patches, and the ratlike face is obvious in the live animals. Although viable crosses have been produced in the laboratory, both histological data and attempts at breeding indicate that the F; generation is sterile. On the basis of karyologi- cal studies (Raicu and Bratosin, 1968) as well as electrophoretic and chromatographic analysis of the blood serum, Raicu et al. (1968, 1969) stated that the Rumanian hamster, Mesocricetus newtoni, was a different species from the Syrian hamster, Mesocricetus auratus, and the data given above reinforce this conclusion. The Rumanian hamster is smaller and darker than the Turkish and Kurdistan animals, and the facial appearance in the live Rumanian animals is distinctive, but these differences are not sufficient for specific separation. Crosses between Rumanian hamsters and ani- mals from Malya in central Turkey or from Kurdistan produce viable young, but the F; generation is sterile. Furthermore, the occurrence of diabetes in the Rumanian-Turkish cross, and the small litters and failure of reciprocal crossing in the Rumanian- Kurdistan matings, are evidence that the crosses encounter biologi- cal barriers sufficient to separate them as species. When Lehman and Macpherson (1967) described the karyotype of their hamsters from Kurdistan, the animals were tentatively iden- tified by the British Museum as Mesocricetus brandti from skins and skulls. Our subsequent comparison with skins and skulls of hamsters from Malya reveal that the two groups resemble each other closely, though the Kurdistan animals are lighter on the dorsal sur- face. Color photographs of these animals alive, lent through the kindness of Dr. R. E. Billingham of the Wistar Institute, indicate that the animals do not possess the ratlike face which seems to be typical of hamsters from Rumania. After karyotypic examination of hamsters from Kurdistan, Malya, Rumania and Syria, Todd et al. (1972) concluded that the karyotypes of animals from Kurdistan and Malya most closely resembled one another. The evidence is persuasive that the animals from Malya and An- kara and the Kurdistan hamsters are all Mesocricetus brandti. The lighter dorsal pelage of the Kurdistan animals, when compared 1977 LABORATORY STUDY OF THE TURKISH HAMSTER It to the two colonies from central Turkey and the specimens in the British Museum, suggest that the Kurdistan animals may be a sub- species of M. brandti. In this regard, it is of interest that Mr. Anat, who is a trained field naturalist, has written us that the animals in the northeast area of Turkey near Kars are larger and darker than hamsters from central Turkey. Several subspecies of M. brandti may exist in Turkey, Armenia and northwest Iran, separated by various geographic barriers. We conclude that there are three distinct species of small hamsters of the genus Mesocricetus in areas south and west of the Caucasus. Mesocricetus newtoni inhabits the Dobruja area in the eastern part of Rumania and Bulgaria and, according to Hamar and Schutowa (1966), probably is not found west of the Danube in Rumania. Mesocricetus brandti ranges across Anatolian Turkey into the northern part of Iraq and the northwestern portion of Iran. Hamar and Schutowa (1966) and Aharoni (1932) report that M. brandti is found south of Aleppo and Aharoni indicates the southern limit is at the latitude of the Sea of Galilee. In contrast to this wide dis- tribution, M. auratus appears to be restricted to the area of the type locality of Aleppo, Syria. For definitive information on the distribution of Mesocricetus in and north of the Caucasus, one must turn to investigators who had access to live specimens or collections in Russia. Although there are differences of opinion on the number of subspecies involved, Ognev and Heptner (1927), Vinogradov and Argiropulo (1941), Hamar and Schutowa (1966) and Gromov et al. (1963) all agree that the genus in this area is clearly separable into two species, referred to here as M. brandti and M. raddei. M. brandti are smaller than M. raddei and there is no black on their bellies. In their karyological studies, Vorontsov and Krjukova (1969) and Ivanov (1969) also concluded that M. brandti and M. raddei were specifically different. The M. brandti studied by Ivanov came from the Daghestan area and Gromoy et al. (1963) state that this species is found in the plains and foothills of Daghestan. Gromov et al. (1963) and several other Russian investigators remark on the discontinuity of the distribution of M. brandti in the Caucasus, and we find nothing in the literature to indicate that it is actually sympatric with M. raddei in any area. Todd et al. (1972) reviewed the reported karyotypes of M. brandti from Daghestan, Armenia, Iranian Azerbajzan and Erevan, and of a specimen which was probably from Iran. They presented a haploid idiogram comparing the karyotypes of the Kurdistan animals with 12 BREVIORA No. 442 our hamsters from Malya, M. newtoni from Rumania and labora- tory M. auratus. The diploid number (2N) of M. newroni was 38, that of M. auratus was 44, and that of the various specimens of M. brandti was 42. They concluded that the differences in the karyotypes of the samples of M. brandti, including the fundamental numbers, was sufficient to suggest that the designation M. brandti was being applied to a group of “cryptic” species. At the time their data was collected, it was not known that the karyotypes of hamsters from Malya and Ankara differed both in diploid and in fundamental numbers, though the animals appeared phenotypically similar and produced fertile crosses in the laboratory. Zimmerman (1970) emphasizes the frailty of karyological data when used by itself to determine taxonomic relationships and the breeding success be- tween the Malya and Ankara group reinforces this concept. Recent evidence indicates that some rodents may be genetically isolated though phenotypically similar. Spalax (Nevo and Shkol- nik, 1974), Perognathus (Patton, 1972), Thomomys (Patton, 1973) and Peromyscus (Schmidly and Schroeter, 1974) are some of the rodents which have marked chromosomal differences in groups from restricted, contiguous areas. Patton and Dingman (1970) point out that the diversity in karyology often occurs in burrowing rodents with limited dispersibility and small breeding populations. Field reports, particularly those of Argiropulo (1939), and|our observa- tions of “field” conditions in the laboratory indicate that hamsters are fossorial rodents, though their way of life is not comparable to that of an animal such as Thomomys, which spends virtually its whole existence underground. Aharoni (1932) and Argiropulo (1939) both emphasize that Mesocricetus is found in very specific types of habitat and Argiropulo states that the distribution of brandti in Armenia is limited to specific altitudes. Mr. Anat has written us that there was no available open water in the areas where he trapped our animals, yet we find that hamsters in the laboratory which are fed laboratory rat chow cannot survive without water to drink. Thus, brandti in some areas must depend on metabolic water and water in their food, and this must limit their distribution. Further- more, Argiropulo states that they never live in wooded areas. These factors, coupled with their slow locomotion, may limit brandti to very specific types of habitat and isolate one group from another, which may explain why the karyotype of each population of this species examined to date differs from the other. 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 13 Aharoni (1932) points out that the Syrian hamster is confined to the immediate vicinity of Aleppo and suggests that the very dry climate has resulted in the recognizably different M. auratus. This population appears to have been isolated long enough to become specifically distinct, a conclusion that is supported. by the cross- breeding experiments. Evolutionary factors of less influence may be causing the differences in karyotypes of M. brandti without producing reproductive isolation or obvious differences in pheno- type. : NATURAL HISTORY Published observations of M. brandti under natural conditions are scanty, the most complete being that of Argiropulo (1939) who studied this species in the Caucasus. Because his paper is written in Russian, a brief summary is included here, supplemented with field notes taken by Mr. Anat when he was trapping hamsters for us in Turkey. Both sources agree that the habitat of M. brandti consists of dry, rocky steppe country sometimes bordering culti- vated fields. These hamsters do not inhabit wooded or bushy areas, and appear to avoid areas of high humidity and dampness. In the Caucasus they are found at altitudes as high as 2800 meters. Al- though they may be seen at dawn and dusk, they are mainly noc- turnal and feed principally at night. In the wild, they are relatively fearless, for Argiropulo observed that they were apt to stand on their hind legs facing an intruder in the agonistic posture so often seen in the Syrian hamster. In order to observe these animals under as natural conditions as possible, a compound measuring 160 X 280 centimeters (cm) and filled with clay soil to an average of 30 cm was set up in our labo- _Tatory. One part of the area was made higher than the remainder by the addition of rocks up to three kilograms in weight, and part of the flatter surface was planted with ground pine. A sunken dish served as a water supply and the bare soil was moistened periodically. During the month of July, two male and three female five-week- old M. brandti were released simultaneously into the compound. Within a few hours there were numerous holes in the soil and all of the animals were underground. For the next four months, no animal was ever seen in the broad daylight, though they were seen above ground when light was failing in the evening or under dim artificial illumination at night. 14 BREVIORA No. 442 During the period of observation, the animals were given Purina laboratory chow(Ralston Purina Co., St. Louis, Mo.) as food, which was thrown at random on the ground. Most of the food which was not eaten was stored underground during the night. Argiropulo noted that M. brandti stores both grain and grasses in its burrows. M. R. Murphy (personal communication) reports that Mesocricetus auratus obtains grain by standing on its hind legs, grasping the stalk with its front feet or pushing it down, and cutting the stalk with its in- cisors. The animal then eats the fallen grain or puts it in its cheek pouches. Numerous stalks of tall grasses with ripened ears were pushed into the soil of the compound to simulate growing grain and the animals were observed in the evening under dim illumination. In spite of the fact that these hamsters were the sixth generation of ani- mals raised in the laboratory, with no exposure to any growing grain, they cut down the stalk and consumed the kernels as described above. After the five hamsters had been in the compound for 130 days, they were removed using box traps, and a diagram of their tunnel system was constructed by squirting silicone foam (Froth Pak, Insta-foam Products Inc., Addison, Ill.) down the various holes and permitting it to harden. Once hardened, the earth was scraped away and a positive cast of the underground galleries was obtained. Although these casts show several points of interest, it must be borne in mind that the hamsters were limited in the depth of their digging by the floor of the room. At the time that the casts of the burrows were made, we were unaware of the work of Argiropulo (1939) with detailed descrip- tions of burrows which he excavated in the Caucasus. However, within the limitations imposed by the artificiality of the com- pound, the two series of burrows were remarkably similar. Both Argiropulo and Anat note that the main entrance of the burrow, when on a flat surface, is an almost vertical shaft extending a meter or more in depth. The vertical shaft usually curves quite abruptly to a horizontal plane, and the galleries and rooms branch from this shaft. Mr. Anat indicates that the hamster burrows can always be distinguished from those of Microtus or Citellus, which may inhabit the same area, because the burrows of the latter two enter the ground at a slant. When a hamster burrow was made in a bank, Argiropulo found that it was nearly hori- zontal. This type of tunnel was usually abandoned in a few days after other tunnels had been dug and the space filled in — 1977 LABORATORY STUDY OF THE TURKISH HAMSTER i) with dirt and feces. Both in the artificial situation and in the natural state there were several underground chambers. One or more of these chambers were used for storage of food. Another chamber was a nesting area which in the natural state was filled with dried grass. In the artificial compound, the nesting material was ground pine and shreds of a cardboard box that had been placed in the area as a surface shelter. Argiropulo found that one chamber was used exclusively as a latrine, but no such chamber was found in the artificial compound. To observe the method of digging employed by the hamster, a narrow, rectangular, clear Plexiglas box measuring 62 X 31 X3.5cm (inside dimensions) was constructed. The box was placed on edge with its long side down and filled with earth to a depth of 17cm. A hamster placed on top of the earth was squeezed so that both of its sides touched the sides of the box. The animal was capable of mov- ing forward or backward, or of turning around, but it could only rest comfortably with its body parallel to the long axis of the box. After several minutes of exploration, the hamster always com- menced digging in the earth. The rapidly moving front feet were used to displace the earth, with motions like a dog burying a bone. The loose dirt accumulated under the chest and abdomen. To move this dirt, the hind feet were thrust backward together, while the front feet were braced. During this maneuver, the hamster arched its back and lifted its head. If the animal was in a tunnel, the head- lifting served to tamp the earth on the roof of the tunnel. As the tunnel or the surface excavation became longer, the earth at the head of the excavation was moved to the tail by a series of parallel kicks of the hind feet accompanied by back arching and head lifting, as described above. After each set of kicks, the hamster moved backward about one-half the length of its body and resumed moving the earth. Use of the mouth and cheek pouches for moving dirt was not observed, though the animals often took small quantities of earth in their mouth as if to taste it. The rapidity with which hamsters can move earth and dig tunnels marks these rodents as truly fossorial, and indicates that hamsters kept in laboratory cages are denied a significant part of their nor- mal behavioral pattern. The lifting of the head to tamp the roof of the tunnel apparently has functional significance, for Argiropulo observed that the inside of the tunnels which he excavated were very smooth. 16 BREVIORA No. 442 LABORATORY CARE AND REPRODUCTION Turkish hamsters in this colony are now kept in individual 17 X 17 X 24 cm floored cages and given ample shavings for bedding. During breeding 23 X 23 X 38 cm cages are used. The hamsters do not thrive in wire bottomed cages. The animals are given water and Purina laboratory rat chow ad lib. with a slice of raw apple and 30 g rolled oats weekly. In 1968-1969 the colony was greatly reduced due to “wet tail” diar- rhea and osteomalacia. The diarrhea was partially controlled by adding 0.05 per cent tetracycline hydrochloride (Polyotic, Cyana- mid) to the drinking water for a period of not longer than two weeks. Various diets, one consisting wholly of Purina mouse breeder chow, have been experimentally tested, but the diet outlined above has been most satisfactory. The cause of osteomalacia has not been determined, but since it no longer occurs in the colony, it proba- bly was due to an insufficiency in the diet exacerbated by an intes- tinal infection which, in its most virulent form, resulted in “wet tail” and death. Although it rarely occurs in M. auratus, several M. brandti suf- fered from malocclusion, so that the upper and lower incisors grew unchecked by wear. The unrestrained growth was remarkably rapid. In a three-month-old hamster the upper incisors had grown slightly more than a full circle with one incisor penetrating and exiting from the roof of the mouth. The lower incisors of a one-year-old hamster with malocclusion were trimmed periodically and the ex- cess tooth material was measured. Growth averaged | cm per month compared to growth of approximately 0.4 cm per month reported by Sarnat and Hook (1941) in the much larger thirteen-lined ground squirrel (Citellus tridecemlineatus). Pairs of hamsters from five weeks to two years of age were exposed to each other for breeding in every month of the year. The young- est female to produce a litter was 50 days old when she gave birth, and animals seven to eight weeks of age reproduce successfully though they have not yet attained full growth. Hamsters reach their peak of fecundity at about one year of age, and most of our breeding is carried out with animals of this age. The tests of one- year-old males are large during the breeding season and histological examination shows active spermatogenesis. Females of this age almost invariably have normal estrous cycles. In the second year, the female estrous cycles are less regular during the breeding season 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 17 though active spermatogenesis usually occurs in the males. Breed- ing is notably less successful in these animals. We have not at- tempted to breed three-year-old hamsters. In our laboratory (Boston, Mass.), animals exposed to natural day length usually were not in breeding condition between the months of November and March. During this period the females which were examined were apt to have anestrous vaginal smears with a preponderance of non-nucleated squamous cells. The testes of some rhales were atrophic with no histological evidence of sper- matogenesis, and no stored sperm in the epididymides. Other animals of both sexes had all the anatomical indications of full breeding condition. The testes were the same size as those of sum- mer animals and histological examination revealed spermatogenesis, while biopsy of the epididymides showed motile sperm. Vaginal smears indicated normal estrous cycles. In 1971, pairs of hamsters were exposed to each other from Janu- ary to September. As was expected, successful breeding was low for the first three months of the year, improved in the next two months and then declined. Thus, from January 20 to April 3 there were |1 litters out of 33 exposures. Successful matings peaked in May, with seven litters born out of 13 exposures. From June | to September 3, there were only 11 litters born in 34 exposures. From 1966 to 1971 various changes in daily illumination and diet were carried out in order to discover the optimum conditions for breeding this species in the laboratory. During the first two years after their arrival, the animals were maintained on 10 hours of il- lumination in each 24 hour period. Breeding success was high dur- ing the first summer (1966) with a total of 109 young in 16 litters being produced by 23 pairings from the wild-caught group of six males and seven females. During the next summer there was a notable decline in successful matings. Of a total of 79 pairings with 42 males and 45 females, only 111 young in 15 litters were produced. In the case of unsuccessful pairings, the female always appeared to be the unwilling one of the pair. In the difficult years of 1968 and 1969 the colony barely main- tained a steady population, but by 1970 a successful diet had been developed, the daily lighting had been standardized to the illumina- tion of this latitude, and attempted breedings were usually limited to the spring and summer months. Under this regime, breeding success improved, with 17 litters in 34 matings in 1974. 18 BREVIORA No. 442 Hoffman and Reiter (1965) and Elliott et al. (1972) have reported that the reproductive cycle of the male Syrian hamster is responsive to photoperiod. A controlled experiment to test this responsiveness in the Turkish hamster has not been carried out, but the birth of litters in January and February, with the day length near the mini- mum, shows that the photoperiodic effect is not absolute. Data presented below indicate that hibernation is involved in a much larger segment of the yearly life of the Turkish hamster than is the case in the Syrian species. Turkish hamsters which are exposed to a constant temperature of 22°C \in the laboratory during the winter months are living in an environment which they would never en- counter for long periods in the natural state. It is not surprising that photoperiod alone does not control the seasonal sexual cycle under these circumstances. The breeding behavior of the Syrian hamster has been described by Murphy and Schneider (1970) and the behavior of the Turkish hamster is similar. When the animals were paired in the afternoon, there was usually some preliminary exploration, but not much ac- tivity occurred until nightfall. In some cases, however, an immedi- ate antipathy developed and violent fighting occurred at once. After the first half hour of exposure, fighting was not as frequent, but it could occur at any time. If the animals were kept together for more than four days the chances of fighting increased, perhaps be- cause the female was pregnant. Fighting was always violent, with the female biting the male near the head region and then clawing him with both front and rear feet. The male, in spite of the punish- ment he received, often followed the female and attempted to copu- late, only to be attacked again. If the male retreated, the female often pursued him and bit him on the flanks and hind quarters. Attempts to reduce the damage to the male by providing various shelters were unsuccessful because of the male’s persistence in pur- suing the female. Often the punishment received by the male was fatal. At autopsy the cuts and bruises did not seem sufficiently severe to cause death and it is possible that exhaustion was a con- tributing cause. The estrous cycle of the Turkish hamster is similar to that de- scribed by Kent (1968) for the golden hamster. The cycle is of four days duration and estrus is characterized by the appearance of large numbers of nucleated epithelial cells in the vaginal smear. Gestation is 15 days in length and the litter size has varied from one to 13 with the average being six (N young= 1021, N litters= 177). 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 19 No female has produced more than three litters during one breeding season and Argiropulo (1939) reports that two litters a year is maxi- mum for wild animals in the Caucasus. The female becomes ex- tremely apprehensive before the birth of her litter, and partial darkness and ample bedding area necessity. Evenif left undisturbed she may kill and eat the young, and if she is handled during parturi- tion the destruction of the litter is inevitable. Perhaps a littering cage which approximated the fossorial existence in the wild would reduce this nervousness, but this has not been tested. Hamsters are born blind and naked, but they quickly grow a protective coat. By the twelfth to thirteenth day the eyes open and the young eat rolled oats, ground laboratory chow and drink from the water bottle. The young are weaned by the mother before the twentieth day, and we routinely separate the young from the mother at four weeks of age. The young live together amicably until about the seventh week, when fighting starts to occur. Usually one animal is the aggressor, but if it is removed from the litter, another soon takes up the role. The attacked animal is bitten violently, resulting in large wounds from tears in the tender skin, and these are often fatal. Presumably, under natural conditions this aggressive behavior causes dispersion of the litter with little harm to its members, but in the laboratory it is a serious problem. Although we have used several types of shelters, we have not been able to keep Turkish hamsters in collec- tive cages and must separate and maintain them in single cages at about six weeks of age. Obviously this is a serious disadvantage for maintaining a large colony. The young hamsters grow rapidly, and healthy litters average 91 g in weight with a range of 50 to 141 g (N = 411) at six to seven weeks of age. Animals born after June do not attain sexual maturity until the following spring. AGING Turkish hamsters in captivity tend to become obese after the first year of life and sometimes weigh over 200 g, with our heaviest recorded animal attaining 285 g. It is unlikely that hamsters in the wild, with limited food supply and an active life, would ever reach this weight. The median life-span of 43 animals which were bred and kept in the laboratory at a room temperature of 22 + 3°C was 670 days with a range of 231 to 1399 days. After about two years of age, the 20 BREVIORA No. 442 pelage usually loses its glossy smoothness and the animals present a scruffy appearance. The effect of hibernation on aging in this or any other species of mammal is unknown and an experiment is in progress to determine if these hamsters age less rapidly during hibernation. HIBERNATION During the past ten years, quite complete records of the colony of M. brandti have been maintained, so that some information can be gleaned concerning the putative factors which control hibernation. Previous observations on M. auratus in this laboratory (Lyman, 1954; Lyman, unpublished observations) provide comparison be- tween the two species. In those studies, all hamsters were moved toa cold room kept at 5 + 2°C and lighted 8 to 10 hours a day. The animals were housed in individual cages with ample shavings and food and water ad lib. Hibernation was monitored by dusting the backs of hibernating Syrian hamsters with shavings. If the shavings were still in place at the next observation, hibernation was assumed to be continuous. This technique is common among students of hibernation (Mrosov- sky, 1971) but is not always accurate. Because hamsters often bur- row in their bedding before hibernation, shavings occasionally be- come lodged on the back and the hamster may be erroneously scored as being in continuous hibernation. Following the technique of Johnson (1931), extraneous material (rolled oats) was placed on the backs of the hibernating Turkish hamsters. Arousal from hiber- nation invariably displaced this material. The result of this differ- ence in technique is to make the Syrian hamsters appear to be “bet- ter” hibernators than they actually are. The timing and period of cold exposure was not the same for M. auratus and M. brandti. The former were moved to the cold at various times of year and remained there until they hibernated or died. In contrast, because it was necessary to enlarge the M. brandti colony, most of these animals were exposed to cold in the autumn and removed to the warm room for breeding in the spring. The fact that the M. auratus were moved to the cold in all seasons of the year should have no effect on the comparisons given below. Smit-vis and Smit (1963) have theorized that there is a seasonal preparation in this species which results in a greater tendency for 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 21 hibernation in February. The evidence is not persuasive, however, for it is based on 26 animals, two of which are excluded from the basic computations. The paper relates the month in which the ani- mals were exposed to cold to the lapse of time before hibernation occurred, but there are no data for April through July or for Sep- tember and October, and there is data for only one animal in August and two in March. From our records of over 2800 Syrian hamsters, placed in the cold in every month of the year, we adduce no con- vincing evidence that the tendency to hibernate is seasonal in this species. In studies such as this, an as yet unresolved problem concerns the criteria which should be used to distinguish “good” from “poor” hibernators. There are at least five interdependent factors which can be considered in such a comparison. These are: 1) the number of animals in a group which hibernate during the period of exposure to cold, 2) the span of time between cold exposure and hibernation, 3) the length of the hibernating season, 4) the amount of time spent in hibernation compared to the amount of time in the cold, and 5) the length of the period during which an animal remains in con- tinuous hibernation. Concerning the fifth factor, Pengelley and Fisher (1961) have shown that the lengths of the periods or bouts of hibernation are quite predictable with individual Citellus lateralis, but these lengths may vary greatly ina group. We find that the same situation obtains in Mesocricetus so that the maximum time of un- interrupted hibernation is more definitive than an average of the lengths of the bouts of hibernation. Until more is known about the subtle physiological nuances which determine whether an animal does or does not hibernate, it is not practical to attempt to assign meaningful values to the five factors listed above, nor to make statistical comparisons. Therefore, in the following studies as many factors as possible are considered but no attempt is made to weight their importance. In a group of 373 Syrian hamsters exposed to the cold, 68 per cent or 252 hibernated and 121 animals died before ever entering the hibernating state. The shortest period before hibernation occurred was three days of cold exposure and the longest was 218 days (Ly- man, 1954). Once hibernation occurred, the period during which M. auratus hibernated lasted about three months. An accurate figure on the percentage of time spent in hibernation cannot be obtained because many of the animals were used in acute experi- ments and the best hibernators were usually chosen. The longest 22 BREVIORA No. 442 recorded bout of hibernation was 21 days and the usual period of hibernation was a week or less (Lyman, 1954). The hamsters from which these data were collected were orig- inally obtained from commercial dealers and hence must be de- scendants of the trio which were the foundation stock of the labo- ratory hamsters used today (Adler, 1948). Twelve M. auratus, which were the first generation of animals obtained in the field near Aleppo by Dr. M. Murphy, were exposed to the cold in the autumn of 1971 under the conditions described above. Although the sample is small, there was no detectable difference in the tendency of this group of animals to hibernate when compared to laboratory ani- mals. Thus, forty years of domestic breeding with no selection pres- sure for hibernation has not changed the pattern of hibernation in M. auratus. These “native” Syrian hamsters provide some information con- cerning the percentage of time spent in hibernation. The group was moved to the cold on November 15, 1971 and taken from the cold on May 26, 1972. No animal hibernated before January and hibernation occurred only 11.3 per cent of the total time. The best months for hibernation were January through April, during which time the animals were in hibernation 13.7 per cent of the time. Between the years 1971 and 1975, a group of 318 M. brandti were exposed to cold for periods of 145 to 172 days. Two hundred ninety- three or 92 per cent hibernated at some time during this period. Eight animals died without hibernating and the remaining 17 did not hibernate at all. Thirty-one animals entered hibernation within 24 hours after exposure to cold. Once hibernation started, the period during which this species hibernated lasted about five months, but for some animals the hibernating season was as long as 10 months. In the entire group, 36 per cent of the time in the cold be- tween November 15 and April 15 was spent in hibernation. The longest period of uninterrupted hibernation was 28 days, with the average bout of approximately the same duration as M. auratus. Thus M. brandtiis a “better” hibernator than M. auratus in all cate- gories except the average Jength of the bout of hibernation. The records of over 400 M. brandti which have been exposed to cold reveal no obvious factors which control or influence hiberna- tion in this species. However, they do give a clearer picture of the pattern of hibernation and its variability, and obviate the need for further research in some aspects of the problem. 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 23 As reported for M. auratus (Lyman, 1948), M. brandti does not undergo the period of autumnal fattening and lethargy which is so typical of the Marmotini that hibernate. Like M. auratus, M. brandti almost invariably lose weight during the first two or three weeks of exposure to cold, with the heavier animals losing the greater amount. Some animals, especially the young of the year, are apt to regain and exceed the original weight if kept in the cold for several months. The amount of hibernation which occurs during this time is not correlated with the change in body weight. Examples from the spring and summer months demonstrate that this species will hibernate at any time of the year. Certainly, if com- pared to Marmotini such as Citellus lateralis or C. tridecemlineatus, any seasonal change in the tendency to hibernate is very poorly de- fined. These citellids, if experimentally naive, rarely hibernate when exposed to cold during the spring and summer months and almost invariably hibernate in the autumn. In contrast, of 66 M. brandti which were in the cold in June, 75 per cent hibernated at some time during the period and, in July, 77 per cent of 53 cold- exposed animals hibernated. The majority of our animals were placed in the hibernaculum between October | and January 2. In a sample of 257 animals, no particular one of the four months was favored for onset of hiberna- tion or for the number of days spent in the hibernating state. Wheth- er exposure to cold was begun in early October or in January, hi- bernation became much less frequent by the beginning of April and, when removed from the cold room in May, most of the animals had ceased hibernating. No organized experiment to test the effect of photoperiod has been carried out, but it is clear that the timing of the onse* of hiberna- tion does not depend exclusively on this factor, since hamsters hi- bernated in June when the daily illumination was close to its peak. Although 9.7 per cent of a sample of 318 animals hibernated with- in 24 hours after being exposed to cold, these animals did not neces- sarily hibernate for a greater number of days than animals which started hibernation at a later date. Age does not affect the ability to hibernate, for a comparison between animals ranging from 3.5 to 36 months of age revealed no differences in the onset or pattern of hibernation. In spite of efforts to standardize conditions, the frequency and pattern of hibernation changed from year to year in the colony as 24 BREVIORA No. 442 a whole and with individual animals. There is no simple explana- tion for the fact that individuals failed to hibernate with six months of cold exposure one year, and hibernated under the same condi- tions the following year. Nor can the observation that the colony exposed to cold hibernated only 16 per cent of the time in 1970 and 39 per cent in 1971 be readily explained. Conditions in the cold room are maintained as identically as possible from year to year, but changes such as shifts in caretaking personnel are inevitable. Even differences in handling the animals during cleaning of the cages might influence the onset of hibernation. It was observed that the first shipment of M. brandti hibernated 67 per cent of the first sojourn in the cold and only 42 per cent in the second. This suggested that environmental factors might play an important role in the control of hibernation. The ability of ani- mals to store food had already been shown to have an effect on hi- bernation in M. auratus (Lyman, 1954) and this factor was included in the experimental design. M. brandti between five weeks and six months of age were placed in the following experimental conditions. As often as possible, sexes and litter mates were matched. On July 8 five animals were placed in the “natural” compound described above. For all other groups, each individual was put in a separate cage (23 X 23 X 38 cm) and given ample shavings for bedding, and water ad lib. A control group of 13 animals was given Purina rat chow pellets ad /ib., which they were able to store in a corner of the cage, while another group of six was given ground chow in unspillable food cups which pre- vented these animals from storing food. Twelve other cages were fitted with exercise wheels, and the number of revolutions for each wheel was monitored by an event recorder (model A620X, Esterline Angus Instrument Co., Indianapolis, Ind.). The animals in six of these cages were given rat chow pellets ad /ib. while the others were fed from unspillable food cups. All animals received 14 hours of light daily at the start of the experiment, and this was reduced by one hour approximately every 16 days. On November 12 all animals were moved to smaller individual cages (17 X 17 X 24 cm) with shavings, water and rat chow pellets ad /ib., and on November 15 all were moved in these cages to the cold room, where the daily il- lumination was 9 hours. Thus, after the preparation period, all ani- mals were exposed to the same conditions. Records of hibernation were kept for the following 137 days. 1977 LABORATORY STUDY OF THE TURKISH HAMSTER 25 As has been reported for many other species, the use of the ac- tivity wheel varied greatly from animal to animal. The nocturnal habits of M. brandti were verified, for virtually no activity was recorded during the daylight hours. There was no correlation be- tween the tendency of an animal to run in the wheel and its subse- quent hibernation. All of the hamsters hibernated at some time during their exposure to cold, but no one group showed a tendency to enter hibernation earlier or to have longer bouts of hibernation. The percentage of time spent in hibernation was as follows: “natu- ral” enclosure, 56 per cent; activity wheel, food hoarding, 43 per cent; no wheel, food hoarding, 37 per cent; wheel, no hoarding, 34 per cent; no wheel, no hoarding, 25 per cent. These results suggest that the conditions prior to cold exposure may have an influence on the subsequent amount of hibernation and that “natural” conditions increase the tendency to hibernate. However, none of the factors tested are crucial to the onset of hibernation. Experiments such as these are easily carried out with M. brandti. Because this animal hibernates readily and can be bred in the labo- ratory, it offers a unique opportunity to study the effects of various factors using matched pairs of known age and lineage. ACKNOWLEDGMENTS This research was supported by grants from the National Institutes of Health and contracts with U.S. Air Force. In this ten year study many people contributed their help. We are particularly grateful to Jane L. Bihldorff, Michael F. Carr, Helen Cheever, and Delphi M. Toth. We owe great thanks to Dr. N. Kala- bukhov of the Far East Scientific Center of the Academy of Sciences, Vladivostok, USSR, who has been most helpful in giving advice and references concerning the species of Mesocricetus in the USSR. We are indebted to Barbara Lawrence for her critical review of the manu- script, but any errors or ambiguities are uniquely our own. LITERATURE CITED ADLER,S. 1948. Origin of the golden hamster Cricetus auratus asa laboratory ani- mal. Nature, 162: 256-257. AHARONI, B. 1932. Die Muriden von Palastina und Syrien. Z. Saugetierk., 7: 166-240. ALTMAN, P. L., AND D. S. Ditrmer. 1964. Biology data book, Fed. Amer. Soc. Exp. Biol., Washingion, D. C. p. 95. 26 BREVIORA No. 442 ArGIROPULO, A. I. 1939. The distribution and ecology of certain mammals of Armenia. (translation) (Russ.) Acad. Sci. U.S.S.R., Armenian Branch, Trans. Biol. Inst., 3: 47-57. ELLERMAN,J.R. 1941. The families and genera of living rodents, Vol. 2. London: British Museum (Nat. Hist.) pp. 1-690. 1948. 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Pineal gland: influence on gonads of male hamsters. Science, /48: 1609-1611. Ivanov, V.G. 1969. The chromosome complements of Mesocricetus of the Cauca- sus and their taxonomic position. Jn N. N. Vorontsov (ed.), The mammals, evolution, karyology, taxonomy, fauna (translation). Acad. Sci. USSR, Si- berian Branch. pp. 105-106. Jounson, G. E. 1931. Hibernation in mammals. Quart. Rev. Biol., 6: 439-461. KenT, G. C., Jr. 1968. Physiology of reproduction. Jn R.A. Hoffman, P. F. Robinson, and H. Magalhaes (eds.), The golden hamster, its biology and use in medical research. Ames, Iowa: lowa State University Press. pp. 119-138. LEHMAN, J. M.,AND J. MACPHERSON. 1967. The karyotype of Kurdistan hamster. J. Hered., 58: 29-31. Lyman, C. P. 1948. The oxygen consumption and temperature regulation of hibernating hamsters. J. Exp. Zool., /09: 55-78. 1954. Activity, food consumption and hoarding in hibernators. J. Mamm., 35: 545-552. Mrosovsky, N. 1971. Hibernation and the hypothalamus. New York: Appleton- Century-Crofts. vi + 287 pp. Murpnuy, M. R., AND G. E. SCHNEIDER. 1970. Olfactory bulb removal eliminates mating behavior in the male golden hamster. Science, 167: 302-304. Nevo, E., AND A. SHKOLNIK. 1974. Adaptive metabolic variation of chromosome forms in mole rats, Spalax. Experientia, 30: 724-726. Ocnev, S.1.,AND V.G. HEpTNER. 1927. A newsubspecies of hamster from Daghe- stan (Mesocricetus raddei, Nehr.) and some remarks on the Russian species of Mesocricetus (with a key to them). Ann. Mag. Nat. Hist. ser. 9, /9: 142-150. Pam, J., W. K. Si-vers, AND R. E. BILLINGHAM. 1967. The problem of histo- compatibility in wild hamsters. J. Hered., 58: 40-44. Patron, J. L. 1972. A complex system of chromosomal variation in the pocket mouse, Perognathus baileyi Merriam. Chromosoma (Berl.), 36: 241-255. 1973. Patterns of geographic variation in karyotype in the pocket gopher, 1977 LABORATORY STUDY OF THE TURKISH HAMSTER Di Thomomys bottae (Eydoux and Gervais). Evol., 26: 574-586. Patton, J. L.,AND R. E. DincMan. 1970. Chromosome studies of pocket gophers, genus Thomomys. II. Variation in T. bottae in the American southwest. Cyto- genetics, 9: 139-151. PenceLtey, E. T., AND K. C. FisHer. 1961. Rhythmical arousal from hibernation in the golden-mantled ground squirrel, Citellus lateralis tescorum. Can. J.Zool., 39: 105-120. Raicu, P., AND S. Bratosin. 1968. Interspecific reciprocal hybrids between Mesocricetus auratus and M. newtoni. Genet. Res. Camb., //: 113-114. Raicu, P., M. Hamar, S. BraTosin, ANDI. Borsan. 1968. Cytogenetical and bio- chemical researches in the Rumanian hamster (Mesocricetus newtoni). Z. Saugetierk., 33: 186-192. Raicu, P., M. Ionescu-VARo, AND D. Duma. 1969. Interspecific crosses between Rumanian and Syrian hamsters. J. Hered., 60: 149-152. Raicu, P., M. Ionescu-VARo, M. NICOLAESCU, AND M. KiriLLova. 1972. Inter- specific hybrids between Romanian and Kurdistan hamsters. Genetica, 43: 223-230. Ripcway,R. 1912. Color standards and color nomenclature. Washington, D. C.: Ridgway, 53 plates & 44 pp. SarnatT, B. G., AND W. E. Hook. 1941. Effect of hibernation on rate of eruption and dentin apposition in the ground squirrel. Proc. Soc. Exp. Biol. Med., 46: 685-688. ScHMIDLY, D. J., AND G. L. SCHROETER. 1974. Karyotypic variationin Peromyscus boylii (Rodentia: Cricetidae) from Mexico and corresponding taxonomic im- plications. Systematic Zool., 23: 333-342. Smit-vis, J. H., AND G. J. Smit. 1963. Occurrence of hibernation in the golden hamster, Mesocricetus auratus Waterhouse. Experientia, 19: 363-364. Topp, N. B., C. W. Nixon, D. A. MULVANEY, AND M. E. ConnELty. 1972. Karyo- types of Mesocricetus brandti and hybridization within the genus. J. Hered., 63: 73-77. VERESHCHAGIN, N. K. 1959. The mammals of the Caucasus. Academy of Sciences of the USSR, Zoological Institute. Academy of Sciences of the Azerbaidzhan SSR. Jerusalem: Israel Program for Scientific Translation. pp. 1-816. VINOGRADOV, B. S.,AND A. I. ARGiRopuULO. 1941. Fauna of the USSR. Mammals. Key to rodents. Zoological Inst. Acad. Sci. USSR. Jerusalem: Israel Program for Scientific Translation. 230 pp. Vorontsov, N. N., AND E. P. Kruukova. 1969. The karyotypical differentiation of Mesocricetus raddei and taxonomical relationships of members of the genus Mesocricetus. In N. N. Vorontsov (ed.), The mammals, evolution, karyology, taxonomy, fauna (translation). Acad. Sci. USSR, Siberian Branch. pp. 107- 109. ZIMMERMAN, E.G. 1970. Karyology, systematics and chromosomal evolution in the rodent genus, Sigmodon. Publ. Mus. Mich. State Univ., Biol. Ser. 4(9): 389-454. teers ae Ji SS) f BREV I Gime A Museum of C ompar ative Zoolo gy US ISSN 0006-9698 CAMBRIDGE, MASS. 27 MAY 1977 NUMBER 443 EPIGONUS TREWAVASAE POLL, A JUNIOR SYNONYM OF EPIGONUS CONSTANCIAE (GIGLIOLTI) (PERCIFORMES, APOGONIDAE) GARRY F. MAYER! AND ENRICO TORTONESE? ABSTRACT. The holotype of Pomatomichthys constanciae Gigholi was reexam- ined and compared with specimens of Epigonus trewavasae Poll from equatorial west Africa, northwest Africa, Madeira, and the western Mediterranean. The two nominal forms were not distinguishable at the species level; thus, the name Epigonus constanciae should be used for the species under consideration. A description and discussion of the geographic variability of FE. constanciae is provided. INTRODUCTION Questions about the status of Pomatomichthys constanciae have recently arisen as the result of increased interest in deep-sea members of the family Apogonidae. P. constanciae was originally described from a single specimen from the Straits of Messina (Gigholi, 1880). The description was incomplete, unillustrated and included several inaccuracies. Although Giglioli designated the fish a new species, he indicated that the specimen resembled juveniles of the more common Mediterranean apogonid, Epi- gonus telescopus (Risso). Perhaps because of Giglioli’s comment and because of the dearth of information about Mediterranean continental slope ichthyofaunas, P. constanciae was generally accepted as a junior 'MESA New York Bight Project, National Oceanic and Atmospheric Admin- istration, Old Biology Building, State University of New York, Stony Brook, N.Y. 11794, U.S.A. 2Museo Civico di Storia Naturale, “Giacomo Doria,” Via Brigata Liguria 9, Genoa, Italy. nN BREVIORA No. 443 synonym of E. telescopus (e.g., Jordan and Evermann, 1896; Jor- dan, Evermann and Clark, 1930; Schultz, 1940; Tortonese and Queirolo, 1970; Tortonese, 1973). Goode and Bean (1896) were the last authors of a comprehensive treatise to give P. constanciae consideration as a separate species. Cavaliere (1956) claimed to have compared the juveniles of Epigonus telescopus and Poma- tomichthys constanciae, but Tortonese and Queirolo (1970) and Tortonese (1973) deemed the study of doubtful validity. In 1974, Mayer published a revision of the genus Epigonus. Although the paper included P. constanciae as a synonym of E. telescopus, it pointed out that several features of P. constanciae were reminiscent of traits characteristic of Epigonus trewavasae Poll. Yhe same work reported E. trewavasae from the Mediter- ranean for the first time and urged that further studies be under- taken on the relationship between E. trewavasae and P. con- stanciae. The current paper stems from a reexamination of the P. con- stanciae holotype and a comparison of this specimen with E. tre- wavasae from equatorial west Africa, Mauritania, Madeira and the western Mediterranean. Our findings indicate that P. constan- ciae and EF. trewavasae are conspecific and that the correct name for the species should be Epigonus constanciae (Giglioli). METHODS Specimens were examined according to techniques outlined by Mayer (1974, 1975) and by Hubbs and Lagler (1958). Statistical analyses were performed on a Wang Model 720 Programmable Calculator at the Department of Marine Science, University of South Florida. A package program was used to calculate routine meristic parameters, such as means and standard deviations. Linear regression analyses were undertaken on mensural data using Ricker’s method for Functional (GM) Regressions (1973: 412-414). To insure linearity, scatter diagrams were plotted for all morphometric data. Characters exhibiting non-linear scatters, as determined by inspection, were excluded from consideration. Meristic characters exhibiting slight variability are reported in the text as the value of the character followed by the number of individuals exhibiting that character. Data exhibiting greater variability are presented in tabular form. Mensural data are 1977 E. CONSTANCIAE DESCRIPTION 3 presented as linear regressions and/or as ratios on standard length (SL) or head length (HL). Specimens for this study were obtained from the following institutions: IRSN: _ Institut Royal des Sciences Naturelles de Belgique, Brussels ISH: Institut fiir Seefischerei, Hamburg ISTPM: Institut Scientifique et Technique des Péches Maritimes, La Rochelle MCZ: Museum of Comparative Zoology, Cambridge, Massa- chusetts MF: Museo Municipal do Funchal, Madeira MSNG: Museo Civico di Storia Naturale, Genoa MZF: Zoological Museum, Florence USNM: National Museum of Natural History, Washington, D.C. ZMB: Zoologisches Museum der Humboldt Universitat, Berlin SYSTEMATICS Mayer (1974: 183-186) provided a comprehensive description of Epigonus trewavasae. The following discussion of E. constan- ciae represents a recapitulation of the former account, modified to reflect data obtained from a larger, more representative sam- ple of individuals. Readers are referred to Mayer (1974) for synonymies, accounts of ontogenetic change and in-depth com- parisons with congeners. Figure 1. Epigonus constanciae holotype, MZF 3089, 115 mm SL. 4 BREVIORA No. 443 Epigonus constanciae (Gigholi, 1880). Figure 1. Pomatomichthys constanciae Gigholi, 1880: p. 20; original descrip- tion; Straits of Messina. Holotype examined: MZF 3089. Epigonus trewavasae Poll, 1954: p. 91; figure 27; original descrip- tion; NOORENDE III, Sta. 52: 06°08’S, 11°30’E, 280-290 m. Holotype examined: IRSN 209. MATERIAL EXAMINED IRSN 209 (1: 153.9 mm, holotype Epigonus trewavasae, NO- ORDENDE III Sta. 52, 06°08’S, 11°30°E, 280-290 m):"IRSN 16549 (1: 97.1 mm, Golfe de Caghari, 540 m); IRSN 16550 (4: 75.5-98.6 mm, Golfe d’Oristano, no depth recorded); ISH 386/64 (3: 144.6-153.2 mm, WALTHER HERWIG Sta. 137/64, 15°51’N, 1l6°S1’W, 200 m); ISTPM_ uncatalogued (10: 129.9-158.9 mm, THALASSA Sta. U30, 17°56’N, 16°32’W, 224-322 m); ISTPM uncatalogued (1: 149.0 mm, THALASSA Sta. X007, 20°59’N, 17°37’W, 250-330 m); ISTPM uncatalogued (1: 148.0 mm, THA- LASSA Sta. X020, 19°08’N, 16°48’W, 280-468 m); MCZ 48849 (le 76:9 mm, LA RAFALE Cr. I, Sta. 27, 11°2s:Ne eae 200 m); MF 3694 (1: 117.6 mm, from stomach of Polymixia nobilis); MF 7231 (1: no SL, from stomach of Aphanopus carbo); MF 8213 (1: no SL, from stomach of 7. nobilis): ME I25 18k 124.5 mm, from stomach of P. nobilis); MF 14695 (1: 131.3 mm, from stomach of 4. carbo); MF 15650 (1: no SL, from stomach of A. carbo); MF 16067 (2: 120-127 mm, from stomach of carbo); MF 17088 (1: 125.5 mm, from stomach of A. carbo): MF 18204 (1: 97.0 mm, from stomach of A. carbo); MF 20599 (1: no SL, from stomach of A. carbo); MSNG 38155 (1: 89 mm, Messina, Sicily); MZF 3089 (1: 117 mm, holotype of Pomato- michthys constanciae, Straits of Messina); USNM 207723 (2: 98.6-100.9 mm, LA RAFALE Cr. I, Sta. 27, 11°25°N) 17-21 aANe 200 m); USNM 207724 (1: 70.9 mm, LA RAFALE-Gr, I, Stave2re 11°25’N, 17°21’W, 200 m); ZMB 23101 (1: 148.9 mm, MORS Sta. 136/72, 18°36’S, 11°29’E, 275 m); ZMB 23102 (1: 11625 ame MORS'Stavl/5/72,.138°05 So 276E- 360) im): DIAGNOSIS In the past, £. constanciae has been mistaken for F. telescopus (Risso) and F&. pandionis (Goode and Bean). The distribution 1977 E. CONSTANCIAE DESCRIPTION 5 of E. constanciae also makes confusion likely with FE. denticulatus Dieuzeide. FE. constanciae may be distinguished from the above species by the presence of VII-I, 9 dorsal fin elements, a pungent opercular spine, lingual teeth and six to eight pyloric caeca. E. constanciae most closely resembles EF. pectinifer Mayer, a species found in the Caribbean Sea, the Gulf of Mexico and the western Pacific Ocean. The former is distinguished by the pres- ence of numerous awl-like fill rakers (29-36) and elongate DI and All fin spines (12.7-16.5 per cent and 13.5-16.8 per cent SL, respectively}. DESCRIPTION See Tables | and 2 for morphometric and meristic data. Body elongate, fusiform in larger individuals; body depth 21.1-30.0 per cent SL. Head moderate, length 32.2-38.9 per cent SL; height 16.2-18.7 per cent SL. Snout tapering to a rounded point, 7.0-9.1 per cent SL. Lower jaw 13.9-17.6 per cent SL; equal to or projecting slightly beyond upper jaw; bearing two nubs of variable prominence on anterior surface of mandible near symphysis. Maxilla reaching |/3-1/2 length of eye; ventro- lateral surface of maxillary head bearing sharp, posteriorly re- Table |. Epigonus constanciae regression parameters. u= functional (GM) regression coefficient; v= Y intercept of GM regression line, Y = uX + v; b=predictive regression coefficient +95 per cent confidence interval; a = Y intercept of predictive regression line, Y = bX + a; n = number of specimens. uv ud a a n HL 0.391 4.033 0.348 + 0.026 3.240 33 Head height 0.191 -1.810 0.189 + 0.011 1.566 30 Body depth 0.332 8.043 0.324 + 0.027 7.062 33 Eye diameter 0.168 0.976 0.164+0.015 0.367 BY Snout length 0.078 0.333 0.074 + 0.010 / 0.922 32 Maxillary length 0.161 0.384 0.157 + 0.013 0.076 34 Lower jaw length 0.174 1.280 0.167 + 0.017 0.428 36 Interorbital width 0.090 0.910 0.087 + 0.008 1.300 36 Caudal peduncle length 0.269 0.333 (0.264 + 0.018 0.242 35 Caudal peduncle depth 0.145 3.418 0.139 +0.015 2.673 34 Dol 0.156 1.219 0.150 + 0.015 0.546 31 All 0.156 0.547 0.149 + 0.019 ().282 27 PI 0.145 (0.298 0.142 + 0.009 0.233 35 6 BREVIORA No. 443 Table 2.) Epigonus constanciae meristic data. X= mean: SD= standard devia- tion: n= number of specimens. cS Range SD n Pectoral fin rays 17.65 16.19 0.62 40 Gill rakers Total 32.43 29 36 eal) 40 Upper arch 9.18 8 10 0.64 39 Lower arch 23} 20=25 1.30 39 Lateral line scales 47.74 46 50 0.93 34 Pyloric caeca 7.00 6-8 ().64 35 curved process, may be obscured by overlying skin. Eye longer than high, reaching upper profile of head, 41.1-49.1 per cent HL. Interorbital width 8.7-10.9 per cent SL. Caudal peduncle mod- erate, length 24.3-28.8 per cent SL; depth 9.7-13.8 per cent SL. Dentition subject to ontogenetic and geographic variation (see Maver, 1974: 185 and below). Teeth small, conical; present on premaxillae, mandibles and vomer; palatines edentulous or dentigerous; tongue dentigerous, bearing triangular patch of glossohyal teeth along midline and broad. closely-spaced patches of teeth parallel to margins: endo- and ectopterygoids edentulous. Opercle dominated by pungent, bony spine; surmounted by one or more poorly ossified spinelets. Preopercle well-ossified; angle narrowly produced; ornamentation subject to geographic variation (see below). Interopercle and subopercle weakly ser- rated or unserrated. Gill rakers awl-like, 29-36 on first gill arch, 8-10 on upper portion, 20-25 on lower portion (see Table 2). First dorsal fin. VII (38): second dorsal fin I; 8 (2), 1) 95GeaE I, 10 (1). Anal fin I, 8 (1), Il, 9 (36); ventral fins 1, 5 (40), soft rays extending to level of anus. Pectoral fin 16-19 (see Table 2). DiI variable, 1.7-3.2 per cent SL: Dsl, All, and Pal longvand pungent: 12.7-16.5 per cent, 13.5=16.8 per cent and’ I13:0=l5% per cent SL? respectively. Preserved specimens yellow-brown; mouth light: branchial and peritoneal membranes black: dorsal fin membranes dark. Viscera often sheathed by heavy. vellow-brown fat deposits. Description based on forty specimens 70.9 158.9 mm SL. 1977 E. CONSTANCIAE DESCRIPTION 7 NOTES ON THE HOLOTYPE OF E. CONSTANCIAE As mentioned earlier, Giglioli’s description of Pomatomichthys constanciae 1s incomplete and includes a number of inaccuracies. Tortonese and Queirolo (1970) figured and redescribed the holo- type but did not fully characterize the fish. One aspect of the present investigation involved reexamining Giglioli’s type speci- men, now in rather poor condition. Measurements and counts obtained from the holotype appear in the Appendix. Qualitative features not mentioned in previous descriptions include: 1. Lower jaw slightly prominent with two nubs at symphysis. 2. Small teeth on premaxillae, mandibles, vomer, palatines and tongue: lingual dentition composed of a central tooth patch and a series of elongate tooth patches parallel to the borders of the tongue. 3. Opercular spine well-developed; two poorly developed spine- lets above, but well-separated from the spine. 4. DiI and AI spines short; DoI, AII and PI spines long. Longest ventral fin rays reach anus. If previous FE. trewavasae descriptions (e.g., Mayer, 1974; Poll, 1954) are compared with P. constanciae data presented in the Appendix and above, it becomes apparent that the two forms are synonymous. Giglioli’s holotype bears all characters diag- nostic of EF. trewavasae including VII-I, 9 dorsal fin elements, long fin spines, 18 pectoral rays, numerous awl-like gill rakers, a pungent. bony opercular spine, mandibular chin nubs and lingual teeth. The species is correctly called £. constanciae be- cause Pomatomichthys is a junior synonym of Epigonus, and constanciae has precedence over frewavasae. The fish was named after Giglioli’s wife, Costanza. DISTRIBUTION E. constanciae is found from the coast of equatorial west Arfica to the Straits of Messina. The species has also been taken off Madeira and probably occurs off the Canaries and Azores. E. constanciae appears most abundant on upper portions of the continental slope between 200 and 400 m. 8 BREVIORA No. 443 GEOGRAPHIC VARIATION Mayer (1974: 185-186) briefly commented on geographic vari- ation in Epigonus trewavasae and suggested that Mediterranean and African populations may represent subspecies. At the time, sufficient material was not available to draw firm conclusions about the validity of infraspecific categories. Nevertheless, variations were noted in qualitative characters such as the promi- nence of chin nubs, the development of vomerine and palatine teeth and the occurrence of preopercular armor. The consolidation of Pomatomichthys constanciae and Epi- gonus trewavasae brings the question of geographic variation into sharper focus, because holotypes of the nominal species were taken from opposite ends of the &. constanciae range. An analysis of geographic variation in FE. constanciae was accom- plished by dividing the specimens examined for this study into three geographic groups: a Mediterranean series, a Madeiran series and an African series. The African series was the largest group and included fishes taken along the west African coast from 18°S to 21°N. The lumping of west-central African and northwest African populations follows Maurin’s conclusion (1968: 78) that the Cape Blanc-Arguin Bank region represents a major northern faunal limit for demersal forms with tropical affinities. Table 3. Comparisons of meristic data from African, Mediterranean and Madeiran populations of Epigonus constanciae. Atr. = African popu- lation, Med. = Mediterranean population; Mad. = Madeiran popula- tion. See text for explanation. COEFFICIENTS OF DIFFERENCE Afr. vs. Med. Afr. vs. Mad. Med. vs. Mad. Pectoral fin rays 0.33 0.48 0.20 Gill rakers Total 0.63 1.24 0.26 Upper arch 0.28 0.59 0.14 Lower arch 0.57 1.27 0.39 Lateral line scales 0.50 0.61 0.16 Pyloric caeca 0.12 0.38 0.09 1977 E. CONSTANCIAE DESCRIPTION 9 MERISTIC CHARACTERS Meristic data comparisons were accomplished by calculating Coefficients of Difference (C.D.’s) (Mayr, 1969: 189-193) for the three geographic groups. The index indicates the degree of non- overlap between two populations for a given character. Popula- tions exhibiting a non-overlap of 90 per cent or more (C.D.2 1.28) are considered distinct at the subspecies level. Table 3 presents data from the current study. As indicated, none of the differences observed are significant at the subspecies level. MORPHOMETRIC CHARACTERS Analogous examinations of mensural data were undertaken using regression techniques. Individual values obtained from Mediterranean and Madeiran specimens were compared to re- gression data from African populations by means of Student’s t-tests. Results of the tests are summarized below. Mediterranean and African populations are very similar and do not appear distinct at the subspecies level. Only one Mediter- ranean data point fell outside the 99 per cent confidence limits of the African group and most data points fell within the 95 per cent or 98 per cent limits. The Madeiran population is more distinct. Madeiran E. con- stanciae appear characterized by shallower bodies, shorter heads and shorter jaws than African forms. However, part of the di- vergence may be an artifact of the Madeiran specimens’ poor condition. All were obtained from the stomachs of large, pred- atory fishes and many were considerably damaged. Additional ‘data are required to assess the taxonomic significance of differences exhibited by the Madeiran specimens. QUALITATIVE CHARACTERS Geographic variability reported by Mayer (1974) for qualita- tive characters was confirmed by the present study. Character States such as prominent chin nubs, weakly armored preopercular margins, edentulous palatines and vomerine tooth patches lack- ing posterior median extensions are prevalent, but not uniform among fishes from equatorial west Africa. In contrast, poorly 10 BREVIORA No. 443 developed chin nubs. spinous preopercular margins, dentigerous palatines and vomerine tooth patches with posterior median extensions are common among Mediterranean specimens. The frequency of “northern” and “southern” character states varies with latitude. Madeiran and Mauritanian specimens tend to be intermediate, with the former more closely resembling Mediter- ranean fish and the latter more similar to specimens from equa- torial Africa. Because characters exhibit continuous variation along the range of the species, and because populations at spe- cies’ distribution limits exhibit a mixture of character states and are not uniformly “northern” or “southern”, qualitative characters do not support the division of FE. constanciae into formal sub- species. CONCLUSIONS The picture that emerges for Epigonus constanciae is one of a polymorphic species whose range extends from the eastern equa- torial Atlantic to the western Mediterranean. This pattern has been reported for other organisms by many authors (e.g.. Briggs, 1974; Maurin, 1968: Tortonese. 1964, 1960; Ekman, 1953). The prevalence of polymorphism among Atlantic and Mediterranean populations has been discussed by Tortonese (1964: 103-104; 1959 387). Although genetic differences between populations cannot be ruled out, Tortonese suggests that Atlantic-Mediterranean phe- notypic dissimilarities may be induced by regional differences in temperature regime and food availability. It has been suggested elsewhere (Maver. 1972) that FE. constan- clae may be among the least vagile species of the genus Epigonus. Adults appear strongly associated with the substrate. Neither pelagic larvae nor pelagic juveniles are known for this species, although such forms have been discovered for several congeners. The smallest known specimen of E. constanciae is a 29.8 mm SL juvenile (IRSN 16548) taken by a bottom trawl off Sardinia. The latter data suggest that #2 constanciae may have either a short-lived pelagic phase or may lack a pelagic phase entirely. [he hypothesis of limited vagility fits the pattern of phenotypic variability observed for FE. constanciae. Individual populations adapt to local conditions and evolve characteristic traits. Breeding may be assumed to occur primarily between members of the same or 1977 E. CONSTANCIAE DESCRIPTION 1] neighboring populations, thus preserving local phenotypes. Rela- tively isolated populations, such as that from Madeira, are more unique in appearance but still conform to phenotypic patterns resulting from latitudinal gradients in environmental or genetic factors. ACKNOWLEDGEMENTS This work was initiated while the senior author was an Asso- ciate in Ichthyology at the Museum of Comparative Zoology and was completed during his tenure at the Department of Marine Science, University of South Florida. We are indebted to these institutions as well as to the Museo Civico di Storia Naturale for logistical support and encouragement. The authors also wish to thank the following institutions and individuals for providing study materials: X. Missionne, Institut Royal des Sciences Natu- relles de Belgique; G. Krefft, Institut fur Seefischerei: C. Maurin and J. C. Quero, Institut des Péches Maritimes; K. F. Liem and R. Schoknecht, Museum of Comparative Zoology; G. E. Maul, Museo Municipal do Funchal: B. Lanza and M. Bucciarelli, Flor- ence Zoological Museum; E. A. Lachner, Smithsonian Institu- tion: and C. Karrer, Zoologisches Museum, Berlin. Special thanks goto J. C. Briggs, R. C. Baird, G- Krefft and G. Smith for criti- cally reviewing the manuscript. BIBLIOGRAPHY Briccs. J. C. 1974. Marine Zoogeography. New York: McGraw-Hill Book Gok 475) pp: CAVALIERE. A. 1956. Su rari stadi larvali e giovanili di Pomatomichthys con- stanciae Gigl. e relativa differenziazione da stadi giovanili della specie affine Pomatomus telescopium Risso. Boll. Pesca Piscicult. Idrobiol., (n.s.) //(1): 121-127. EKMAN. S. 1953. Zoogeography of the Sea. London: Sidgwick and Jackson, Ltd. 417 pp. Gictiott, EH. 1880. Elenco dei Mammiferi, degli Uccelli e dei Rettili Ittiofagi Appartenenti alla Fauna Italica e Catalogo degli Anfibi e dei Pesci Italiani. Firenze: Stamperia Reale. 85 pp. Goopr. G. B.. axp T. H. BEAN. 1896. Oceanic Ichthyology Washington. D.C.: Smithsonian Institution. 553 pp. Husps. C. L. anp K. F. LAGLER. 1958. Fishes of the Great Lakes Region Ann Arbor: Univ. Mich. Press. 213 pp 12 BREVIORA No. 443 JORDAN. D. S.. AND B. W. EVERMANN. 1896. The fishes of North and Middle America. Bull. U.S. Nat. Mus., 47(1): 1-1240. as Sey Be . AND H. W. CLARK. 1930. Check List of the Fishes of North and Middle America. Appendix X, Report to the United States Commis- sioner of Fisheries for 1928. Washington, D.C.: U.S. Government Printing Office. 670 pp. MAURIN. C. 1968. Ecologie ichthyologiques des fonds Chalutables atlantiques (de la baie ibéro-marocaine a la Mauritanie) et de la Mediterranée occidentale. Theses presentées a la Faculté des Sciences de Université de Nancy. No. d’Ordre: A. O. 2 182. 145 pp. Mayer. G. F. 1972. Systematics, functional anatomy, and ecology of the cardinalfish genus Epigonus (Apogonidae). Ph.D. Thesis. Harvard Uni- versity. 190 pp. 1974. A revision of the cardinalfish genus Epigonus (Perciformes, Apogonidae). with descriptions of two new species. Bull. Mus. Comp. Zool. 146(3): 147-203. 1975. Results of the research cruises of FRY “Walther Herwig” to South America. XXXIX. The epigonine fauna of the South Atlantic, with a key to the genera and a redescription of Rosenblattia robusta Mead and De Falla. Arch. Fisch. Wiss., 26(1): 13-28. Mayr. E. 1969. Principles of Systematic Zoology. New York: McGraw-Hill, Inc. 428 pp. Port. M. 1954. Poissons. IV. Teleostéens acanthoptérygiens (premiére partie). Rés. Sci. Expéd. Océanogr. Belge Eaux Cét. Afr. Atl Sud, 4(3A): 1-390. Ricker. W. E. 1973. Linear regressions in fishery research. J. Fish. Res. Bd. Canada, 30: 409-434. Scuuttz. L. P. 1940. Two new genera and three new species of cheilodipterid fishes, with notes on other genera of the family. Proc. U.S. Nat. Mus.. 88(3085): 403-423. TORTONESF. E. 1959. General remarks on the Mediterranean deep-sea fishes. In: International Oceanographic Conference, 1959, Preprints, M. Sears (ed.). Washington, D.C.: Amer. Assoc. Adv. Sci. 386-387. 1964. The main biogeographical features and problems of the Med- iterranean fish fauna. Copeia, /964(1): 98-107. 1973. Apogonidae. In: Check-lst of the Fishes of the Northeastern Atlantic and of the Mediterranean. Vol. 1, J. C. Hureau and Th. Monod (eds.). Paris: UNESCO. 365-367. — AND L. C. QuriroLto. 1970. Contributo allo studio dell’ ittiofauna del Mar Ligure orientale. Ann. Mus. St. Nat. Genova, 78: 21—46. 1977 E. CONSTANCIAE DESCRIPTION 13 APPENDIX Meristic and morphometric data obtained from the holotype of Epigonus constanciae, MZF 3089. Measurements are in millimeters. Meristic data Dorsal fin VII-I, 9 Pelvic fin TES Anal fin 11,9 Lateral line 46 Pectoral fin 18 Gill rakers 30 Morphometric data SE 115 Interorbital width 125 HL 37 Maxilla 16 Head height 19.5 Caudal peduncle length 29 Body depth 30 Caudal peduncle depth 13 Eye U7) Dol 16 Snout 8 All 17 Pol 15 SOP ee AR 1 8 jop6 Museum of Comparativg,Zoology US ISSN 0006 KAIMERSITY CAMBRIDGE, MASS. 27 MAY 1977 NUMBER 444 STATIONS OF THE THAYER EXPEDITION TO BRAZIL 1865 — 1866 MYVANWY M. DICK! Asstract. In April of 1865, a group of scientists from the Museum of Compara- tive Zoology led by Louis Agassiz embarked for Brazil. The purpose of this expedi- tion was to study, in situ, the fauna and flora of that country. Members of the ex- pedition, divided into three groups, explored the major part of Brazil from the Rio Tocantins eastward to the coast. Although the major part of the work was done on the distribution and relationships of the fresh water fishes (Agassiz’s prime interest) many superb collections were made of plants, invertebrates and vertebrates. Thor- ough geological surveys provided much information on the structure of Brazil. This paper traces and maps the numerous stations where specimens were collected. Based on modern and nineteenth century maps, and on notes and papers recording the routes followed and collections made by the expedition, it represents a compilation of all this information into a single, indexed directory. This directory should facili- tate the future labors of individuals working with the Thayer Expedition material. INTRODUCTION Louis Agassiz had for many years wanted to observe in situ the fauna and flora of Brazil, a study that he knew would be substan- tially encouraged by the Brazilian government. The opportunity to do so, however, had never presented itself. This cherished dream became a reality in 1865 through the understanding and generosity of Nathaniel Thayer, a Boston businessman keenly interested in science. Thayer assumed the expenses of Professor and Mrs. Agas- siz and six assistants for an expedition to Brazil lasting several months. The scientific staff from the Museum of Comparative Zoology ‘Museum of Comparative Zoology 2 BREVIORA No. 444 included John G. Anthony, Assistant Curator of Molluscs (who was with Agassiz in Rio de Janeiro and made extensive collections in this area before illness forced his return to Cambridge); Joel Asaph Allen, Assistant Curator of Birds (later the noted Curator of Birds at the American Museum of Natural History); Frederick C. Hartt and Orestes St. John, geologists trained by Agassiz; George Sceva, Preparator and collector primarily of fossil material; and James Burkhardt, an artist who had worked for many years with Aggassiz. In spite of poor health, Burkhardt was an indefatigable worker and made, according to Agassiz, over eight hundred illus- trations during the expedition. Most of these unfortunately have disappeared, but the remaining examples, which are in the Museum of Comparative Zoology, indicate the high quality of his draft- manship. Agassiz also had extremely able and enthusiastic young volun- teers in his party: Edward Copeland, who accompanied Hartt from Rio de Janeiro to Bahia; Newton Dexter; Walter Hunnewell; William James, the future great professor of philosophy at Harvard University; Stephen Thayer, the son of Nathaniel Thayer; and Thomas Ward. On 2 April 1865 the group left Boston aboard the S.S. Colorado, guests of the Pacific Mail Steamship Company. While at sea, Agassiz gave a series of lectures giving background for the work planned for the months ahead. On 23 April 1865 the S.S. Colorado docked in Rio de Janeiro. Agassiz had arrived at the threshold of his longed-for goal. The area covered by the members of the expedition was extensive. They divided into three groups and explored the major part of Brazil from the Rio Tocantins eastward to the coast; they also explored some of its tributaries, to the borders of Colombia and Peru. Al- though the major part of the work was done on the distribution and relationships of the fresh water fishes (Agassiz’s prime interest), many superb collections were made of plants, invertebrates and vertebrates. Thorough geological surveys provided much infor- mation on the structure of Brazil. The numerous stations where specimens were collected have been difficult to locate because several were small villages or clusters of dwellings on the edge of a river or lake. Many no longer exist and some have names other than those used in 1865. Rivers and 1977 THAYER EXPEDITION 3 lakes that were recognized a hundred years ago are frequently overlooked in recent atlases. Researchers involved with the Thayer Expedition have devoted many hours to tracing these stations and collections. Using both modern and nineteenth century maps, it has been possible to locate them fairly accurately. Information concerning the routes followed and the collections made by mem- bers and associates of the expedition was obtained from publica- tions listed in the bibliography. The locality names used in the text and on the maps are almost always those employed during the period of the Thayer Expedition. Subsequent changes in locality names or spelling are italicized in the index. Having this informa- tion incorporated in a single directory should facilitate the future labors of individuals working with the Thayer Expedition material. Much work needs to be done with the material collected by Agassiz and his corps, especially with the collections of fishes. Many other groups, both plant and animal, also need comprehen- sive study and evaluation to complete the survey of Brazil envis- aged by Louis Agassiz so long ago. MEMBERS OF THE THAYER EXPEDITION 1865 — 1866 MUSEUM STAFF: Louis Agassiz Elizabeth Cary Agassiz Joel A. Allen Ornithologist John G. Anthony Malacologist James Burkhardt Artist Frederick C. Hartt Geologist George Sceva Preparator Orestes St. John Geologist VOLUNTEERS: Edward Copeland Newton Dexter Walter Hunnewell William James Stephen Van R. Thayer Thomas Ward 4 BREVIORA MEMBERS ADDED IN BRAZIL Major M. Coutinho Monsieur D. Bourget Mr. Talisman COLLECTIONS ALSO MADE BY: Captain Anacleto Senhor Augustinho Senhor Barrosa Colonel Bentos Senhor S. E. Pimento Bueno Dr. Pacheco de Silva Major Estolano Senhor Felice Senhor Joao Baptista da Fonseca Senhor Glaziou Senhor Honorio Dr. Justa Mr. Kaulfuss Senhor Antonio de Lacerda Senhor Mariano P. F. Lage Senhor Cicero de Lima Dr. Couto de Magalhaes Dr. Malcher Dr. Mendes Senhor Penna Senhor Joachim Rodriguez Senhor Sepeda Mr. Charles Taylor Father Torquata Senhor Vinhas No. 444 Brazilian Government Corps of Engineers French naturalist residing in Rio de Janeiro Officer, Amazonian Steamship Company Fishes, Amazon River Fishes, Amazon River Miscellaneous, Amazon River Fishes, Rio Trombetas Forest and river fishes, Amazon River Fishes, Rio de Janeiro Fishes, Amazon River Geological observations, miscel- laneous, Ceara Fishes, mammals, environs of Rio de Janeiro Plants, palms, Organ Mountains Miscellaneous, Amazon River Fishes, Rio Parahyba do Norte Andean fossils Miscellaneous, Bahia | Plants, animals, fishes, Rio Novo, | basin Rio Parahyba | Fishes, insects, interior Brazil Miscellaneous, at his direction, upper Amazon River Birds, Amazon River Miscellaneous, Ceara Fishes, Amazon River Fishes, Santarém Miscellaneous, Amazon River Drawings and collections of fishes and insects, Posse Fishes, Amazon River Fishes, Rio Xingu 1977 THAYER EXPEDITION 5 The Emperor, Dom Pedro II, had extensive collections of fishes made for Professor Agassiz from several rivers in southern Brazil. Thomas G. Cary, Mrs. Agassiz’s brother, though not connected with the expedition, contributed material from Montevideo, Buenos Aires and other localities. ACKNOWLEDGMENTS The suggestions made by Dr. Ernst Mayr and Dr. Raymond A. Paynter, Jr.. Museum of Comparative Zoology, concerning the presentation ef material have been greatly appreciated. The gen- erous assistance of Dr. P. E. Vanzolini, Museu de Zoologia da Universidade de Sao Paulo, has been invaluable and I am indeed grateful. Dr. Naercio Menezes, also of the Museu de Zoologia, kindly located six stations that had completely eluded me. To Dr. Bruce B. Collette, United States National Museum, I am in- debted for his critical reading of the manuscript. These kind people all have my sincere thanks. THE THAYER EXPEDITION Louis Agassiz established a laboratory in Rio de Janeiro which the members of the expedition used as a base for their work during the weeks prior to their departure for other areas of Brazil. Nu- merous brief trips were made at this time, including one over the route of Dom Pedro II Railroad, which Agassiz believed would be of great future value for the transportation of large geological and zoological specimens. Zoologists and geologists thoroughly investigated the railroad from its origin in Rio de Janeiro to its terminus in Parahyba, a distance of one hundred miles. On 27 April Professor and Mrs. Agassiz enjoyed their first journey on the railroad as guests of Major Ellison, chief engineer. Two speci- mens from the fresh waters of Brazil were collected at this time, inaugurating what was to be an outstanding assemblage of Bra- zilian fishes. An excursion of several days duration was made by the Agassizes in May, 1865; it was the first extending any distance from Rio de Janeiro. They went by ferry to Maua then by railroad and post coach to Petropolis. From Petropolis they continued along the valley of the Rio Piabanha to Posse, crossed the Rio Parahyba at 6 BREVIORA No. 444 Entre Rios, soon reached the Rio Parahybuna, then went on to Juiz de Fora. This trip provided an excellent opportunity to study the plant life and geological formations and to collect numerous speci- mens of fishes and other vertebrates. Examples of the peculiarities of Brazilian geological formations were observed in Tijuca, about eight miles from Rio de Janeiro, 26-27 May. On 21 June the Agassizes again went to Juiz de Fora, arrived there 22 June, then continued thirty miles farther where they were guests of Senhor Lage at his fazenda. Senhor Lage ac- companied the Agassizes to the Serra da Babylonia. On 27 June they returned to Rio de Janeiro. In June the party divided into units to facilitate the exploration of Brazil. St. John, Allen, Ward and Sceva left for the interior the first week of June and Hartt and Copeland left to explore the coast from the Rio Parahyba to Bahia the following wee. On 25 July, the Agassizes, Major Coutinho, Burkhardt, Monsieur Bourget (a French naturalist residing in Rio de Janeiro engaged by Agassiz as a collector and preparator), Hunnewell and James left Rio de Janeiro on the “Cruzeiro do Sul” for Bahia, where they arrived 28 July. In Bahia they were joined by Dexter and Thayer, who had preceded them. Antonio de Lacerda, their host, had greatly assisted Dexter and Thayer in their collecting. The next collecting station was Maceio on 30 July, then Pernambuco 31 July and on 2 August the Parahyba do Norte as far as the town of Parahyba. They arrived at Ceara 5 August, the guests of Dr. Mendes, and were at Maranhao 6 August. They arrived at Para 10 August, where Mr. Pimenta Bueno arranged living and working quarters from 10 August to 19 August, and the use of a steamer, the “Icambiaba”, for a month between Para and Manaos. In Para, Mr. Talisman, a young Brazilian officer of the Amazonian Steam- ship Company, joined the group. Breves, 20 August, was the first station in Amazonas. Leaving Aturia, they passed into the Rio Tajapurti and stopped at the village of Tajapuru. On 21 August, after two days spent skirting the island of Maraj6, they left the Para River and turned into the mainstream of the Amazon. They went to Gurupa, 22 August, where a collection of forest fishes was made, and on to Porto de Moz on the Rio Xingu, 23 August. A brief non-collecting halt was made at Prainha on 24 August, and they were at Monte Alégre, at the mouth of the Rio Gurupatuba on 25 August. 1977 THAYER EXPEDITION 7 From Santarém, at the mouth of the Rio Tapajoz, Dexter, James and Talisman left, 26 August, to explore the Rio Tapajoz. Bour- get and Hunnewell remained in Santarém, Bourget to make col- lections, Hunnewell to repair his photographic equipment. Agas- siz, Burkhardt, Thayer and Coutinho continued on to Obydos. Before leaving the Rio Tapajoz to join the Amazon, the “Icambiaba” detoured through a narrow channel, Igarapé-Assu. A brief stop was made at Obydos, 26 August. On 27 August they landed at Villa Bella, at the mouth of the Rio Tupinambaranas. By canoe, the group left Villa Bella to explore Lago José Asst until 30 August, proceeding from there to a branch of the Rio Ramos, which con- nects the Amazon to the Rio Madeira. They returned to Villa Bella on 2 September. A rest period was taken from 4 to I! September at Mandaos, on the Rio Negro. There they were joined by Dexter, James and Talis- man. A brief halt was made at Barreira das Cudajas on 12 Septem- ber and on 13 September they made another short halt at Coari. Teffé stands beside a small lake formed by the Rio Teffé before it joins the Amazon. This site was visited on 14 September, then Fonte Boa, 15 September and Sao Paulo de Olivenga, 17 Septem- ber. Here James and Talisman left the group to continue from there to the Rios Iga and Hyutahy by canoe. The farthest point reached by Agassiz was Tabatinga, 19 September. Bourget re- mained in Tabatinga for a month, collecting there and in the sur- rounding country. Agassiz and the rest of the party returned to Teffé, remaining there until 22 October. Agassiz and Coutinho were at a Station on a branch of the Rio Solimoens from 4 October to 6 October. James and Talisman rejoined them at Teffé, 16 Oc- tober and Bourget boarded the “Icambiaba” 22 October. While in Teffé, Dexter made an extensive collection of birds and, with the assistance of Hunnewell, Thayer and local sportsmen, added to the reptile and animal specimens. Agassiz also purchased a large collection of insects at this station. Manaos was reached on 23 October. While at this station, Agas- siz, Coutinho, Burkhardt, Dexter and James, preceded by their host, Senhor Honorio, had a two day, 27 October to 29 October, collecting excursion to Lago Hyanuary on the western side of the Rio Negro. Talisman and Dexter explored the Branco and Negro rivers for six weeks, returning to Manaos 10 December. Thayer and Bourget were at Lago Cudajas for ten days, 27 November to 8 BREVIORA No. 444 6 December (Thayer also collected at Lago Alexo), and James went to Manacapuru for ten days. The Agassizes, Coutinho and Burkhardt left Manaos 10 De- cember to spen ten days at Mauhes. They proceeded down the Rio Ramos to the Rio Mauhes, which runs almost parallel with the Amazon, to the Rio Madeira (which joins the Amazon oppo- site Serpa). The land encircled by these rivers is the island of Tu- pinambaranas. Mucaja-Tuba and another small settlement on the Rio Mauhes were visited 13-15 December, and on 21 Decem- ber the group returned to Manaos. On 26 December the party left Mandos and went up the Rio Negro as far as Pedreira on the “Ibicuhy,” arriving there 28 De- cember. A collection of palms had been made at Taua Péassu and they were picked up on the return to Manaos 31 December. The stay at Mandaos ended about 12 January 1866 and the party returned to Villa Bella 16 January. Collections were made 18 Janu- ary at Lago Maximo, a short walk overland from Villa Bella. A narrow outlet connects this lake to the Rio Ramos. On 21 January the party was again at Obydos, and 22 January at Santarém, which is situated on a point of land separating the Rio Tapajos and the Amazon. They left Santarém 24 January for Monte Alégre, where they stayed until 29 January. Agassiz, Coutinho and a few friends went to the Serra d’Ereré, northwest of Monte Alégre, to see the geological formations. Coutinho went by horseback and Agassiz by canoe, along the Rio Gurupatuba, up a narrow stream known as the Rio Ereré to a point on a line with the Serra and from there proceeded on foot. Senhor Vinhas, a resident of Porto de Moz, had a collection of fishes from the Rio Xingu ready for Agassiz upon his arrival 29 January, so the group quickly continued on to Gurupa, arriving 30 January. They reached Tajapuru 31 January and spent two days there. From 4-27 February they were at Para. On 28 February they left on the steamer “Tabatinga” to explore the great island of Marajo. They visited Soures, then, 29 February to 4 March, Vigia, Baia do Sul and the small island of Tatuatuba. They returned to Para 5 March. On 26 March the party left Para for Ceara where they arrived 31 March. A stop was made en route at Maranhao so that Agassiz and Coutinho might examine the coastal geology with more care 1977 THAYER EXPEDITION 9 than on their previous visit. An excursion to Pacatuba and the Serra da Aratanha, 6-12 April, included the Agassizes, Coutinho and Senhor Pompeo, Government Engineer of the Province. Col- lections were made in the vicinity of Ceara by various people in- cluding Senhor Felice and Senhor Cicero de Lima. They left Ceara 16 April and continued, with a brief stop at Pernambuco, to Rio de Janeiro, where they arrived 25 April. During the time in Rio de Janeiro, 25 April to 2 July, collections were packed and shipped to Cambridge. A few short trips were made, to Petropolis and again along the Dom Pedro II Railroad. The last excursion made by Agassiz was 9-12 June, to the Organ Mountains, accompanied by Mrs. Agassiz, Mr. Glaziou and Dr. Nageli. The group went by boat to Piedade, then walked from there to Theresopolis, collecting along the way. On 2 July Burkhardt, Hartt, Copeland, Sceva, Dexter, James, Hunnewell and Thayer embarked for the United States. Allen, Ward and Sceva left Rio de Janeiro 9 June 1865 under the leadership of Orestes St. John. Their route led to Juiz de Fora, then across the Serra da Mantiqueira to Barbacena (where Ward left the group) through Lagoa Dourada, across the Rio Caranda- hy and the Rio Paraopeba just above the water gap of the Serras da Piedade and Itatiaiassu, to the village of Morro Velho. They tra- versed the basins of the Rio Parahyba, Rio Grande (Rio de La Plata) and Rio Sao Francisco. By way of Sabura, Santa Luzia, Lagoa Sancta and Sette Lagoas they reached Gequitiba. Sceva remained in Lagoa Santa to search for fossils in the caves of the region, but a previous collector had taken most of the ma- terial. He did, nonetheless, prepare an excellent series of mam- mals before returning to Rio de Janeiro. After a few days in Rio de Janeiro, devoted to preparing and packing specimens collected by other members of the expedition, Sceva went to Cantagallo to collect until he rejoined Agassiz in Rio de Janeiro to return to the United States. St. John and Allen left Gequitiba 31 July 1865 and travelled by canoe down the Rio das Velhas, arriving at the junction of the Rio Sao Francisco 28 August. After three days here they continued by boat to Januaria, reaching there 9 September and remaining until 19 September. Allen, in iil health, decided at this point to continue on alone to Bahia, taking with him the material so far collected. Still trav- 10 BREVIORA No. 444 elling by boat he followed the Rio Sao Francisco past Urubt to Chique-Chique, where he arrived 8 October and stayed for several weeks. From Chique-Chique his route led overland to the coast by way of Engenho Velho, Jacaré and Olhos d’Agua, across an arid plain, to Taboleiro de Jacobina, down the “Tombador” (a precipitous defile) to Jacobina valley. From Jacobina he went to the Serra da Terra Dura, Arraial do Riacho do Jacuhype, Feira de Sta. Anna, Espelto (Espelho?) and Cachoeira, then to Bahia, a journey of many weeks. He did not reach Bahia until the end of November. On 15 December Allen left Bahia for Cambridge. St. John followed the Rio Séo Francisco to Barra, then went along the valley of the Rio Grande to the Rio Préto, Santa Rita and Paranagua. Several days were spent at Paranagua before con- tinuing on along the valley of the Rio Gurgueia to Manga. The party followed the Rio Paranahyba to Sao Goncallo where many specimens (primarily birds, reptiles and insects) were taken. There- zina, capitol of the state of Piaui, was the next station and excellent collections of fishes were made from the Rio Poty, a tributary of the Rio Parnahyba. From Therezina St. John continued on to Caxias, proceeded to Maranhdo by way of the Rio Itapicuru and arrived at Maranhao 8 January 1866. Illness kept St. John in Maranhfo until February, when he left to join Agassiz in Para. His geological observations were as careful and as valuable as his zoological collections. After Ward separated from his associates at Barbacena he con- tinued on to Ouro-Préto, Mariana and Santa Barbara, then down the Rio Piracicaba to the Rio Doce. He followed the river almost to the point where it meets the Rio Antonio. Crossing the Serra das Esmeraldas he arrived at the basin of the Rio Jequitinhonha and explored several branches of the river after passing Diaman- tina. He proceeded cross-country to the town of Rio Pardo and the Rio Pardo. He crossed the Rio SAo Francisco at Januaria, then continued northwestward until he reached the Rio Tocantins, which he followed to Para. From Para Ward returned to the United States. Hartt visited Brazil twice, first as a member of the Thayer Expe- dition, and, later, on a private expedition. Although he and Cope- land made extensive collections, particularly of marine inverte- brates and fishes, their major interest was the geology of Brazil. This has been comprehensively treated by Hartt, both from per- 1977 THAYER EXPEDITION 11 sonal observation and from the work of other geologists, in his book, The Scientific Results of a Journey in Brazil, 1870. Con- siderable information is also included on the common species of plants and animals and areas of fossil beds. Hartt and Copeland did not go directly from Rio de Janeiro to Bahia as planned. Their first attempt took them as far as Nova Almeida north of Victoria, but they were forced to return to Rio de Janeiro for lack of mules and money. After again working near Rio de Janeiro they embarked on a sailing vessel for Sado Matheos, stopping wherever possible along the coast. From Sao Matheos they went to the Rio Doce, returned to its mouth and then back to Sado Matheos. They proceeded north to Belmonte via the Rio Jequitinhonha and then returned to Porto Alegre, then north to Bahia. Rather than complicate the survey of the area covered by Hartt and Copeland the following route description is given (with the exception of their return to Porto Alegre) in direct sequence. The environs of Rio de Janeiro were thoroughly explored by Hartt and Copeland. They examined the area of the Serra do Mar, which borders the coast from the state of Sao Paola to the state of Rio de Janeiro, and also the Serra da Mantiqueira, partly separated from the Serra do Mar by the valley of the Rio Parahyba do Sul. Within the bay of Rio de Janeiro the Ilha de Paqueta, Ilha das Cobras, Ilha das Enxadas and Ilha do Governador were explored. They visited several places near Rio de Janeiro, includ- ing the hills that run westward to Lagoa de Freitas, separated from Corcovado by the valley of Botafogo. They also explored Trés Irmaos, southwest of Corcovado, the Gaira to the west and the Tijuca range to the north, which is separated from Corcovado by the pass of Boa Vista. They covered the entire length of the Dom Pedro II railroad. The Rio Macacu was followed to Porto Villa Nova, then along the Cantagallo Railway to Porto das Caixas and the terminus at Cachoeria. Between Rio Janeiro and Cabo Frio observations were made at Marica, Lagoa de Marica, Lagoa de Saquarema (east of Porto Negra) and Lagoa de Araruama. After Cabo Frio they stopped at Os Buzios, Cabo de Sao Thomé, Macahé, the Ilhas de Santa Ana (which are a few miles offshore from Macahé) and Campos on the Rio Parahyba do Sul. Just south of Campos is the Lagoa Feia into which the Rio Macacu empties. Several lagoons have formed 12 BREVIORA No. 444 along this coast, including large ones such as Rio Iguassu and Lagoa do Campello. They followed the Rio Muriahé, a tributary of the Rio Parahyba, for several miles then followed the Rio Parahyba from Campos to Sao Fidélis. From here they travelled northward through the valley of Vallao Grande to Bom Jesus on the Rio Itabapuana, which divides the states of Rio de Janeiro and Espiritu Santo. They went down the Rio Itabapuana, past Porto da Limeira, to the mouth of the river. Hartt and Copeland continued northward to Lago Maroba, then along the coast to the town and river of Itapémirim and the surrounding area, then on to Piuma, the Rio Benevente and Guara- pary. Stops were made at Ponta da Fructa and Ponta de Jecu en route to the bay of Espiritu Santo, Villa Telha (on the bay of Espi- ritu Santo) and Victoria. Fifty miles northwest of Victoria they ex- plored the Rio Santa Maria and Sao Leopoldina then went on to Carapina, Santa Cruz and the Rio Doce, which they traversed from its mouth to Porto do Souza. They went down the river to Linhares and Lagoa Juparanda (connected to the Rio Doce by a deep chan- nel called the Rio Juparanaa). Returning to the coast they continued northward, stayed briefly during November 1865 in the vicinity of Barra Secca, then passed the Lagoa Marircu and the Rio Azeites at As Azeites on the way to Sao Matheos. Hartt journeyed to the Fazenda do Capitao Grande on the Rio Braco do Norte, which joins with the Braco do Sul to form the Rio Sao Matheos. On the return trip he mapped the river as far as SA€o Matheos. Further north along the coast they noted the Rio Itahunas. From Porto Alégre they went up the Rio Mucury to Santa Clara and from that point travelled westward, using the Minas road, to the headwaters of the Rio Mucury near Poté. Farther west they passed the watershed dividing the Rio Mucury and the Rio Jequi- tinhonha. They arrived at the valley of Jequitinhonha in April 1866 via the Rio Setubal. A detour was made from the Rio Setubal to Alto do Bois before they proceeded cross-country to Calhao. After arriving at Calhao, Hartt made a visit to Minas Novas, re- turned to Calhao, and with Copeland, went down the Rio Aras- suahy and the Rio Jequitinhonha to Belmonte. From Belmonte, Hartt and Copeland returned to Porto Alégre, before again follow- ing the coast north. 1977 THAYER EXPEDITION 13 Observations were made of the terrain between the Mucury and Peruhype rivers. They visited Sao Leopoldina, about ten miles upstream, and Villa Vicosa, four miles below. The route continued northward along the coast to the Rio Caravellas, Prado, the Rio Jucurucu, Porto Seguro and Santa Cruz. After again reaching Belmonte they went on to Cannavieiras and the Rio Pardo. They ascended the river to the head of navigation, passing Cachoeirinha do Rio Pardo and the fazenda of Sisterio. They stopped briefly in Ilhéos on the Rio Cachoeira and in Camamu, en route to Bahia. From Bahia they ascended the Rio Paraguassu as far as Cachoeira, passing the valley of Iguapé. The area surrounding Bahia received close attention before Hartt and Copeland returned to Rio de Janeiro, where they joined Agassiz and other members of the expedition for departure to the United States. 14 BREVIORA No. 444 INDEX Name Map Abrolhos, Parcel dos 3A Alto dos Bois 3A Amarante (San Gongallo) 5 Antonio (Santo Antonio), Rio ) Arancho 11 Arary (Arari) 11 Arassuahy (Aracuai), Rio 3A Armacao 3B Arraial do Riacho do Jacuhype 6 Arupuana, Rio 9 As Azeites 3B Assu Grande 9 Aturia, Rio (Furo do Aturid) 9 Azeites, Rio 3B Bagro do Rio das Velhas 4,6 Bahia, (Salvador) 3A, 4,6; 11 Barbacena 4,7 Barra (/tabapoana) 3B Barra (Villa do Barra) 4,6 Barra Secca (Séca) 3B Barreira das Cudayas, (Codajas) 10 Belmonte 3A, 6 Belmonte, (Jequitinhonha), Rio 3A Benevente, Rio 3B Boa Vista, Pass | Bom Jardin (Jardim) 6 Bom Jesus (Bom Jesus do Itabapoana) 3B Botafogo l Braco do Norte, Rio 3A Braco do Sul (Mantena), Rio 3A Branco, Rio 10 Breves 9 Cabo de Sao Thomé (Tome) 3B Cabo Frio 3B, 6 Cabralia (Santa Cruz) 3B Cachoeira (Caxoeira) 3A, 6 Calhao, (/tira) 3A 1977 THAYER EXPEDITION Name Camamu Cameta Campos Canindé, Rio Cannavieiras Cantagallo (Cantagalo) Cantagallo (Cantagalo), Railroad Capitao Grande Carandahy (Carandai), Rio Caravellas (Caravelas), Rio Carolina Caxias Caxoeira (Cachoeira) Caxoeira, Rio Ceara (Fortaleza) Chique Chique (Xique Xique) Coari Codajds (Barriera das Cudajas) Coeira, Rio Colonia Leopoldina (Sao Leopoldina) Contas, Rio de Corcovado Cudayas Curucga Curupira, Lago Diamantina Doce, Rio Dom Pedro II Railroad Ega (Teffe) ( Tefé) Engenho Velho Entre Rios Ereré, Rio Espelto (Espelho?) Espiritu Santo, Bay Estacao d’Entre Rios (Trés Rios) Fatuarana Fazenda Sen. Lage Feira de Sant’Anna (Santana) LOS) Oo D>} > COW PR WN WN W NA W OO > ee, Ww > > 16 BREVIORA Name Fonte Boa Formosa Fortaleza (Ceara) Furo do Aturia (Rio Aturia) Furo do Tajapuru (Rio Tajapuru) Gaira Gavia (Gadvea) Gequitiba (Jequitiba) Gongogi, Rio Grande, Rio Guarapary (Guaraparl) Guaxindiba or Itahunas (/taunas), Rio Guruguéia (Gurguéia), Rio Gurupatuba (Parieutuba), Rio Gurupu (Gurupa) Hyanuary, Lago Hyapura (Japura), Rio Hyavary, (Javary) (Javari), Rio Iga, Rio Igarapé-Assu (Acu) Igarapé Grande Iguapé, Rio Iguassu (/guacu), Rio Ilha das Cobras Ilha das Enxadas Ilha de Marajo Ilha do Governador Ilha Grande do Jutai Ilha Paqueta Ilhas de Santa Ana (Santana) (Anna) Ilhéos (//héus) Ttabapoana (Barra) Itabapuana (Jtabapoana), Rio Ttacoatiare (Serpa) Itapémerim (/tapemirim), Rio Itapicuru, Rio Itatiaiassu (/tatiaiugu), Serra do Ttaunas (Guaxindiba or Itahunas), Rio No. 444 a. OCnWWWaAW PL — Oo mw Ww W— COK O — mm Prw fon 1977 THAYER EXPEDITION Name Itim, Rio Jacaré, Rio Jacobina Jacuipe (Jacuhype), Rio Januaria Japurd (Hyapura), Rio Jatuarana Javary (Javari) (Hyavary), Rio Jequitiba (Gequitiba) Jequitinhonha (Belmonte), Rio José Asst (Acu), Lago José Fernandez, Lago Jucurucu, Rio Juiz de Fora Juparanaa, Rio (Lagoa Juparana) Jutahy (Jutar), Rio Lagoa de Araruama Lagoa de Freitas (Lagoa Rodrigo de Freitas) Lagoa de Marica Lagoa de Saquarema Lagoa do Campello (Campelo) Lagoa do Maximo (Lago Maximo) Lagoa Dourada Lagoa Feia Lagoa Juparana (Rio Juparanaa) Lagoa Maricu Lagoa Sancta (Santa) Lago Cudajas (Codajas) Lago Curupira Lago do Bota Lago Hyanuary (January) Lago Irandubo Lago January (Hyanuary) Lago José Assu (Agu) Lago José Fernandes Lago Manacapuru Lago Maroba Lago Saraca 18 BREVIORA No. 444 Name Map La Plata (Grande?), Rio 6 Linhares 3B Macacos, Rio dos 9 Macacu, Rio 2 Macahé ( Macaé) 3B Maceio 11 Madeira, Rio 9 Mamuru (Tupinambaranas), Rio 9 Manacapuru 10 Manaos (Manaus) 9, 10 Manga 5 Manhuassu (Manhuagu), Rio 3B Mantena (Braco do Sul), Rio 3A Marajo, Ilha de 9 Maranhiao (Sdo Luts) 1] Marafion, Rio 10 Mariana / Marica, Lagoa do, Ilhas 3B Mateus, Rio 3A Maua (Bardo de Mauda) 2 Mauhes (Maués), Rio 9 Minas Novas 3A Monte Alegre 9 Morro Velho 4,6 Moura (Pedreira) 10 Mucayja-Tuba (Mucajd) 9 Mucury (Mucuri), Rio 3A Mucury Pequeno, Rio 3A Mugiquisaba (Mogiquicaba) 3A Muriache (Muriahé) (Muriaé), Rio 3B Mutum, Rio 3B Nazareth 9 Negro, Rio 10 Novo, Rio 3B Obydos (Obidos) 9 Olhos d’Agua 6 Organ Mountains (Serra dos Orgdos) 2 Os Buzios (Cabo dos Buzios) 3B 1977 THAYER EXPEDITION Name Ouro-Préto (Ouro Préto) Pacatuba Paqueta, Ilha de Para (Belém) Parahyba (Paraiba) do Norte, Rio Parahyba (Paratha) do Sul, Rio Parahybuna (Paraibuna), Rio Paramirim, Rio Paranagua j Paraopeba, Rio Parcel dos Abrolhos Pardo Pardo, Rio Parguassu (Parguacu), Rio Parintins (Villa Bella) Parnatba (Parnahyba), Rio Pedreira (Moura) Pernambuco (Recife) Peruhype ( Peruipe), Rio Petropolis Philadelphia (Teofilo Otoni) Piabanha, Rio Piauhy (Piaut), Rio Piedade Piracicaba, Rio Pirahy (Pirat), Rio Po-Asst (Canal de Poacu) Ponta da Fructa (Fruta) Ponta de Jecu Ponta Negra Porto Alegre Porto da Limeira Porto das Caixas Porto de Moz Porto do Souza Porto Nacional Porto Seguro Posse 20 BREVIORA No. 444 Name Map Poté 3A Poty (Puty) (Por), Rio 5 Prado 3A Prados 4 Prainha 9 Préto, Rio Pt Puty (Poti), (Poty), Rio 5 Ramos, Rio (Parand do Ramos) 9 Recife (Pernambuco) 11 Rio Antonio (Santo Antonio) a Rio Arupuana 9 Rio Arassuahy (Aracuat) 3A Rio Azeites 3B Rio Aturia (Furo do Aturid) 9 Rio Belmonte (Jequitinhonha) SAU, Rio Benevente 3B Rio Brago do Norte 3A Rio Brago do Sul (Rio Mantena) 3A Rio Branco 10 Rio Caninde 5 Rio Carandahy (Carandat) 4 Rio Caravellas (Caravelas) 3A Rio Caxoeira 3A Rio Coeira 3A Rio das Velhas SA ASO, Rio de Contas 3A Rio de Janeiro 1, 3,745 6,57 Rio Doce 3Be7) Rio dos Macacos 9 Rio Ereré 9 Rio Gongogi 3A Rio Grande 6 Rio Grande (La Plata?) 4 Rio Guaxindiba or Itahunas (/taunas) 3B Rio Gurgueia (Gurugueia) 5) Rio Gurupatuba ( Parieutuba) 9 Rio Hyapura (Japurd) 10 Rio Hyavary (Javary) (/Javari) 10 1977 THAYER EXPEDITION Name Rio Hyutahy Rio Iga Rio Iguapé Rio Iguassu Rio Itabapuana (/tabapoana) Rio Itanheim Rio Itapemérim Rio Itapicuru Rio /taunas“(Itahunas or Guaxindiba) Rio Itim Rio Jacaré Rio Jacuipe (Jacuhype) Rio Japurd (Hyapura) Rio Javary (Hyavary) (Javari) Rio Jequitinhonha (Belmonte) Rio Jucururt Rio Juparanaa (Lagoa Juparana) Rio Jutahy (Juraz) Rio Macacu Rio Madeira Rio Manhuagu (Manhuasst) Rio Mantena (Braco do Sul) Rio Marafion Rio Mateus Rio Mauhes (Maueés) Rio Mucury (Mucuri) Rio Mucury Pequeno Rio Muriahé (Muriaé) (Muriache) Rio Mutum Rio Negro Rio Novo Rio Paraguassu Rio Parahyba (Paraiba) do Norte Rio Parahyba (Paraiba) do Sul Rio Parahybuna (Paraibuna) Rio Paramirim Rio Parnahyba ( Parnatba) Rio Paraopeba Swill 21 22 BREVIORA Name Rio Pardo Rio Parnatha (Parnahyba) Rio Peruhype ( Peruipe) Rio Piabanha Rio Piauhy Rio Piracicaba Rio Pirahy Rio Piuma Rio Poty (Poti) (Puty) Rio Préto Rio Ramos (Paranda de Ramos) Rio Samambaia (Sambaia) Rio Santo Anténio (Antonio) Rio Sao Francisco Rio Sao Matheos (SG Mateus) Rio Setubal Rio Solimoens (Solim6es) Rio Tajipuru (Furo do Tajipurit) Rio Tapajoz (Tapajos) Rio Tocantins Rio Trombetas Rio Tupinambaranas (Mamuru) Rio Una Rio Vacaria Rio Xingu Sabura (Sahara) Salvaterra Samambaia (Sambaia), Rio San Gongallo (Amarante) San Luiz (Maranhao) San Mateo (S@o Mateus) (Matheos) San Paolo (S@o Paulo de Olivenga) Santa Anna Islands (//has de Santana) Santa Barbara Santa Caterina (Catherina) Santa Clara Santa Cruz (Cahralia) Santa Luzia WN wNm'! Sys 2 oe) WO OonNOCDOCFWWWNANUON NWN N ee) 4 o> mD> w Ny oO Ff ON n > Re (0) // 1977 THAYER EXPEDITION Name Santarém Santa Rita de Cassia (Villa da Santa Rita) Santo Antonio (Antonio), Rio Sao Fidélis Sao Francisco, Rio Sado Leopoldina (Col6nia Leopoldina) Sao Matheos (S@o Mateus) Sao Paulo de Olivenca (San Paola) S40 Tomé, Cabo de Serpa (/tacoatiare) Serra da Aratanha Serra da Babylénia ( Babilonia) Serra da Mantiqueira Serra da Piedade Serra das Esmaraldas Serra da Terra Dura Serra de Cupati Serra de Monguba Serra d’Ereré (Ereré) Serra do Itatiaiassu (/tatiaiucu) Serra do Mar Serra dos Orgdos (Organ Mountains) Sette Lassas | Setubal, Rio Silva Sisterlo Solimoens (Solimdes), Rio Soures (Soure) Tabatinga Taboleiro de Jacobina Tajipuru, Rio (Furo do Tajipuru) Tapajoz (Tapajos), Rio Tatuatuba Taua Péassu (Tauapegacu) Teffé (Ega) (Tefé) Teofilo Otoni (Philadelphia) Teresopolis (Theresopolis) Therezina (7eresina) Map 3B SAS 4595027 3B 3B 10 3B 9 11 3B 3B, 4, 7 4,6 24 BREVIORA Name Tijuca Tocantins, Rio Tombador Tonantins Trés Irma&os Trés Rios (Estagao d’Entre Rios) Trombetas, Rio Tupinambaranas (7upinambarama) Tupinambaranas (Mamuru), Rio Una, Rio Urubu Vacaria, Rio Vallao Grande (Valdo Grande) Velhas, Rio das Vereda do Mocambo (Mocambo) Victoria (Vitoria) Vigia Villa Bella (Parintins) Villa da Santa Rita (Santa Rita de Cassia) Villa do Barra (Barra) Villa (Vila) Velha Villa (Vila) Vigosa Vitoria (Victoria) Xingu, Rio Xique Xique (Chique Chique) Ypiranga (/piranga) NPORDWYWARROODRPWWIADWOOON- SAN >) wo > wD > 4, 67 ON on on 1977 THAYER EXPEDITION 7) REFERENCES AGASSIZ, LOUIS, AND ELIZABETH CARY. 1868. Journey in Brazil. Boston, Ticknor and Fields. 540 pp. ALLEN, Jort A. 1916. Autobiographical Notes and Bibliography of Scientific Publications. New York, American Museum of Natural History. pp. 11, 215. CamsripGeE. Harvard University Archives. William James papers. . Museum of Comparative Zoology. Scientific catalogs and specimens from various departments. . Museum of Comparative Zoology Archives. Louis Agassiz papers. . Museum of Comparative Zoology Archives. “List of Fishes” (by Orestes St. John). EIGENMANN, C: H. 1917. The American Characidae. Memoirs of the Museum of Comparative Zoology, 43. Hartt, C.F. 1870. Scientific Results of a Journey in Brazil. Boston, Fields, Os- good and Company. xxiii + 620 pp. INstITUTO BRASILEIRO DE GEOGRAFIA. 1972. Carta do Brasil Ao Milionesimo. Rio de Janeiro. Lurie, Epwarp. 1960. A Life in Science. Chicago, University of Chicago Press. pp. 345-350. THARP, Louise HALL. 1959. Adventurous Alliance. Boston, Little, Brown and Company. pp. 16-190, 204-210. Times AtLas. 1957. Volume 5, Cambridge, Houghton, Mifflin Co. Maps 115, Gs V7 Sapp s-xi1; 235,517 UniteD States BOARD ON GEOGRAPHIC NAMES. 1963. Gazetteer, Brazil. Wash- ington, D.C., U.S. Government Printing Office. 915 pp. VANZOLINI, P. E. AND N. PAPAVERO. 1968. Indice Dos Toponimos Contidos Na Carta Do Brazil. Sao Paulo, Fundac4éo de Ampara a Pequisa do Estado de Sao Paulo. 197 pp. Witcus, A. Curtis. 1972. Nineteenth Century Travelers: Louis and Elizabeth Agassiz. Americas 24 (2) : 25-32. No. 444 BREVIORA Olaues ap OLY ‘| aundI4 — SVLIFUA IC VQOVI (eH, VLISIA VWOd 0 ee _ WHIWO ? = OOOAVLO ave, IN OdVAOOYOO aS Cod VONrIL vuaoo Wd WHIT advddId WOVXNA SVC WHT ees : ee, AA by YOCYNYAAOD Od WISI ne VLaNOVd VHTI THAYER EXPEDITION 1977 (OLR “WeH Jaye) peosprey [[ O1paq Woq 7 aindi4 4 e a) N D OWIGNVS ad O14 WHOAN WLNOd SAG : é Sy y ‘“ u & “ Ox, LL & ne 7 NK %%, & 4 ae iS An 4 oS ay iv) IS & eS ms So YS One le °, oe es A Py Ss & & De ey 5 Ey SF a: aa fo PNY y e SW7O a 9g 2K 2 SI Tod0ubad | if / VHNVAVId OY OTTVOVINVO , ‘ No. 444 BREVIORA purjadod pure Wey — asay OO 0} BIYeY WO] ysBPOD S31" 0S 3493 0140, ———__ Sv11aAvaa | ‘| SOH10¥av VOSA qarey <2 2 " VWI 104031 VINWS SSO, S ‘> wn is) ONIND Sd AUN is = OdVud/> MED S ec 41 Ory & Ze Pg SWAON SYNINY > $09 YS Ody, oF S JLy 49 Of d ON 00 08vyg SSS O¥ND3S O14Od f SK OvHIWo * VaVeYvE VINVS VEVSINDIDNN {- 31LNOW139 6 OV nssv-0d YHNOHNILINO 3 SVUIBIAWNNW O° ove” BaUvd rr x» oauvd 0 > op ah SA3H & Ss S SS) » & = S m = (pny ~ > SVLNO9 30 o1y a) 01344 O18 VIHV Gr Vul30HOV9° ASSYNOVUWd OW AdAHNOVE OW 3d¥N9!I Olu ALIS THAYER EXPEDITION 1977 puvjado,y pure Wey — ollouer ap ory 0} seunye}] OlY WO] sBPOD “g ¢€ JNSIY S S31IN 0S ae? aed J nS a + = o1u4 OBv i oyovnuy aon soizna so Ofouranve 30 OW VNNW WINS 301 vow 72 3HVOVN 3SS0d & “— sodosauaHi> { = > nsswnal owé~ Cdvia4 voov 4 018 3WOHL OYS OBY S Uw 8 Vey 7 : Sodmv) 40% ak 30a ww : 0113dWv9 00 vooVT yw Go 1 110 WT 9% yyos 9 39V1'N3S VON3ZV4 VaOMVN OV? i> ons os VINOTAQ¥a Va Vuuas VNVNdVBV AL! Ol aan oN aanvan ovtwa VUISNDLLNVER VO VaRaS 4, 01404 4 vNaoveuve 29, WIY3W3dV LI sar woa wy 3LN3A3N38 O1Y %o VWNId O14 “4 WNId10d037 VINWS° 7 ZNYO WINVS Vy 30 BREVIORA No. 444 + PARANAGUA = ILLA DA SANTA RITA RIO PRETO BAHIA BAGRO DO RIO DAS VELHAS SETTE LASSAS * LAGOA SANCTA SANTA LUZIA ABURA MORRO VELHO SERRA DO ITATIAISSU SERRA DA PIEDADE CARANDAHY LAGOA DOURADA BARBACENA 50 MiLt. + PRADOS \ JUIZ DE FORA R Re oooh SERRA DE MANTIQUEIRA annev® RIO DE JANEIRO Figure 4. Rio de Janeiro to Paranagua — St. John THAYER EXPEDITION MARANHAO. i} 2, %, 4. % Lp “Y CAXIAS THEREZINA 2 < ° o we) 2 m s ABY ave Le he aT 7 \ RY S 0 saya lke S y = Ovd Ovs, ognanval OV % oS - Wan v108 00 OovINV 34431 ON OOW “hy Mindvown ' “Oo, 2 areas NWNY34 3SOP 'b, vn ? S $30 7 AMWONWAH OOV1 Vy. 09v7 2 2 KS : - 4 © = SS Ry vot 018 S\NSSW3d WAWL %, a] SS a j — % zt , a XsNIINvNOL Ue G > a vv, l4vdno 30 vaNas SN3OWIN0S O14 Vula¥dad 1977 THAYER EXPEDITION 37 S) § : z s g Qu Figure 11. Coast, Maranhao to Bahia — Agassiz et al. > - - 2 . * » , = _— x _ MUS, COMP. ZOOL: LIBRARY BRE V.EGR A Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, MASS. 30 NOVEMBER 1977 NUMBER 445 NATURAL HISTORY OF CERION. VII. GEOGRAPHIC VARIATION OF CERION (MOLLUSCA: PULMONATA) FROM THE EASTERN END OF ITS RANGE (HISPANIOLA TO THE VIRGIN ISLANDS): COHERENT PATTERNS AND TAXONOMIC SIMPLIFICATION. STEPHEN JAY GOULD! AND CHARLES PAULL? ABSTRACT. The eastern cerions (Hispaniola to the Virgin Islands) have, following the conventional practice for this diverse genus, been split into seven allopatric taxa. We reject this traditional scheme on the basis of a multivariate morphometric analy- sis (19 characters, 20 shells per sample) of 23 samples from all major areas of Cerion’s eastern range. We first show that eastern cerions are distinct from other members of the genus by patterns of covariation among their morphometric measures. We then demonstrate by canonical analysis that populations of each island have a dis- tinct morphology. Were there no coherent patterns among islands, current nomen- clature might be supported. However, the first canonical axis (59 per cent of all information) for an analysis by islands arrays the populations in perfect geographic order — from egg-shaped, finely and copiously ribbed shells in the east, to more cylindrical, apically pointed shells with fewer, stronger ribs in the west. The mor- phological direction of this cline approaches common cerion forms further west (particularly on Cuba); unique morphologies are isolated at the eastern extreme of the range. In addition, a plot of Mahalanobis vs. geographical distance shows a remarkably tight relationship (r = 0.96) between geographic and morphological distance. Therefore, we synonymize all living eastern cerions into the single taxon C. striatellum (“Férrusac” Guerin-Meneville), reserving C. rude (Pfeiffer) for the rather different St. Croix fossil. I. INTRODUCTION Cerion has attracted the attention of eminent naturalists ever since Linnaeus designated its type species in 1758. W. H. Dall (1905), H. A. Pilsbry (1902), P. Bartsch (1920), Ludwig Plate 'Museum of Comparative Zoology, Harvard University. 2Rosenstiel School of Marine Science. University of Miami. p) BREVIORA No. 445 (1906, 1907), W. J. Clench (1957), and Ernst Mayr (1956, 1963) invested substantial segments of their careers trying to understand the protean diversity of this unusual genus containing more than 600 described species and subspecies. So far Cerion has stood firm against all efforts to find coherence in the geographic distribution of its exuberant morphological variation. Elegant, oversimplified stories have collapsed (Plate, 1906), while more circumspect evo- lutionists speak of “crazy-quilt” distributions and random trans- port.by hurricanes (Mayr and Rosen, 1956). The burden of avail- able names has generally precluded a search for order and simplicity in the areas of Cerion’s greatest diversity — Cuba, the Bahamas, and the Cayman Islands. Yet order and simplicity demonstrably exist when we trade museum trays and lists of names for the map- ping of recognizable elements and their interactions in the field (Gould and Woodruff, in press, and in preparation on Little Ba- hama Bank, New Providence, Great Exuma, and Long Island, Bahamas). As another strategy, we might wage some preliminary skirmishes before attacking the central areas of Cuba and the Bahamas, since Cerion displays markedly less variation in three peripheral areas of its distribution. The isolated populations on Aruba, Bonaire, and Curagao have received much attention (Baker, 1924; Hum- melinck, 1940; de Vries, 1974; Gould, 1969). Geographic variation in the single species, Cerion uva (L.), reflects both isolation (each island has a characteristic, if subtly differentiated, morphology) and local habitat; patterns have been stable for at least 50 years (Baker, 1924; Gould, 1969). This coherence allows hope for sensi- ble interpretations elsewhere. The second peripheral area is con- tiguous with central regions: the Florida Keys with their single species, Cerion incanum (Binney). The third peripheral area is larger, more diverse, also contiguous to major centers, and equally unstudied: the essentially linear array of islands running from Hispaniola through Mona Island and Puerto Rico, to Necker and Anegada in the Virgin Islands (Fig. 1). Here 11 names are avail- able for a basic morphology that all students of Cerion have recog- nized as unique to this Eastern area (Pilsbry, 1902). These eastern cerions provide a superb opportunity for modern study; they repre- sent a situation intermediate between the relative simplicity of Cerion uva in the Leeward Islands (where we cannot practice the 1977 EASTERN CERIONS 3 art of taxonomic rectification, since no one ever successfully split the single species) to the complexity of New Providence Island with its 82 named species. - T 70°w 66°W S| TORTUGA 2 a 2 0UNE | RAND NES HISPANIOLA ANEGADA WIRSNDS \ PUERTO 3 © WN “weoxer oa Ke RICO | open }__*"__ gna -&% BEANE ST CROIX J Fig. 1. The distribution of eastern cerions. Crosses indicate localities used in this study for the large islands only. Moreover, the linear arrangement of islands leads to simple hy- potheses of clinal distribution in our search for coherence, while their geographic range (980 km from Western Haiti to Anegada) permits a truly regional, not merely local, study. i. SYSTEMATICS OF EASTERN CERIONS Collectors of eastern cerions report that these snails share the habitat conventionally assigned to the genus (though our own ob- servations of Bahamian forms demonstrate a much wider range); they live near the coast in limestone regions. Traditional wisdom proclaims that the animals are active on moist evenings, feeding predominantly on fungal mycelia in decaying vegetation; in drier conditions, they secrete an epiphragm and attach to plants, remain- ing immobile for long periods (several months in dry laboratory conditions). Beyond these basic facts of distribution, we know vir- tually nothing about the ecology and life history of any Cerion, including the eastern forms. (D. S. Woodruff has been studying two populations on Abaco Island, Bahamas during the past three and one-half years; our first adequate data will soon be available.) Heretofore, the eastern cerions have been defined by concho- logical characters alone. These snails have a unique and consistent morphology (Fig. 2), rendered in most taxonomic descriptions by two characters: 4 BREVIORA No. 445 1) An unusually obtuse apex, giving the entire shell a barrel or egg-shaped appearance (contrasting with the more common cylin- drical form of species with pointed apices and rapid attainment in ontogeny of a definitive width; or the triangular form of species with pointed tops and continually increasing width). 2) Fine, abundant and regular ribs, evenly covering the entire surface of the shell. Fig. 2. Representative eastern cerions. Top row Hispaniola; second row, Mona Island (left) to Anegada (right). The large specimen at the extreme right is fossil C. rude from St. Croix. Note the major clinal trends from coarsely ribbed, apically pointed species in the west (Hispaniola) to copiously and finely ribbed, barrel- shaped forms in the east. Note also, the peculiarities of individual islands as de- scribed in the text —i.e., the nearly circular aperture of Puerto Rican specimens. Top row, starting from left: 1) holotype of “C. rortuga” (Acad. Nat. Sci. Phil. No. 146701 — locality 22 of this study); 2) holotype of “C. vumaense ferruginum” S b yp (M.C.Z. No. 76230); 3) paratype of “C. vumaense™ (M.C.Z. No. 185945 — locality 18 of this study): 4) holotype of “C. vunaense saona” (Acad. Nat. Sci. Phil. No 47200). Bottom row: I) holotype of “C. monaense” from Mona Island (M.C.Z No. 171019 — locality 12 of this study): Puerto Rican specimen of C. srriatellum from locality 7; 3) from Necker Island, Virgin Islands, locality 1: 4) from Anegada. Virgin Islands, locality 2. C. rude is from lot No. 60564 Acad. Nat. Sci. Phil. © rude specimen is 34.7 mm in height 1977 EASTERN CERIONS 5 Pilsbry (1902), in the last comprehensive attempt to establish subgroups within Cerion, defined the eastern forms as one section among I1I in the subgenus Strophiops. He included two extrane- ous forms within the group: 1. Cerion antonii reported by Kiister (1847) from the South American coast in Guyana. This species has been dutifully listed by all students of Cerion; comments about its anomalous geogra- phy abound. But we do not believe that it exists. The shells were described from a collector’s cabinet, not from field observations. Mistaken labels in such cabinets once gave other cerions a proud range from India to China. We have no field records of living Cerion in South America. Moreover, Kiister’s shells have not been seen, so far as we can tell, by any subsequent author. All records of C. antonii depend upon the single poor figure published by Kiister and reproduced by Pilsbry (1902) in making his assignment. 2. C. caymanense Pilsbry from Grand Cayman. According to Pilsbry (1902), C. caymanense differs from other members of the eastern group by its less obtuse apex and stouter ribs. But these are the very characters that should place it out of the group; for obtuse apices and fine ribs are the defining traits of eastern cerions. We do not understand why Pilsbry didn’t include C. caymanense with the more similar Cuban group of C. cyclostomum (Kiister). The remaining taxa of Pilsbry’s eastern group all reside in the geographic area of this study. Current nomenclature divides the eastern cerions according to accepted criteria within the genus: specific names have been awarded to allopatric populations of recognizable morphology. Moving from east to west among living species, cerions of the Virgin Islands (Anegada and Necker) and Puerto Rico belong to the same taxon, Cerion striatellum — al- though the phenetic distance between Puerto Rican and Virgin Island samples is greater than that between most other species in the group (see section VI). Some confusion has surrounded the appropriate name for this taxon. Pupa striatella, attributed to Férussac but never described by him, was defined by Guerin- Meneville in the 1829 Regne Animal of Cuvier (Mollusca, p. 16, plate 6, fig. 12). But Guerin-Meneville listed only “the Antilles” for a locality, and the task of deciding which cerions merited the desig- nation fell to later authors. Kiister(1847) and many others attributed Guerin-Meneville’s name to the Puerto Rican specimens, but Poey 6 BREVIORA No. 445 and others applied C. srriate//um to a species from Cabo Cruz, Cuba and used Sowerby’s (1875) C. crassilabre for Puerto Rico. Pilsbry (1902) followed Poey, but changed his mind later (1943), awarded a new name to the Cuban species (C. cabocruzense Pilsbry and de la Torre), and preserved the earlier C. striatellum for the Puerto Rican form. We follow Pilsbry’s later opinion and consider C. crassilabre as a synonym of C. striatellum. This is a happy reso- lution since we will synonymize all living eastern cerions under this name, and since thick lips are decidedly not characteristic of many populations, whereas all are abundantly ribbed. Moving westward from Puerto Rico, we next encounter Cerion on Mona Island. Clench (1951, p. 274) awarded the Mona Island population their own name (C. monaense) for their coarser costae and smaller size compared with Puerto Rican relatives. Taxonomy of the Hispaniolan forms is more exuberant, but the defining criteria — size, ribbing, relative width, and shape of apex —have not been enlarged. Kiister’s name (1847), Pupa striatella minor from Haiti, is the earliest available, but all subsequent au- thors have ignored Kiister’s inadequate description, and we will not challenge this tradition. Pilsbry and Vanatta described the first full species in 1895 (p. 210): Cerion yumaense from the Dominican Republic — not so wide, more pointed apically, and more coarsely ribbed than the Puerto Rican populations. Maynard (1896) fol- lowed soon afterwards with Strophia ferruginea from Haiti, gen- erally ranked as a subspecies of C. yumaense by later authors (e.g., by Pilsbry, 1902). Two additional subspecies accrued in later years: C. yumaense sallei (Pilsbry and Vanatta, 1896, p. 325; first attrib- uted to C. striatellum, but later, and more appropriately, switched to C. yumaense by Pilsbry, 1902), and C. vuwmaense saona (Va- natta, 1923). Definitions involve little more than size: sa//ei is quite small, while saona is larger, with coarser costae than either sa/lei or the nominate subspecies. Finally, Pilsbry and Vanatta (1928) erected Cerion tortuga for more strongly colored, thicker lipped specimens from the nearby island of Tortuga. In summary, living eastern cerions are generally ranked in seven taxa: C. striatellum (“Férussac” Guerin-Meneville) for Puerto Rico and the Virgin Islands, C. monaense Clench for Mona Island (be- tween Puerto Rico and Hispaniola), and C. yumaense Pilsbry and Vanatta [with subspecies C. 1. rumaense, C. \. ferruginum (May- 1977 EASTERN CERIONS 7 nard), C. y. sallei Pilsbry and Vanatta, and C. y. saona Vanatta] and C. tortuga Pilsbry and Vanatta for Hispaniola and associated islands. They are distinguished by little more than variation in the key characters used to define the entire eastern group: coarseness of ribbing, obtusity of the apex, and width of the shell. A somewhat different fossil form has long been known from St. Croix — Cerion rude (Pfeiffer). [Pfeiffer’s (1855) second taxon, C. latilabre, has been properly synonymized by all later authors as nothing more than a short specimen of C. rude.] C. rude is consid- erably larger than other eastern forms. It also departs from its near- est neighbors in the very characters distinguishing them from other Cerion — C. rude has a more pointed apex and fewer, coarser ribs than most living eastern Cerion. Yet its general appearance is still closer to the living eastern cerions than to any other group within the genus. Jacobson (1968) states that the fossil shells are buried in soils and often appear at the surface after preparation of the land for cultivation of cane. He attributes their extinction to cul- tivation and burning (itself, primarily, for cultivation), but we have no firmly documented record of St. Croix Cerion living in historic times. III. MATERIALS, METHODS, AND INTENTIONS Following the protocol of Gould, Woodruff, and Martin (1974), we made 19 measurements on 20 specimens (when available) for 23 samples of eastern cerions. All taxa but C. yumaense sallei are represented in our samples. Our measures attempt to capture the standard taxonomic characters of adult size, whorl sizes and num- bers, size and shape of the aperture, shape of the spire, and pat- terns of ribbing. With the exception of one sample personally col- lected on Anegada (by C.P.), all shells come from museum collec- tions (see Table 1). In all but two cases (Maynard’s paratypes of C. yumaense ferruginum, and fossil C. rude — where we amalga- mated whatever we could find from the collections of four museums to form an adequate sample), we restricted ourselves to large, recent samples collected en masse without obvious preference for large, unusual or attractive specimens. We gathered these data for a mul- tivariate study of geographic variation in eastern cerions. We wished to address the following issues of distinction (at several levels) and coherence. 8 BREVIORA No. 445 Table |. Samples used in this study.* Necker Island, 220391 Anegada, 229017 Anegada, collected C. Paull Anegada, 203778 Near Tower Guanica Insular Forest, Puerto Rico, 212303 Tamarindo Beach, Puerto Rico, 212302 Guanica Insular Forest, Tower Road, Puerto Rico, 212293 Cabo Rojo Colony No. 2, Puerto Rico, 212298 Pt. Criollo, Guanica, Puerto Rico, 216702 10. Cayo Maguey, La Paruera, 212292 11. Cabo Rojo Colony No. 1, 212297 12. Mona Island paratypes, 171020 13. Mona Island, 190144 14. Mona Island, Isabella Anchorage paratypes, 184052 15. N.W. Saona Is., Hispaniola, 98829 16. Lighthouse Cabo Engafio, Altagracia, Hispaniola, 251314 17. North Shore Bahia de Yuma, Hispaniola, 251313 18. Yuma River, paratypes, Hispaniola, 181967 19. Romana Prov. 0.7 mi. east of Macao, Hispaniola, 250638 20. N.E. Grande Cayemites Is., Hispaniola, 251161 21. Juanilo, Altagracia, Hispaniola, 25132 22. Tortuga Island ANSP 146701 and 146702 23. St. Croix [composite sample: MCZ. Harvard: ANSP, AMNH (N_.Y.) and Field Museum (Chicago)]. SED SION A soe IS) *Samples 1-21 (with the exception of sample 3 collected by C. Paull) are from the Dept. of Molluscs, Museum of Comparative Zoology (lot number indicated). Sam- ple 22 is from the Academy of Natural Sciences, Philadelphia. Conventional iden- tifications follow: samples I-11, C. striatellum; 12-14, C. monaense; 15, C. vumaense saona; 16-19, and 21, C. vumaense; 20, C. ferruginum; 22, C. tortuga; 23, C. rude. 1. Distinction of eastern cerions from other sections of the genus: Are evident differences in morphology matched by pat- terns of covariation not encountered in other cerions? 2. Distinction among islands: Do the morphologies of eastern cerions record unambiguously the island of their occurrence? If so, we may identify isolation by geography as the primary corre- late of morphological variation in these animals. 3. Distinction among samples: A long tradition in Cerion studies (amply affirmed by all our qualitative observations) holds that nearly every local population has developed its own recognizable morphology. (The older conchologists chose to recognize this pri- 1977 EASTERN CERIONS 9 mary observation by granting each form its own specific name.) Does such local differentiation exist among the eastern cerions? 4. Coherence: If cerions are distinct by island (issue 2 above), can we detect any interisland pattern (clinal or otherwise) that might suggest a common control correlated with geography (selec- tion along climatic gradients, gene flow, or isolation by distance, for example)? The discovery of such a pattern will not permit us to distinguish among these potential explanations, for a pattern of coherence does not specify its cause, and museum specimens will not settle the issue. If we find no such pattern, current taxonomy granting autonomy to each major island must be maintained. If overall patterns exist, some consolidation may be in order, since hypotheses of direct interaction or similar responses of a single system to common factors would gain strong support from such patterns. The subsequent definition of our measures follows Gould et al. (1974, pp. 522-524) except for 6 and I1. Cerion is a biometrician’s delight for three reasons: 1) It preserves its entire ontogeny on fully exposed whorls. 2) It ceases growth with a definitive adult aperture. Thus we measure patterns of covariation among truly standardized adults. 3) We can specify comparable stages of ontogeny among speci- mens because the boundary between protoconch and accretionary shell provides an unambiguous, biological criterion for numbering whorls. Our characters can record both terminal sizes (for reason 2 above) and standardized traits of intermediate growth stages. Our measures include: 1. Width of protoconch. 2. Width at the end of the fourth whorl. 3. Total number of whorls of the adult shell (with the termina- tion of the protoconch taken as the Oth whorl). 4. Number of ribs on the fourth whorl. 5. Number of ribs on the sixth whorl. 6. Number of ribs in 40 micrometer units (2.22 mm) at the end of the first whorl. 7. Length of the adult shell, apex to lower apertural tip. 8. Maximum width of the adult shell. 9. Height of the protoconch. 0. Total height of the shell at the end of the fourth whorl. 10 BREVIORA No. 445 11. Height from the end of whorl 4 to the end of whorl 6. 12. Width of the umbilicus. 13. Width of the apertural lip at its widest point (measured parallel to the plane of the aperture). 14. Thickness of the apertural lip at its thickest point (measured perpendicular to the plane of the aperture). 15. Height of the aperture. 16. Width of the aperture. 17. Protrusion of the aperture beyond the junction of aperture and terminal whorl (see fig. 5 of Gould et al., 1974) —a measure of both adult size and intensity in change of adult coiling. 18. Tilt of the aperture —a “pure” (ratio) measure of change in coiling to produce the adult aperture. 19. Weight of the shell. IV. DISTINCTION OF EASTERN FORMS FROM OTHER CERIONS: PATTERNS OF COVARIATION The distinction of eastern cerions on traditional criteria of static adult morphology has long been recognized (Pilsbry, 1902). We have been experimenting with the promising, but rarely-used cri- teria of “dynamic morphology,” or differences in the structure of covariation among variables (Gould, 1969; Gould et al., 1974). We have, so far, studied in detail only the C. bendalli Pilsbry and Va- natta group of Abaco cerions (Gould et al., 1974). We wish to re- port some interesting differences between these and the eastern cerions. For seven samples of C. bendalli from Abaco we studied factor loadings on oblique axes (following varimax rotation) in separate R-mode analyses of each sample. We found that five axis solutions (recovering 70 to 75 per cent of the total information) yielded con- sistent and sensible interpretations with sufficient reduction of di- mensions to form interesting clusters. We grouped each variable with the axis of its highest loading. In all seven samples, we ob- tained the same five groups, each defined by the unvarying asso- ciation of two measures (see previous list of measures), and the more variable inclusion of others: 1. Adult size with whorl number (3) and shell height (7) as its focus. 1977 EASTERN CERIONS 11 2. Ribbing, with ribs on the fourth (4) and ribs on the sixth (5) whorl. This group is by far the most distinct among Abaco shells. It always occupies the second or third axis and never includes other measures (except, in two cases, the anomalous ratio measure of apertural tilt, where there can be no consistency, for measure 18 loads once positively, once negatively, and both times weakly). 3. Early whorl heights, with protoconch height (9) and height at the fourth whorl (10). 4. Later whorl sizes, with height of the middle whorls (11) and width at the fourth whorl (2). 5. Apertural lip, with lip width (13) and thickness (14). We performed similar R-mode oblique factor analyses for 21 eastern samples with sufficient specimens, using the DUVAP pro- gram (with provision for missing data) as in Gould et al., 1974. Five axes encompass 71 to 90 per cent of all information in these samples. No new groupings appear with any consistency or fre- quency. Associations three through five occur with less regularity but similar composition (in 14, 10, and 11 samples respectively). We note two outstanding differences between eastern and Abaco cerions: 1. In Abaco samples, whorl number and shell height are invari- ably associated as the focus of a group representing adult size. We find this association in only 9 samples of eastern cerions. In 16 samples (including some of these 9), whorl number appears in neg- ative association with one or more measures of shell size at stand- ardized early whorls (measures 1, 2, 9, 10, and 11). Such a negative association appears in no sample of Abaco C. bendalli. We assume that this disparity reflects a difference in the control exerted via growth over final adult size. A shell may reach large size either by growing more whorls or by growing larger whorls. If final size is strictly controlled, we anticipate a negative interaction of whorl number and whorl size — as in the eastern cerions (and in our one sample of a second Abaconian species, Gould et al., 1974, p. 528). In a jigsaw puzzle with rigid borders, increase in the size of one piece must be matched by decreasing size in others. But if size is not so strictly controlled, and if all factors that increase it work in concert (or at least independently), then we expect no negative interaction —as in the seven samples of Abaconian C. bendalli. If the borders of the jigsaw puzzle can expand, then all pieces might increase together. [2 BREVIORA No. 445 We do not know which of our characters best measures the elusive property of overall “size,” but let us consider shell weight as a possible surrogate: For all nine samples of Abaconian C. bendalli (only seven were factor analyzed), the mean coefficient of variation for shell weight is 21.00 (range from 15.51 to 25.85). Twenty-one eastern samples yield a mean C.V. of 16.82 (range from 9.9 to 30.2, with only three samples above 20.0). This differ- ence is significant at the 5 per cent level (t = 2.33 at 28 d.f.). Adult weight within samples is less variable for eastern than for Abaco cerions. Final size may be more strictly controlled in the eastern cerions. C.V.’s for weight exhibit an interesting heterogeneity among eastern cerions. Hispaniolan samples have a mean C.V. of only 12.41 (range from 9.9 to 14.3), while all other samples average 19.02 (range from 15.0 to 30.2, no overlap with Hispaniola). The lower value for Hispaniola is correlated with stronger negative in- teraction between whorl number and whorl size. In factor analyses of all seven Hispaniolan samples, for example, whorl number (3) always sorts in negative association with shell height from the fourth through sixth whorls (11). We find this negative interac- tion in only 5 of the remaining 14 groups. 2. In Abaco Cerion, the ribbing measures always form a tight cluster, completely distinct from all other variables. Their asso- ciation in eastern cerions is equally strong, but their group often includes other measures, almost invariably of adult size. In 18 of 21 samples, other measures group with rib numbers. Shell width (8) and umbilical width (12) join in four samples, width at the fourth whorl (2) and lip width (13) in three (but never lip thick- ness), apertural width (16) and apertural height (15) in two. This discovery supports our intuition that the “ribs” of eastern cerions —with their even spacing, complete coverage of the shell, and regularity of form — are not controlled in the same manner as the stronger, more irregular, and more widely spaced ribs of most other cerions. The more common pattern of other species may record environmental fluctuations and the pauses in growth that accom- modate to them. Perhaps the ribbing in eastern cerions reflects a more regular, internal metabolic pattern correlated with general growth (and final size) of the animal. 1977 EASTERN CERIONS 13 We gain some support for this speculation from coefficients of variation. In Abaco C. bendalli, mean C.V. for ribs on the fourth whorl is 14.75 (range from 12.27 to 20.94), and for ribs on the sixth whorl, 14.83 (range from 11.93 to 17.00). Comparable figures for the same measures in eastern cerions are 8.57 (range from 3.67 to 13.59) and 8.68 (range from 3.42 to 11.27). This distinction cannot be explained by differences in absolute numbers of ribs, for the ranges are comparable in both groups (mean rib number varies from 33 to 80 in Abaco samples and from 25 to 78 in eastern sam- ples). V. DISTINCTIONS AMONG ISLANDS FOR EASTERN CERIONS The traditional taxonomy of eastern cerions records primarily a distinction among islands (though we note with some surprise that no one ever split the Virgin Island from the Puerto Rican cerions). We have already reported an interesting difference in variability of shell weight between Hispaniola and other areas. Differences in variation and covariation should be pursued more vigorously (if only because they are so widely neglected); but for now we proceed along traditional lines and examine differences in form. We performed a canonical analysis on all 23 samples (373 speci- mens) using D/DA, a program written by John Rhoads, Depart- ment of Anthropology, Yale University (see Gould et al., 1974). We treated each sample separately, without any prior identification of its island. Table 2 records the order of discriminating power for variables expressed in univariate ANOVA. We find a perfect cor- respondence between this order and the traditional criteria of tax- onomic distinction: strength of ribbing (variables 4, 5), width of the shell (8, 13, and 16), and obtusity of the apex (10). (Shells bearing obtuse apices are high at the end of the fourth whorl; whorls enlarge quickly and begin their rapid translation down the axis of coiling early in ontogeny. Triangular apices are lower at the end of the fourth whorl; the “triangle” records a slower increase in whorl size and a later incidence of the strong allometry that gives Cerion its characteristic shape by increasing height relative to width during post-juvenile to pre-adult growth.) The matrix of means for islands also records the rough order of discrimination. (Table 3 — 14 BREVIORA No. 445 Table 2. Univariate ANOVA: discriminatory power of individual variables for all 23 samples F-ratios with 22 and 350 degrees of freedom. All are significant. Name Number F-ratio 4th ribs 4 208.5 6th ribs 5 157.0 aperture width 16 102.9 width 8 102.9 4th height 10 101.8 lip width 13 67.2 aperture height 15 54.1 height i 53.0 Ist ribs 6 52.4 4-6th height 1] 52.1 umbilicus 12 45.2 4th width 2 40.4 whorls 3 33.6 lip thickness 14 22 protrusion 17 22 protoconch width l 19.5 protoconch height 9 18.8 tilt 18 33 not the mean of sample means, but the mean of ungrouped shells for each island.) The first three canonical axes account for 87 per cent of the vari- ance among sample means (45.1, 29.6 and 12.3 per cent respec- tively; the fourth axis falls below 5 per cent). Figure 3 displays the projection of sample centroids upon the first two axes. Sam- ples clearly group by islands; all islands are widely separated with the exception of Hispaniola and Mona. The introduction of axis three completes the pattern; Mona and Hispaniola are now distinct (Fig. 4). We need only three components of variation to sort 23 samples unambiguously into five larger groups. The strength of determination by island is well recorded in the classification matrix: 309 of 373 specimens are correctly classified, and every misclassi- fied specimen sorts with another sample of its island. If we redo the analysis with islands rather than samples, as groups, not a single specimen is misclassified. Each lies closest to the centroid of its own island. Geographic isolation seems to be the primary correlate of morphological variation in eastern cerions. 1977 Table 3. N in mm. Number of specimens . protoconch width 4th whorl width total whorls 4th ribs 6th ribs Ist ribs height width protoconch height 4th height . 4th-6th height umbilical width lip width lip thickness aperture height aperture width . protrusion tilt weight EASTERN CERIONS Necker 7.63 2.10 1.20 Anegada 59 Puerto Rico 140 Mona 40 His- paniola 110 15 Matrix of mean values for islands. Weight in grams. Linear measures St. Croix The matrix of factor pattern for the analysis by samples (Ta- ble 4) defines the uniqueness of each island and exhibits the mor- phological bases of separations in Figures 3 and 4. (The factor pattern matrix of standardized partial regression coefficients per- mits us to discern how the original measures vary together in dis- tinguishing samples along the canonical axes.) We did not include shell weight (variable 19) in this analysis because we could not weigh the St. Croix fossils with their permineralized shells and filled interiors. 16 BREVIORA No. 445 6reP {P| e S V4 P Sat | . cP *p | a \ sts 2 = Plaly ———— all = iL = ! 1 ! Sl Ne Aes -6 -4 \ =e O 2 4 6 8 10! l2 \ AXIS | re M “ / es wm-2b La H, N x / \ Mee He A l Heats \ ae a NH -6t Fig. 3. Plots of centroids for all samples on first two axes of a canonical analy- sis (74.7 per cent of all information) to show distinction of islands and coherence within an island. Islands are circumscribed by minimum convex polygons. A- Anegada, N-Necker, P-Puerto Rico, M~Mona, S-St. Croix, and H-Hispaniola. H,-Cerion tortuga (locality 23), H;i-C. yumaense ferruginum (locality 20), HC. yumaense saona (locality 15). ae H aK eet He P p é con eePye Hs P/ eP © ep °5 - =I [a -6 =O pulse O 2 4 6 8 He) l2 ie AXIS | Ep ine) hs | Le AY Hy x< | dq Mee Ss ~e N 4 Me e -6 Fig. 4. Axis I vs. axis 3 for the same canonical analysis depicted in Figure 3. Code letters same as in Figure 3. Note that Mona and Hispaniola (overlapping in Fig. 3) are distinguished by the third axis. 1977 EASTERN CERIONS 17 Table 4. Factor pattern (structure of covariance) for original variables on first five canonical axes for all 23 samples. No. Name l 2 3 4 5 protoconch width ~0.0430 0.1082 —0.3254 0.2116 -0.0632 l 2 4th width -0.0239 0.2443 -0.4745 0.1357 -0.1085 3 whorls -0.0047 0.2851 -0.1552 -0.4931 -0.1116 4 4th ribs 0.6196 0.4860 0.2486 -0.0291 -0.1228 5 6th ribs ~ 0.4526 0.5397 0.2748 -0.0553 -0.2221 6 Ist ribs 0.2427 0.3124 0.0459 0.2449 0.4016 7 height -0.0968 0.3027 ~0.4599 -0.1611 -0.4270 8 width -0.0607 0.5167 -0.5713 —0.1082 -0.2853 9 protoconch height -0.1590 -0.1151 -0.1863 —0.0339 -0.0540 10 4th height 0.3893 —0.2690 -0.4891 0.1371 0.1004 11 4 -6th height 0.1832 -0.2792 -0.2773 0.4856 —0.2967 12 umbilicus ~0.0624 0.2278 -0.4813 -0.3020 0.2144 13. lip width —0.2000 0.4035 -0.2115 0.3895 0.1607 14 lip thickness ~0.0726 0.2054 -0.1453 0.4387 -0.1463 15 aperture height -0.0788 0.3720 ~0.3568 -0.0482 ~0.4348 16 aperture width -0.1126 0.5692 —0.3769 —0.0336 -0.1512 17 protrusion —0.0602 0.2759 -0.0882 -0.0274 -0.0037 18 tilt ~0.0180 0.0311 0.0417 -0.1369 -0.1053 The first axis separates Virgin Island samples from all others (see Figs. 3, 4). Its factor pattern records high values for measures of ribbing and whorl heights (10 and 11). Indeed, the Virgin Island samples exceed all others in their number of ribs and the obtuseness of their apices (Fig. 2). The negative loading of protoconch height (9) also reflects the obtuse apex of Virgin Island cerions (see mean values in Table 3). High protoconchs are followed by slowly ex- panding whorls that join the embryonic shell to produce a pointed apex with modest whorl heights; low protoconchs precede rapid expansion (high values of 10-11) and build more obtuse apices. The second axis separates Puerto Rican samples (positive pro- jections) from Mona Island and Hispaniola (negative projections). The factor pattern contains high positive loadings for two groups of variables: measures of ribbing (4 and 5) and shell widths, par- ticularly aperture width (8 and 16). Whorl heights (10 and 11) load negatively. The cerions of Puerto Rico can be distinguished visually by their more circular apertures; this feature translates to high values for aperture width in our quantitative analysis. The St. Croix specimens (which also project strongly and positively on 1s BREVIORA No. 445 axis two) are wide as a simple function of their large size. The ribbing measures set the major contrast between Puerto Rican samples and Mona and Hispaniolan samples. Puerto Rican cerions are abundantly ribbed (though not so abundantly as Virgin Island samples), while Mona and Hispaniolan cerions have the fewest (though strongest) ribs among eastern cerions. Puerto Rican spe- cimens also have fairly triangular apices. This feature translates as low whorl height (10 and I1) in our measures; the negative load- ing of measures 10 and II assures the high positive projection of Puerto Rican samples on the second axis. The third axis separates Mona and Hispaniola to complete the distinction among islands. Mona and Hispaniolan samples fell together with negative projections on the first two axes for essen- tially the same reason — their small number of ribs. Several meas- ures of size — particularly of widths (2, 8, 12, and 16 —load strongly and negatively upon the third axis. Hispaniolan shells are small and especially narrow (note low mean values of 2 and 8 in Table 3); they project positively upon an axis with negative loadings for these measures. Mona samples are large and wide; since measures for size and width have negative loadings, these samples project negatively upon the axis. Subsequent axes continue the patterns of distinction by island. The fourth axis, for example, separates Mona, having high pro- jections, from St. Croix, having low projections. Highest loadings are for height from whorl 4 through 6 (positive) and number of whorls (negative). St. Croix specimens with their maximal num- ber of whorls and minimal fourth to sixth whorl heights project negatively. Mona samples with their distinctively low number of whorls project positively. The fifth axis separates Virgin Island samples into their two areas of Necker and Anegada. The Necker Island sample projects strongly and negatively upon an axis with positive loadings for its low number of ribs on the first whorl (rela- tive to Anegada), and high negative loadings for various measures of shell size. VI. DISTINCTION AMONG SAMPLES A persistent tradition of Cerion studies holds that virtually every local population bears its own distinctive morphology. This re- markable phenomenon has inspired the burden of an exuberant 1977 FASTERN CERIONS 19 specific nomenclature for these animals. Nothing we have seen in several years of study contradicts this conventional observation. We therefore predicted that samples within islands should be dis- tinct in separate canonical analyses for each island. Our results confirm this prediction with great accuracy. The classification ma- trix for four Virgin Island samples correctly assigns 63 of 65 spe- cimens to their correct sample (96.9 per cent). Only 3 of 40 speci- mens from Mona Island lie closest to the centroid of a sample not their own (92.5 per cent are correctly classified). Of specimens from Hispaniola, 101 of 110 are correctly allocated among eight samples (91.8 per cent). Only Puerto Rico falls below 90 per cent success — 116 of 139 specimens in seven samples (83.5 per cent). We have, in summary, established a hierarchy of distinction among the eastern cerions. Their basic design is strikingly different —both in form and in pattern of covariance — from other ceri- ons. Islands are distinct. Samples within islands can also be sep- arated with little overlap. VII. AUNIFYING THEME ANDA TAXONOMIC PROPOSAL The distinction of islands does not preclude the existence of pat- tern among them. We concluded that geographic separation serves as the primary correlate of morphological distinction among the eastern cerions. But does the degree of morphological separation reflect geographic distance? If so, we might expect a clinal distri- bution of morphology, since the islands form an essentially linear array. If we find no coherent pattern, taxonomic distinction by island should be maintained (with, God forbid, a new taxon for the Virgin Islands). The output of our canonical analysis for islands includes a ma- trix of Mahalanobis distances (D2) — a measure of separation be- tween centroids with adjustments for variance and covariance (Ta- ble 5). Each entry is a measure of morphological distinctness (based on all variables) between shells of two islands. If we plot D2 (see Sneath and Sokal, 1973, pp. 127, 128) against geographic separation (defined, for any island pair, as the distance between our two closest sample localities), we obtain a remarkably tight and linear relationship (r = .96). This calculation (Fig. 5) excludes the St. Croix fossils as representatives of another time with no expected relationship to current trends. When we plot the St. Croix 20 BREVIORA No. 445 Table 5. Matrix of Mahalanobis distances (D2) for analysis using all specimens grouped by islands. N A P M H S Necker 0 Anegada 18.87 0 Puerto Rico 125333 102.90 0 Mona 155.33 134.73 54.66 0 Hispaniola 164.32 131.32 38.00 S23 0 St. Croix 214.23 187.83 65.86 65.29 57.11 0 200 150 MAHALANOBIS D2 oO (e) On (e) | | O 100 200 300 400 500 DISTANCE IN KM Fig. 5. Mahalanobis distance (morphological) vs. geographic separation for eastern cerions. Note the tight correlation and the departure of St. Croix fossils from the trend. A~Anegada, N-Necker, P-Puerto Rico, M~Mona, H~-Hispaniola, and S-St. Croix. 1977 EASTERN CERIONS Di S HM p AN aes) Vay -4 =2 0 2 4 6 8 Te Fig. 6. Projections on the first canonical axis for a discriminant analysis by islands. This axis explains more than half (58.8 per cent) of all information. Note the perfect correspondence of morphological gradation and geographic position for the modern samples. Only the St. Croix fossils depart from the trend. Code letters as in Figure 5. - distances on Figure 5, they depart markedly from the regression line of modern samples. Geographic and morphological distance are closely correlated in living eastern cerions. But this relation- ship does not guarantee any graded trend of morphology along the linear array of islands. In fact, we already have a test for clines in the results of our canonical analysis. In our canonical analysis for islands, the most significant dis- criminator (the first axis) explains 58.8 per cent of all information. Figure 6 shows the array of projections. Living cerions are arranged in perfect geographic order; only the fossils of St. Croix depart from the pattern. The morphological gradation runs from egg- shaped, finely and copiously ribbed shells with very obtuse apices (Virgin Islands) to more cylindrical, apically pointed shells with fewer, stronger ribs (Hispaniola) —see the discussion of factor patterns in Section V. The direction of the cline also meets our expectations. Virgin Island shells represent the extreme expression of morphological features that characterize eastern cerions; they are also geographically furthest from any contact with non-eastern congeners. The Hispaniolan specimens, situated in closest prox- imity to non-eastern cerions, depart most strongly from modal east- ern morphology towards the more “normal” form of the genus. We are not addicted to speculative biogeography, and we offer no preferred hypothesis for this pattern; we do not see how mu- seum specimens can distinguish among such alternatives as selective gradients and gene flow, and we know virtually nothing about the history of colonization. We only note that the strikingly discordant position of the St. Croix fossils may indicate a fairly recent origin for the graded series of modern forms. (The St. Croix fossils are morphologically furthest from their nearest modern relatives on Necker and Anegada.) 22 BREVIORA No. 445 In the light of this pattern, we see no reason to maintain the taxonomic distinction among living eastern forms. Some common factor accounts for most of the morphological variabiuty among these populations. We therefore consider all of the following names as synonyms of the first-named form, Cerion striatellum (“Feérru- sac” Guerin-Meneville), 1829: C. monaense Clench, 1951; C. yu- maense Pilsbry and Vanatta, 1895 (with its four subspecies); C. ferruginum (Maynard, 1896 —ranked as a subspecies of C. yu- maense by most later authors); and C. tortuga Pilsbry and Vanatta, 1928. We maintain, for now, Cerion rude (Pfeiffer), 1855 for the St. Croix fossils. The St. Croix shells depart completely from modern trends in geographic variation. Indeed, this departure serves aS a strong support for taxonomic identity of recent popu- lations — for it argues against any appreciable antiquity for the modern pattern. Moreover, the St. Croix shells, while closer to the eastern mode than to any other morphotype of Cerion, vary from this mode in the direction of more conventional Bahamian and Cuban forms — shells are larger, more cylindrical, and more sparsely and strongly ribbed. As another argument for the tax- onomic unity of modern forms, we note that the morphological distances among islands are trifling compared with the distinction of several forms that interbreed freely throughout the Bahamas. In fact, and incredible as it may seem in the light of Cerion’s fan- tastic morphological variety, we have located no unambiguous case of sympatry without interbreeding for the entire genus (two of nine reported cases have a fair chance of validation in our opinion). VIII. CONCLUSION Studies of Cerion have been dominated by the conviction that no regular patterns of geographic variation exist. All leading stu- dents have invoked haphazard transport by hurricanes to support a notion of “crazy-quilt” distributions (Mayr and Rosen, 1956). We now present the first evidence for simple, graded, coherent variation over a large part of Cerion’s range. Moreover, S. J. Gould and D. S. Woodruff, working in the northern Bahamas during the past four years, have found strikingly consistent pat- terns of within-island variation on all islands visited (Abaco, Grand Bahama, New Providence, Great Exuma, and Long). The “ribby” morphotype, for example (called C. glans (Kiister) and 1977 EASTERN CERIONS NO eS) about 50 synonyms on New Providence, C. abacoense Pilsbry and Vanatta and 3 synonyms on Abaco and Grand Bahama, etc.), al- ways inhabits coastal areas at the edges of the Pleistocene banks; while the mottled morphotype inhabits interior coasts (with respect to the banks) and interior areas. The two morphotypes intergrade in a variety of interesting ways at their contacts. The discovery of simple and coherent patterns, both within and between islands, leads to a hope — even an expectation — that some sense will yet be made of Cerion. Ix. ACKNOWLEDGMENT This work was supported by NSF Grant BMS 72-02213. REFERENCES BAKER, H. B., 1924. Land and freshwater molluscs of the Dutch Leeward Islands. Occ. Pap. Mus. Zool. Univ. Mich. No. 152, 158 pp. BARTSCH, P., 1920. Experiments in the breeding of Cerions. Papers Dept. Marine Biol. Carnegie Inst. Wash. No. 282, 55 pp. CLENCH, W. J., 1951. Land shells of Mona Island, Puerto Rico. J. Conchyliologie 90: 269-276. , 1957. A catalogue of the Cerionidae (Mollusca — Pulmonata). Bull. Mus. Comp. Zool. 116: 121-169. Dat, W. H., 1905. Fossils of the Bahama Islands, with a list of the nonmarine mollusks. in G. B. Shuttuck (ed.), The Bahama Islands. N. Y., MacMillan, pp. 23-47. DE Vries, W., 1974. Caribbean land molluscs: notes on Cerionidae. Stud. Fauna Curagao and other Carib. Islands 45: 81-117. GouLb, S. J., 1969. Character variation in two land snails from the Dutch Lee- ward Islands: geography, environment, and evolution. Syst. Zool. 18: 185-200. GouLtp, S. J.. D. S. WoopRuUFF, AND J. P. MARTIN, 1974. Genetics and morpho- metrics of Cerion at Pongo Carpet: a new systematic approach to this enigmatic land snail. Syst. Zool. 23: 518-535. GuféRIN-MENEVILLE, F. E., 1829. Iconographie du regne animal de G. Cuvier, Mollusques. Paris, Fortin. HuMMELINCK, P. W., 1940. Mollusks of the genera Cerion and Tudora. Stud. Fauna Curagao, Aruba, Bonaire, and Venezuelan Islands 2: 43-82. JACOBSON, M. K., 1968. The land Mollusca of St. Croix, Virgin Islands. Sterkiana 32: 18-28. Kuster, H. D., 1847. Die Gattungen Pupa, Megaspira, Balea und Tornatellina. Conchylien-Cabinet (2), 1, pt. 15, 194 pp. MAYNARD, C. J.. 1896. Monograph of the genus Srrophia. Contributions to Science, vol. 3, Newtonville, Mass. 24 BREVIORA No. 445 Mayr, E., 1963. Animal species and evolution. Cambridge, Mass., Harvard Univ. Press, 797 pp. Mayr, E., AND C. B. ROSEN, 1956. Geographic variation and hybridization in populations of Bahama snails (Cerion). Novitates Am. Mus. Nat. Hist. No. 1806, 48 pp. PFEIFFER, L., 1855. Beitrage zur Molluskenfauna Westindiens. Malak. Blat. 2: 98-106. Pitssry, H. A., 1902. Family Cerionidae. Manual of Conchology, Second Series, Pulmonata, Vol. 14. Philadelphia, Acad. Nat. Sci., pp. 174-281. , 1943. Note on Cerion striatellum. Nautilus 57: 34-35. Pitssry, H. A. AND E. G. VANATTA, 1895. New species of the genus Cerion. Proc. Acad. Nat. Sci. Phila. 47: 206-210. , 1896. Catalogue of the species of Cerion, with descriptions of new forms. Proc. Acad. Nat. Sci. Phila. 48: 315-338. , 1928. Land shells of Tortuga Island, Haiti, and a new Haitian Oleacina. Proc. Acad, Nat. Sci. Phila. 80: 475-478. PLATE, L., 1906. Die Artbildung bei den Cerion-Landschnecken der Bahamas. Verhandl. deutsch. zool. Ges. 16: 127-136. . 1907. Die Variabilitat und die Artbildung nach dem Prinzip geographi- scher Formenketten bei den Cerion-Landschnecken der Bahama-Inseln. Arch. Rassen Gesellsch. Biol. 4: 433-470, 581-614. SNEATH, P. H. A., AND R. R. SOKAL, 1973. Numerical Taxonomy. San Francisco, W. H. Freeman, 573 pp. SowerBy, G. B., 1875. Monograph of the genus Pupa. Conchologia Iconica, vol. 20. VANATTA, E. G., 1923. Two new Santo Domingan land shells. Proc. Acad. Nat. Sci. Phila. 75: 359-360. “US. COMP. ZOOL LIRRARY BREVIORA UNIVER ERSITY Museum of Comparative Loology US ISSN 0006-9698 CAMBRIDGE, MASS. 20 DECEMBER 1978 NUMBER 446 pein ene aac tn ene es AS nee ee i Be ee ee ee A NEW INTERPRETATION OF THE MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN FROM THE ALBIAN OF TEXAS PERCY M. BUTLER! ABSTRACT. A reappraisal has been made of tribosphenic teeth from the Albian of Texas, previously described by Patterson, Slaughter and Turnbull. At least six genera are represented. Pappotherium and Holoclemensia, the most common gen- era; are sufficiently alike to be placed in the same family, Pappotheriidae. They have four molars as in marsupials and complex premolars as in placentals. They cannot be classified either as marsupials or placentals but are considered to belong to a separate evolutionary line (Pappotherida, new order). Two forms of smaller size, Kermackia texana and Trinititherium slaughteri gen. et sp. nov., whose lower molars are at a more primitive stage of evolution than the Pappotheriidae, are placed in a new family Kermackiidae (Aegialodontia, new order). Slaughteria erup- tens gen. et sp. nov., the only species for which there is evidence of tooth-replace- ment, is placed provisionally in the same family. A new infraclass, Tribotheria, is proposed to include mammals with tribosphenic molars that are not classifiable as marsupials or placentals. INTRODUCTION Patterson (1956) reported on a collection of teeth found in the Greenwood Canyon, near Forestburg in eastern Texas. Many of the molar teeth are of tribosphenic pattern. They are mid-Creta- ceous (Albian), the oldest tribosphenic molars known at that time. The Greenwood Canyon fauna is intermediate in age between the late Jurassic Purbeck and Morrison faunas and the late Cretaceous Djadokhta fauna of Mongolia. During this long interval of about 50 million years, the Jurassic mammal orders were largely replaced 'Zoology Department, Royal Holloway College, Englefield Green, Surrey TW20 9TY England. D BREVIORA No. 446 by marsupials and placentals. Only the multituberculates survived into the late Cretaceous in more than smal numbers. The differentiation between marsupials and placentals is believed to have taken place during the earlier part of the Cretaceous. Both marsupials and placentals share the tribosphenic molar pattern. With only the evidence of isolated teeth, Patterson was unable to decide in which infraclass his specimens belonged or whether they represented a group from which both marsupials and placentals would arise. Accordingly, he classified them as “Therian mammals of uncertain infraclass affinities but of Metatherian-Eutherian grade.” Between 1965 and 1971, Slaughter published descriptions of im- portant additional material. He obtained it from Butler Farm, a locality of similar age to the Greenwood Canyon locality. He named two species based on upper molars: Pappotherium patter- soni and Holoclemensia texana. The holotype of P. pattersoni is a fragment containing the last two molars. The holotype of H. texana is a penultimate molar of which the protocone shelf is lack- ing; Slaughter used a complete last molar as a paratype and re- ferred a lower molar to this species. He named a third species, Kermackia texana from another isolated lower molar. Holoclemensia differs from Pappotherium in the arrangement of the buccal stylar cusps of its upper molars (Fig. 1). In Pappother- ium there is a large stylocone (Style B), about as high as the meta- cone. In Holoclemensia, the stylocone is much smaller, but there is a large conical cusp (Style C) opposite the notch between the paracone and the metacone. In the incomplete material at his dis- posal Patterson (1956) mistook Style C for the stylocone. Style C occurs in many late Cretaceous marsupials; it is higher than the stylocone in Albertatherium and A quiladelphis (Fox, 1971). Slaughter (1968b) regarded Holoclemensia as a marsupial, not only because of the presence of Style C, but also because the meta- cone is less reduced than in Pappotherium. In 1968a he described some partly molariform premolars, unlike those of any marsupial. In 1971, he referred to Pappotherium a mandible that contained a newly erupted and a partially erupted premolar together with worn molars. This specimen indicates diphyodonty of the anterior den- tition, characteristic of placentals. He therefore regarded Pappo- therium as a placental. Subsequently Fox (1975) deduced from the 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 3 A Figure |. P, last two upper molars of Pappotherium pattersoni, SMP-SMU 61725, and buccal view of penultimate molar. H, the same, Holoclemensia texana, SMP-SMU 62099 and 62147. A, buccal view of upper molar of Alphadon marshi. Stylar cusp C is indicated by an arrow. shapes of the last two teeth that Pappotherium had only three molars. Lillegraven (1969, 1974) and Hoffstetter (1970, 1972) believe that the marsupial-placental differentiation was due to isolation on dif- ferent continents. Placentals in Asia and marsupials in North America suggest that the placentals that appear in North America towards the end of the Cretaceous are immigrants from Asia. This opinion does not accord with Slaughter’s view that both infraclasses were present on the continent as early as the Albian. It seemed desirable to reexamine the evidence provided by the Texas Albian fauna that affects a crucial stage of mammalian evolution. 4 BREVIORA No. 446 During visits to the United States in 1966 and 1974 I studied specimens from Greenwood Canyon that Patterson described (1956) as well as additional material that Turnbull (1971) described. The latter was subsequently collected from the same locality. The Greenwood Canyon material is housed in the Field Museum of Natural History, Chicago (registration numbers PM —). Of the Butler Farm collection, I saw the specimens described by Slaughter (1965, 1968a) and the jaw containing premolars with molars (Slaughter, 1971). I studied other Butler Farm specimens from casts. The material from Butler Farm is housed in the Shuler Museum of Paleontology, Southern Methodist University, Dallas, Texas (registration numbers SMP-SMU — ). I used a Wild M5 microscope to make camera lucida drawings, at known magnifica- tions, of all the specimens studied. Nearly all the specimens consist of isolated teeth. A primary aim of the investigation was to find the most probable way in which they could be combined into dentitions. The relationships of the species believed to be present in the fauna could then be considered. UPPER MOLARS All the upper molars that I have seen are drawn in crown view in Fig. 2. Except in the holotype of Pappotherium, all are isolated, and their position in the series must be judged from their shape alone. Last molars, whether M3 or M4, are easily recognizable. Reduction of the metacone and loss of the metastyle accompany a reduction of the posterobuccal lobe of the outline. These features are widespread and almost certainly primitive among mammals with tribosphenic molars. On passing forward along the molar series, both in marsupials and placentals, the posterobuccal lobe becomes relatively larger, so that eventually it projects buccally beyond the level of the parastyle; the metacone also enlarges and the metastylar crest becomes longer. The degree of difference be- tween adjacent molars varies considerably between genera and families and to some extent individually. The penultimate molar is preserved in the holotypes of Pappotherium and Holoclemensia (Fig. 2, d, 1). In both, the posterobuccal lobe is less prominent than the parastylar region, though both possess the metastylar crest. 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 5 When measured by the relative distances of the metacone and par- acone from the buccal edge, Holoclemensia has a rather less prom- inent posterobuccal lobe, but the difference between the two genera is not great. Holoclemensia also has a higher metacone than Pap- potherium, a difference that is repeated on the last molar. Of the isolated last molars, PM 1075 (Fig. 2, c), though smaller than the holotype of Pappotherium, agrees with that specimen in structure; PM 1015 (Fig. 2, b) has a less reduced posterobuccal lobe and a larger metacone; PM 1287 (Fig. 2, p) has a smaller pro- tocone shelf than Pappotherium. The posterior surface of the stylo- cone of PM 1015 has a small cusp, indistinctly indicated on PM 1075. Whether this cusp is the equivalent of Style C is uncertain. Style C is comparatively well developed on the last molar of Holo- clemensia, but Turnbull (1971, Fig. 1) misidentified it as the meta- style. PM 1015 and PM 1075 both differ from Holoclemensia and resemble Pappotherium in having a concavity of the anterior out- line lingually to the parastyle. PM 1287, with its small protocone, is clearly distinct from both Holoclemensia and Pappotherium. Several upper molars have the posterobuccal region less reduced than on the penultimate molars of Holoclemensia and Pappothe- rium, and they must be considered as examples of more anterior molars. PM 1000 (Fig. 2, 0) has a well preserved buccal stylar shelf on which Style C is the highest cusp. This identifies the tooth as Holoclemensia, but the metacone is farther from the buccal edge than the paracone is. PM 886 (Fig. 2, m) has lost the parastylar region, but Style C, which is preserved, is as high as the metacone. This tooth is larger than PM 1000 and differs from it in that the paracone and metacone are nearly equidistant from the buccal edge. PM 1004 (Fig. 2, n), poorly preserved, agrees with PM 886 in comparable parts. There is evidence that Holoclemensia had two molars anterior to the penultimate tooth and four molars with all included (Fig. 5, H). PM 999 (Fig. 2, h) resembles the penultimate molar of Pap- potherium in the enlarged stylocone and in the metacone, which is less high than the paracone; however, the posterobuccal lobe of PM 999 is prominent and bears a strong metastylar crest. PM 1238 (Fig. 2, i), of which the anterior part of the stylar shelf is missing, agrees with PM 999 in the metacone and metastyle and is almost BREVIORA No. 446 W= ‘ e Lots’ Wee oe. 7 ‘yy Vd! 2 WK ] NS WAAAY g VU _/\\ J ‘ 0 ‘ \ NNN py Natt \ \ \ , \\\" 1mm fees cere] Pp Figure 2. Crown views of upper molars. a, SMP-SMU 61725 (holotype of Pappotherium pattersoni); b, PM 1015; c, PM 1075; d, SMP-SMU 61725 (holotype of P. pattersoni); e, PM 884; f, PM 1749; g, PM 1325¢ h, PM 999: i, PM 1238: J, SMP-SMU 62402; k, SMP-SMU 62099 (paratype of Holoclemensia texana); |, SMP-SMU 61947 (holotype of H. texana); m, PM 886; n, PM 1004; 0, PM 1000; p, PM 1287. All drawn as right teeth: b, c, e, f, i, 0, p have been reversed. 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN V certainly another example of the same tooth. PM 884 and PM 1749 (Fig. 2, e, f) represent a different type of molar. The meta- stylar region has broken off in both specimens. As in Pappothe- rium, the stylocone is high and the metacone is lower than the para- cone. These teeth are larger than PM 999 and PM 1238 (Fig. 2, h, i), and based on the incomplete crowns, the metacone and para- cone were nearly equidistant from the buccal margin. On PM 999, PM 884 and PM 1749 there is a slight elevation of the marginal ridge in the region corresponding to Style C, but this elevation is very different from the enlarged Style C of Holoclemensia. There are therefore two types of upper molar, with a pattern similar to Pappotherium, that correspond to the supposed first two molars of Holoclemensia — a larger, more symmetrical M2? and a smaller M! with a more prominent metastylar corner. The evidence suggests that Pappotherium has four molars like Holoclemensia (Bigs 50h): PM 1325 (Fig. 2, g) is the lingual part of an upper molar which seems from its size to be more probably referable to Pappotherium than to Holoclemensia. SMP-SMU 62402 (Fig. 2, }) resembles PM 999 and PM 1238 in pattern but differs from them in its smaller size. This tooth may be M! of a smaller species, of which PM 1075 might be the last molar. LOWER MOLARS The allocation of lower to upper molars is particularly difficult in the case of the therians from the Albian of Texas. This difficulty arises because Pappotherium and Holoclemensia have been dis- tinguished mainly by the buccal stylar shelf, which does not occlude with the lower teeth. The protocone, which does occlude, is missing from most upper molars. Patterson and Slaughter have distinguished six types of lower molars, Types |, 2, and 3 by Patterson (unworn examples illustrated in Fig. 3), Types 4, 5 and 6 by Slaughter. Combinations of some of these types seem probable. Teeth cf Type | are characterized by the anteroposteriorly short trigonid, in which the paraconid is lower than the metaconid, and a comparatively wide talonid on which the hypoconid is elevated and placed nearly in line with the protoconid. In Type 1 Patterson 8 BREVIORA No. 446 placed PM 1005 (Fig. 3, H) and the trigonid PM 887; to these may be added the trigonid PM 3877. Slaughter (1965) classified three teeth from Butler Farm as Type 5 (SMP-SMU 62131, 61727, and 61735). These differ from Type | in the less elevated trigonid, which has a greater anterior tilt, and in the narrower talonid. Teeth Figure 3. P, H, K, and T, left lower molars drawn in crown, lingual and pos- terior views. P, Type 2 (PM 96S, reversed), referred to Pappotherium,; H, Type 1 (PM 1005, reversed), referred to Holoclemensia; K, Type 3 (PM 1245, reversed), Kermackia; T, Type 6 (SMP-SMU 61728), holotype of Trinititherium slaughteri. S, SMP-SMU 61992, holotype of Slaughteria eruptens, crown and lingual views. 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 9 of Type 5 were referred by Slaughter (1968b) to Holoclemensia. SMP-SMU 61726, belonging to Slaughter’s Type 4, has a relatively long and narrow talonid with a less elevated hypoconid; it has lost the tops of the trigonid cusps. The differences between Types |, 4, and 5 could be due to serial position in a single species — Type 4 being the last molar, Type 5 the penultimate, and Type | the most anterior molar of the three. A trigonid, PM 966, agrees with Type | in the relative height of the paraconid and metaconid, but the trigonid is narrower transversely and less compressed anteroposteriorly. Probably because of its more open trigonid, Patterson included this specimen with Type 2, but it could be a first molar standing in the same dentition as Type 1 and bringing the number of molars to four (Fig. 5, H). Teeth of Patterson’s Type 2 have a more open trigonid than those of Type |. The metaconid is smaller; it is similar in height to the paraconid. The talonid is narrower than in Type 1. The hypoconid is comparatively low, and it is not placed so far buccally in relation to the protoconid. The talonid is preserved completely only in PM 965 (Fig. 3, P); in PM 948 the hypoconulid has been broken off. PM 930, PM 660, PM 1249, and PM 1119 are isolated trigonids. A feature of Type 2 teeth, not noted by Patterson, is the oblique crest that continues along up the posterior wall of the trigonid al- most to the top; it ends in an intermediate position between the tip of the metaconid and the notch dividing that cusp from the proto- conid. In teeth of Types | and 5, and also in PM 966, the oblique crest is confined to the lower part of the trigonid wall. Type 2 teeth are smaller than those of Types 1, 4 and 5: their trigonid widths range from 0.91 to 1.16 mm compared with 1.26 to 1.46 mm in the other types (measurements from Turnbull, 1971). No teeth of Type 2 have been reported from Butler Farm. It is probable that Types 1, 4, and 5 are referable to Holocle- mensia and Type 2 to Pappotherium. The upper molars of Holo- clemensia are larger, and they have larger metacones which might correlate with large and buccally situated hypoconids on the lower molars. The upper molars referred to Pappotherium are too small to fit lower molars of Types 1, 4, and 5, but their occlusion with Type 2 lower molars is feasible. PM 930, the Type 2 tooth with the smallest and most open trigonid, would be a first molar. The largest examples of Type 2, PM 965 and PM 660, would be second 10 BREVIORA No. 446 molars, corresponding to the largest upper molars. The smaller PM 948 and PM 1249 would be third or perhaps fourth molars (Fig. 5, P). It is to be noted that Crompton (1971) used a Type 4 tooth to illustrate the occlusion of Pappotherium. Patterson’s Type 3 was represented by only one damaged tooth (PM 922), but an unworn specimen (PM 1245; Fig. 3, K) has been recovered subsequently. It is very similar to the holotype of Ker- mackia texana, from which it differs by its higher and more pointed protoconid and metaconid. This difference is due to loss of enamel from the trigonid of the holotype. The trigonid is widely open, and the paraconid points anteriorly. The protoconid is the highest tri- gonid cusp, the paraconid the lowest. From the metaconid arises a strong posterior crest (named Distal Metacristid by Fox, 1975), continuous with the crista obliqua. The talonid is narrower than the trigonid. The hypoconid stands only slightly more buccally than the hypoconulid. The entoconid is small, and the talonid basin is open on the lingual side between the entoconid and the meta- conid. Slaughter’s Type 6 is based on the jaw fragment SMP-SMU 61728. This contains a molar (Fig. 3, T), apparently the last, and the alveoli of the next anterior tooth. Type 6 resembles Kermackia texana in size and general proportions, but the metaconid of Type 6 is much smaller, no higher than the paraconid. As in Kermackia, the crista obliqua runs to the metaconid. As the crest continues up the posterior surface of the metaconid, it is much weaker and nearly vertical. The talonid basin is widely open on the lingual side as in Kermackia, but the entoconid is almost absent, and an additional cusp is present on the crista obliqua. The top of the hypoconid has been broken off. Two trigonids from Greenwood Canyon, PM 1065 and PM 1045, have the same structure. This form is suffi- ciently distinctive to deserve a name, which I propose formally as follows: Trinititherium, gen. nov. Diagnosis. The lower molars differ from those of Kermackia as follows: (1) metaconid and paraconid subequal; (2) an accessory cusp present on the crista obliqua; (3) the crest on the posterior surface of the metaconid weaker and more vertical; (4) entoconid rudimentary. Type species: Trinititherium slaughteri, sp. nov. 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 11 Diagnosis. This is the only known species of Trinititherium. Holotype, SMP-SMU 61728, from Butler Farm. Illustrated by Slaughter (1965, Fig. 4, and 1971, Pl. 1). W. A. Clemens (personal communication) has raised the question whether Trinititherium might represent the posterior molar of Ker- mackia. This would imply a reduction of the metaconid towards the posterior end of the series. Such a process occurs as an aspect of carnassial specialization in Cenozoic carnivorous mammals (But- ler 1947). The metaconid is also reduced in Deltatheridiidae and Stagodontidae in the late Cretaceous. In all these, there is evidence of a sectorial specialization of the paraconid which is not shown by Trinititherium. The metaconid is similar in height to the paraconid in Kielantherium and probably in Aegialodon. In these genera, however, the talonid is shorter and narrower than in Trinititherium and Kermackia. Turnbull (1971) grouped together Types 1, 2, and 5 as Holo- clemensia and Types 3, 4, and 6 as Pappotherium. He included in Holoclemensia the specimens of Type 2 that are here regarded as Pappotherium. He did not consider in detail the morphological features that distinguish Type 2 from Types | and 5. He ascribed the differences to individual variation, to position in the series, and possibly to the presence of more than one species of Holoclemensia. The teeth that Turnbull identified as Pappotherium include those of Kermackia and Trinititherium, together with Type 4, regarded here as the last molar of Holoclemensia. The criteria by which he distinguished Pappotherium from Holoclemensia are those which in the present paper are used to distinguish Kermackia and Triniti- therium from Pappotherium and Holoclemensia. Kermackia had not been described when Turnbull wrote his paper, and his suppo- sition, that all the tribosphenic molars in the collection could be referred to Pappotherium or Holoclemensia, was not unreasonable. There remains to be considered the lower jaw with four teeth (SMP-SMU 61992; Fig. 3, S), referred by Slaughter (1971) to Pap- potherium. Two teeth are simple, and two are molariform. The first molariform tooth is narrower than the second and has less elevated cusps. Slaughter says the first is less worn than the second molariform tooth and identifies it as a premolar. The existence of a molariform premolar at such an early date seems very unlikely. Both molariform teeth are much more worn than the two premo- 12 BREVIORA No. 446 lariform teeth, one of which is in process of eruption. Slaughter’s statement (1971: 137) that X-rays showed no unerupted teeth ex- cludes the possibility that the first molariform tooth is a milk molar. The molariform teeth must be regarded as permanent molars. M, is similar in size to that of Kermackia; M2 is larger than M,, though it is smaller than the smallest specimen of Type 2 (PM 930). On both teeth, the tops of the trigonid cusps are broken off or worn, but the metaconid was evidently higher than the forwardly projecting paraconid. On both molars the talonid is as wide as the trigonid, and the hypoconid is buccally situated in relation to the hypoconulid. The entoconid is as high as the hypoconulid on Mb, but lower on M;. On both molars the anterior end of the crista obliqua has been obliterated through wear, and its relations to the trigonid are uncertain. However, the metaconid of M) carries a strong posterior ridge that resembles Kermackia and differs from Type 2. This specimen cannot be placed in any of the species so far described, and because of its importance I propose a name for it as follows: Slaughteria, gen. nov. Diagnosis. My differs from lower molars of Type 2, referred to Pappotherium, in that (1) the metaconid is higher than the para- conid, and (2) the metaconid has a posterior ridge, probably con- tinuous with the crista obliqua. M> differs from lower molars of Kermackia in that (1) the talonid is equal in width to the trigonid, and (2) the entoconid is larger than that of Kermackia. There is no metaconid on the lower premolars. Type species: Slaughteria eruptens, spec. nov. Diagnosis. This is the only known species of S/aughteria. Holotype, SMP-SMU 61992, from Butler Farm. Illustrated by Slaughter, 1971, Pl. 9. It is a mandibular fragment with two molars and two premolars, the anterior premolar in process of eruption. PREMOLARS The last premolar of S/aughteria is a two-rooted tooth with a well developed, one-cusped heel but no anterior cusp. The anterior premolar is smaller and seemingly has a single root. There are some isolated two-rooted premolars from Greenwood Canyon. PM 1247 (Fig. 4, a) is larger than the last premolar of Slaughteria. It has a 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 13 small posterior heel and an anterior basal cusp which is displaced towards the lingual side, indicating that the tooth is from the lower jaw. PM 1098 (Fig. 4, b) is the posterior part of a lower premolar, probably a little larger than PM 1247 and witha larger, two-cusped heel. PM 1136 (Fig. 4, c) is a much smaller tooth with no anterior ‘Ss Figure 4. Premolars and canine. a, PM 1247, lingual and crown views; b, PM 1098, lingual view; c, PM 1136, ? buccal view; d, PM 931, buccal and crown views; e, PM 1124; f, SMP-SMU 62399, lingual and crown views (from a cast); g, SMP- SMU 61947, lingual and crown views; h, SMP-SMU 61948, buccal and crown views. 14 BREVIORA No. 446 cusp or posterior heel. Its anterior profile is strongly convex, and there is a posterior crest which shows wear. None of the Texas specimens resembles the Wealden specimen that Clemens (1963, Fig. 10) described. The latter is a narrow tooth with divergent roots and a low crown, having the appearance of a milk molar. SMP- SMU 61730 and 61731 (Slaughter 1965, Fig. 5) are triconodont. The submolariform premolars that Slaughter (1968a, 1971) de- scribed are of greater interest. SMP-SMU 61947 (Fig. 4, g) is a lower premolar with a small talonid, a low, lingually placed para- conid, and a low metaconid. A crest on the buccal side from the talonid cusp continues up the posterobuccal surface of the proto- conid. In addition, there is a median talonid crest (=crista obliqua) which terminates at the base of the trigonid. SMP-SMU 62399 (Fig. 4, f) resembles SMP-SMU 61947 in having a small metaconid, but it differs from that specimen in a number of details. The tooth is narrower and the protoconid is taller. The crista obliqua con- tinues up the posterior surface of the protoconid to meet the crest that connects the protoconid with the metaconid. As the paraconid of SMP-SMU 62399 has broken off, the length of the tooth can only be estimated, but it was probably a little shorter than SMP- SMU 61947 (length about 1.4 mm, compared with 1.55 mm). It is unlikely that either of these teeth is a penultimate premolar (conventionally called P;). Among late Cretaceous mammals a metaconid is known on P; only in Gypsonictops. In that genus P; also has a partly molariform talonid, and its possession of a meta- conid can be ascribed to the unusually high level of molarization of the tooth. Such molarization is almost certainly a specialization of Gypsonictops, unlikely to be present in ancestral placentals (Butler 1977). A metaconid occurs on the last premolar (P4) in several late Cretaceous genera: Zalambdalestes, Protungulatum, Procerberus and Batodon. Both the Albian specimens are inter- preted as P;. The differences between them show that they belong to different species. SMP-SMU 61948 (Fig. 4, h) is a triangular upper premolar which Slaughter (1968a) compared with a placental P}. The lingual lobe was broken off, but a small protocone must have been present originally. The paracone is a tall cusp, similar in height to the protoconid of SMP-SMU 62399. There is no metacone. This tooth resembles P3 of Asioryctes, but whether it is a P3 or a P4 is uncer- 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN |S) tain. In Asioryctes, as in Kennalestes and Zalambdalestes, the paracone of P3 is higher than that of P4 and subequal to the proto- conid of Ps. However, P? may have been secondarily enlarged (Butler 1977). There is no metaconid on the lower premolars of Kennalestes and Asioryctes, and comparison with these genera may be misleading. In Cretaceous marsupials, such as Alphadon, the last upper premolar is the largest, and this is true also of Deltathe- ridium. : PM 931 (Fig. 4, d) is an upper premolar which differs greatly from SMP-SMU 61948. It has a metacone on the shearing crest, but no protocone. This last cusp is represented only by an indis- tinct cingulum. The strongly developed shearing crest suggests that PM 931 is a last premolar that functioned against the trigonid of M;. The absence of the protocone suggests that the corresponding lower premolar lacked a metaconid, and therefore that PM 931 did not occlude with either of the known complex lower premolars. It is too large to fit the lower dentition of Slaughteria. There are only two roots. The posterior one is flattened in cross section and ar- ranged obliquely, to support the lingual as well as the posterior apex of the crown. A similar arrangement is known in symmetro- dont molars, but PM 931 is much too large to be referred to Spa- lacotheroides. Probably some of the submolariform premolars belong to spe- cies of which the molars are already known. Only in Slaughteria have premolars been found in association with molars, and there the last premolar has no metaconid. In Slaughteria, the length of the last premolar is 65 per cent of that of M2. The two lower pre- molars with metaconids are as long as or longer than the molars of Kermackia and Trinititherium, and they are therefore probably too large to belong to these forms. Moreover in Kermackia and Trinititherium the crista obliqua on the molars runs directly to the metaconid. The possibility that the submolariform premolars be- long to Pappotherium or Holoclemensia must therefore be con- sidered. The crista obliqua of SMP-SMU 62399 resembles that of Type 2 (supposedly Pappotherium) molars in meeting the protoconid- metaconid crest, whereas that of SMP-SMU 61947 fades out near the base of the trigonid as on Holoclemensia molars. In the late Cretaceous marsupials, Alphadon and Pediomys, the last lower 16 BREVIORA No. 446 premolar is about three-quarters as long as the longest molar, and in the placental Kennalestes the proportion is about 80 per cent. SMP-SMU 61947 is 80 per cent as long as PM 1005, interpreted as M2 of Holoclemensia; the somewhat shorter SMP-SMU 62399 would bear about the same proportion to PM 965, the largest supposed Pappotherium molar. Size as well as structure is therefore consistent with the reference of SMP-SMU 61947 to Holoclemensia and SMP-SMU 62399 to Pappotherium. The upper premolar SMP-SMU 61948 would fit either of the two complex lower premolars. It possesses a strongly developed stylar cusp on the buccal edge, anterior to the metastyle. This cusp might be considered serially homologous with Style C of the molars. For this reason I refer SMP-SMU 61948, tentatively, to Holoclemensia asa. P*or P32: PM 931 (Fig. 4, d) is too small and its shearing crest too longi- tudinal to have functioned against the trigonid of PM 966, the sup- posed M, of Holoclemensia. The absence of the protocone on the premolar suggests that this cusp was relatively small on the molars. Perhaps this tooth belongs to the same species as the last molar PM 1287 (Fig. 2, p) in which the protocone is small. The mandible PM 583 (Patterson 1956, Figs. 10, 11) contains alveoli for four two-rooted premolars. The posterior alveoli are of a size to fit Kermackia molars and are probably too large for pre- molars of that genus. The last pair of alveoli, however, could have held either of the premolars with metaconids, and the simple pre- molar PM 1247 would fit the alveoli for P;. There is a possibility that this mandible belongs to either Holoclemensia or Pappothe- ium. The remarkably large canine alveolus differentiates this speci- men from SMP-SMU 62400 (Slaughter 1971, Pl. 10), a fragment containing a small, procumbent canine. PM 1124 (Fig. 4, e) is an upright two-rooted canine of small size, probably from the upper jaw. The mandible PM 583 contains alveoli for four lower incisors. Slaughter (1971, Pl. 10) described a rather large, spatulate upper incisor. DISCUSSION Number of Genera Represented The material from the Albian of Texas consists almost entirely of isolated teeth. Until more complete specimens are discovered, 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN I7/ any interpretations or conclusions must remain highly speculative. Differences in the molars, premolars, canines and incisors indicate that a number of distinct taxa are present, at least on the generic level. In attempting to allocate the various sorts of teeth, I have avoided the introduction of additional taxa as far as possible. The minimum number of taxa necessary to include the known molars seems to be the following six: 1. Holoclemensia, the largest form, is represented by several upper and lower molars; complex upper and lower premolars are allocated to this form. 2. Pappotherium, a somewhat smaller form, is represented again by several upper and lower (Type 2) molars; a complex lower pre- molar is referred to it. Holoclemensia and Pappotherium are the most common mem- bers of the fauna, but Pappotherium seems to be relatively more common at Greenwood Canyon than at Butler Farm. The tooth- less mandible may belong to either genus. 3. Slaughteria is represented by a single mandibular specimen, which is the only one to show associated molar and premolar teeth; it is smaller than Pappotherium. 4. Kermackia is a small form known only by three lower molars. 5. Trinititherium, similar in size to Kermackia, is known by one complete lower molar and two trigonids. The small upper molar SMP-SMU 62402, and the last molar PM 1075 may belong to either Kermackia or Trinititherium. 6. The last upper molar PM 1287 and the upper premolar PM 931 do not appear to belong to any of the foregoing and probably represent one or two additional species. The Family Pappotheriidae As interpreted here, Pappotherium and Holoclemensia are much alike in the organization of their molar series. Both have four molars, of which the second is the largest, and in the upper jaw, M3 is reduced posterobuccally. M! of Holoclemensia is more sym- metrical than in Pappotherium, and M3 less so, but such differences are to be found frequently in closely related genera of mammals. There is a wide stylar shelf in both genera. The trigonid is ele- vated, and it has a well developed, trenchant paraconid. The talo- nid is narrower than the trigonid. The lower molars have no cingu- 18 BREVIORA No. 446 lum except on the anterior surface of the protoconid. The last lower premolar has a metaconid and, at least in Holoclemensia, the last upper premolar has a protocone. The large Style C of Holoclemensia is represented in some speci- mens of Pappotherium by a minor elevation of the buccal marginal ridge. The metacone of Holoclemensia is larger in comparison with the paracone than in Pappotherium. The lower molars of Holo- clemensia have a more compressed trigonid, a higher metaconid, and a more buccally situated hypoconid. The crista obliqua in Holoclemensia fades out at the base of the trigonid, whereas in Pappotherium it continues up the trigonid wall. These differences seem insufficient to place the two genera in separate families, and I propose to include them both in the Pappotheriidae Slaughter 1965. Relationship of Pappotheriidae to Marsupials and Placentals The Pappotheriidae agree with the primitive marsupial A/phadon in the possession of four molars and in the wide stylar shelf of the upper molars. Holoclemensia resembles Alphadon in having a distinct Style C, but in A/phadon this is smaller than the stylocone (Style B), and it varies in size between the species. In A/phadon M3 is the largest molar, and its posterobuccal apex and metastylar blade are not reduced. A further difference appears to be that the proto- cone shelf of A/phadon is better differentiated with more strongly developed conules. In Alphadon the metacone is as tall or taller than the paracone; Holoclemensia, with its larger metacone, ap- proaches A/phadon in this respect more closely than Pappotherium does. The lower molars of Pappotheriidae resemble Alphadon in the trenchant paraconid and in the connection of the crista obliqua to the middle of the posterior trigonid wall. However, in the marsu- pials the talonid is as wide as the trigonid, sometimes wider. The hypoconulid is in a lingual position, close to the entoconid, and a cingulum, arising from the hypoconulid, passes behind the hypo- conid. In pappotheriids the trigonid, especially in Holoclemensia, is more elevated. Holoclemensia also differs from Alphadon in that the trigonid is more compressed on M2-M,. The metaconid is higher than the paraconid, and the hypoconid is higher in compari- son with the entoconid on M2 and M3. Pappotherium (i.e. Type 2) 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 19 Figure 5. Reconstructions of cheek teeth in occlusion. P. Pappotherium patter- soni; H, Holoclemensia texana; A, Alphadon marshi (a marsupial); C, Cimolestes incisus (a placental). Specimens used in the reconstruction of Pappotherium: M', PM 999; M?, PM 894; M3 and Mé, the holotype; Ps, SMP-SMU 62399; Mi, PM 930; M2, PM 965; M3, PM 948. Specimens used in the reconstruction of Holoclemensia: P*, SMP-SMU 61948; M!, PM 1000; M2, PM 886; M3, the holotype; M4, the paratype; Ps, SMP-SMU 61947; Mi, PM 966; M2, PM 1005; Ms, SMP-SMU 61727; Ms, SMP-SMU 61726. 20 BREVIORA No. 446 agrees more with Al/phadon in these characters, as was noted by Clemens (1966:16—17). Albertatherium (Fox 1971), from the early Campanian, ap- proaches Holoclemensia in that Style C is higher than the stylocone, and the metaconid is somewhat higher than the paraconid. How- ever, the stylar shelf of Albertatherium is narrower than in Holo- clemensia. The metacone and metastyle of M3? are larger, and the conules are better developed. On the lower molars, the trigonid is less compressed; the talonid wider; and the hypoconulid is lingual in position, and the entoconid is higher. As Fox (1971) noted, Albertatherium does not link Alphadon with Holoclemensia. Style C is also enlarged in Aquiladelphis, an early Campanian pediomyid in which the stylocone is reduced to two small cusps. In other re- spects this form resembles Holoclemensia even less than Alberta- therium; for example, the crista obliqua meets the posterior wall of the protoconid. The primitive eutherian molar pattern is believed to be preserved best in the late Cretaceous Cimolestes and Kennalestes (Butler, 1977). Holoclemensia and Pappotherium differ from these in the retention of the fourth molar, in the wide stylar shelf with better developed styles posterior to the stylocone, and in the correspond- ingly weaker development of the protocone shelf. Lower molars of Pappotheriidae agree with those of primitive placentals in the high trigonid and comparatively narrow talonid, in the connection of the crista obliqua to the middle of the trigonid wall, and in the ab- sence of the posterior cingulum. Holoclemensia, not Pappothe- rium, agrees with Cimolestes and Kennalestes in that the metaconid is higher than the paraconid. If the tooth has been correctly re- ferred, the possession of a premolar protocone would constitute another resemblance of Holoclemensia to placentals. However, there is no metaconid on P, in Cimolestes or Kennalestes. Fox (1975) noted a resemblance in the outlines of the last two upper molars between Holoclemensia and the late Cretaceous Del- tatheroides, which has four molars. He concluded that Holocle- mensia must have four molars. Because the penultimate molar of Pappotherium shows less reduction in the metastyle, Fox inferred that only three molars were present in that genus. I find his argu- ment unconvincing; the relationship between the shapes of teeth and their numerical position is too inconstant for such deductions 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 21 to be made. Late Cretaceous didelphids have four molars, and yet the metastyle of their penultimate molar is more developed than in Pappotherium. Deltatheroides, in which the metastyle of M3? is reduced, can be regarded as being in the process of losing its M4, which is absent in Deltatheridium (Kielan-Jaworowska 1975); Holoclemensia might by analogy be considered to be evolving towards the three-molar state characteristic of placentals. It seems impossible to fit Holoclemensia and Pappotherium into a simple metatherian-eutherian framework. They have a mixture of characters which indicate that they belong to neither infraclass. Though Holoclemensia approaches primitive marsupials in some ways, Pappotherium is more like them in others; Holoclemensia also shows some resemblances to primitive placentals. No doubt some of the characters that pappotheriids share with marsupials or placentals are plesiomorphs inherited from a common ancestor, which may include the wide stylar shelf of primitive marsupials and the narrow talonid of primitive placentals. The fourth molar is probably another plesiomorphous character lost in placentals but retained in Deltatheridiidae (Butler & Kielan-Jaworowska, 1973; Kielan-Jaworowska, 1975) as well as in marsupials. Other shared characters may be due to parallel evolution, such as the enlarged Style C. The metaconid of Ps; may also have evolved independently, for it occurs in some placentals but not in others. Placentals have been reported from Khovboor in Mongolia (Beliajeva et al., 1974; Dashzeveg, 1975) in a deposit that is be- lieved to be of Aptian age. The fauna, still undescribed, is said to contain forerunners of the much later Djadokhta fauna. If the ‘Aptian age of these specimens is correct, the divergence of placen- tals and marsupials must already have taken place at that time. The Albian Pappotheriidae would in that case be too late to be the common ancestors of the two infraclasses. Clemens (1971: 174) states that “The evidence . . . does not exclude the possibility that when the tribosphenic grade of dental evolution was attained an adaptive radiation occurred producing lineages in addition to those ancestral to marsupials and placentals.” The Pappotheriidae may well have belonged to one such lineage. During much of the Cretaceous the land surface was more than usually fragmented by epicontinental seas (Lillegraven, 1974; Tel- ford, 1974). North America was divided, from middle Albian time 22. BREVIORA No. 446 onwards, into eastern and western land-masses by the Interior Sea- way. The northern part of the Atlantic was closed: the seaway between Greenland and Canada developed only in the Campanian (Martin, 1973). Europe was separated from Asia by the Turgai Strait. In mid-Cretaceous time, eastern North America and Europe thus formed a single continent from which the mammals from Texas are the only known representatives of their class. Following Lille- graven (1974) one may imagine a wide dispersal, in the early Cre- taceous, of an ancestral group with tribosphenic molars, of which Aegialodon, from the English Wealden, is the only known example. Subsequently isolated on different land-masses, the ancestral group gave rise to the placentals in Asia and the marsupials in western North America (or as Telford (1974) suggests, in South America). Possibly the Pappotheriidae represent another lineage that evolved on the Euramerican continent, but subsequently became extinct. A molar from the Campanian of Champ-Garimond, France (Le- doux et al., 1966; Butler, 1977) might conceivably be a pappotheriid derivative, but it is more advanced, and it might equally well be a placental immigrant from Asia. Kermackiidae, Fam. nov. Kermackia and Trinititherium are distinguished from Pappo- therium and Holoclemensia by the continuation of the crista ob- liqua to the tip of the metaconid, so that on the posterior surface of that cusp it forms a ridge (the distal metacristid of Fox, 1975). The talonid is narrow, its basin is widely open lingually, and the entoconid is small or rudimentary. These genera are at a lower level of molar evolution than the Pappotheriidae. In the latter the con- nection of the crista obliqua to the trigonid has shifted to a more buccal position, the talonid is wider, and the entoconid is larger. Dashzeveg (1975) noted the resemblance of Kermackia to Aegialo- don and Kielantherium, which belong to the family Aegialodonti- dae Kermack (1957). However, these differ from Kermackia in having the talonid reduced in length as well as in width. Because of this difference I propose the family Kermackiidae to include Kermackia and Trinititherium. The Kermackiidae resemble the Aegialodontidae in retaining the distal metacristid but differ from them in that the talonid is equal in length to the trigonid. The 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 23 Aegialodontidae, Kermackiidae and Pappotheriidae represent three stages in the evolution of the talonid (Fig. 6). Slaughteria, which probably possesses a distal metacristid, has a wide talonid, and (on Mb, but not on M_) a fully developed en- toconid like the Pappotheriidae. It may represent another family, but, as some doubt remains over the details of the molar pattern, I prefer to place Slaughteria provisionally in the Kermackiidae. To judge from the relative size of the penultimate premolar, the num- ber of premolars may bave been reduced. S/aughteria is the only Albian therian for which there is evidence of tooth replacement, and on this character, Slaughter (1971) placed it in the Eutheria. However, diphyodonty occurs in pantotheres (Butler & Krebs, 1973), and it must be considered a plesiomorphous character, re- Figure 6. Left lower molars in lingual view, to illustrate possible evolutionary radiation in the early Cretaceous. a, Peramus (Pantotheria); b, Aegialodon; e,Kiel- antherium (Aegialodontidae); c, Kermackia; d, Trinititherium (Kermackiidae); f, Deltatheridium (Deltatheridiidae); g, Holoclemensia; h, Pappotherium (Pappo- theriidae); 1, Alphadon (Metatheria); j}, Champ-Garimond molar; k, Kennalestes (Eutheria). 24 BREVIORA No. 446 tained in placentals and lost (except for the last premolar) in mar- supials. CLASSIFICATION ABOVE THE FAMILY LEVEL A name is required for an infraclass to include those therians that have a functional protocone and have advanced beyond the pantotherian stage, but nevertheless cannot be regarded either as marsupials or placentals. Turnbull (1971) proposed a cohort Tribosphenata, equal in rank with his Marsupiata and Placentata, including all within the infraclass Eutheria. The Tribosphenata comprised two orders: (1) Tribosphena, containing Pappotherium, Holoclemensia, Aegialodon and Peramus, and (2) Zalambdodonta, consisting of a number of placentals and marsupials with zalamb- dodont molar teeth. Since zalambdodont molar patterns have been derived, more than once, from dilambdodont (“euthemorphic’”) pat- terns (Butler 1972), I consider the order Zalambdodonta to be arti- ficial. I propose the name Tribotheria for an infraclass standing between the Pantotheria, on the one hand, and the Metatheria and Eutheria on the other. The Tribotheria differ from the Pantotheria in having reached the tribosphenic stage of molar evolution by the acquisition of a protocone that functioned against the talonid. The infraclasses Metatheria and Eutheria originated from members of the Tribo- theria, but there were other, less diversified lineages, such as the Deltatheridiidae and Pappotheriidae, which may be left in the Tr- botheria. Turnbull’s order Tribosphena was similar in conception, but it included Peramus, which lacks a functional protocone. For Tribotheria with a distal metacristid I propose an order Aegialodontia. This corresponds to stage IIA in the scheme of molar evolution tabulated by Fox (1975, Table 1). The Aegialo- dontia includes the families Aegialodontidae and Kermackiidae. The late Cretaceous Deltatheridiidae may also be placed here. Their lower molars show a considerable resemblance to Kielantherium, and it seems likely that they have an aegialodontid ancestry, from which they have retained the small talonid. Potamotelses (Fox 1975) may represent another family in the Aegialodontia. For the Pappotheriidae, which have lost the distal metacristid, and are at a stage of evolution comparable to the primitive marsupials and placentals, I propose a new order Pappotherida. 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 25 The classification proposed may be tabulated: Subclass Theria Infraclass Pantotheria Infraclass Tribotheria Order Aegialodontia Family Aegialodontidae (Aegialodon, Kielantherium) Family Kermackiidae (Kermackia, Trinititherium, ? Slaugh- teria) Family Deltatheridiidae (Deltatheroides, Deltatheridium) Family ? (Potamotelses) Order Pappotherida Family Pappotheriidae (Pappotherium, Holoclemensia) Infraclass Metatheria Infraclass Eutheria SUMMARY 1. At least six genera of mammals with tribosphenic molars are represented in the collection, but most specimens can be referred to Pappotherium and Holoclemensia. 2. There are four types of upper molar of Holoclemensia pat- tern, and four of Pappotherium pattern, showing that both these genera had four molars. 3. Of the six types of lower molar distinguished by Patterson and Slaughter, Types 1, 4, and 5 are referred to Holoclemensia and Type 2 (except for PM 966) to Pappotherium. 4. Lower molars of Type 3 are Kermackia, and those of Type 6 are named Trinititherium slaughteri gen. et sp. nov. 5. The two lower premolars with metaconids are referred to Holoclemensia and Pappotherium respectively, and the upper pre- molar with a protocone is referred to Holoclemensia. 6. The mandible SMP-SMU 61992, which Slaughter (1971) re- ferred to Pappotherium, has two molars and two premolars. It is named S/aughteria eruptens gen. et sp. nov. It is the only specimen that provides evidence of tooth replacement. 7. Pappotherium and Holoclemensia are united in the family Pappotheriidae. They are neither marsupials nor placentals, but represent a separate line of evolution. 8. Kermackia and Trinititherium are placed in a new family Kermackiidae. They are at a more primitive stage of molar evo- 26 BREVIORA No. 446 lution than the Pappotheriidae. S/aughteria is provisionally placed in the Kermackiidae. 9. A new infraclass Tribotheria is proposed to include mammals with tribosphenic molars that are not classifiable as marsupials or placentals. This infraclass includes two new orders, Aegialodontia and Pappotheria. ACKNOWLEDGMENTS I thank W. Turnbull and B. Patterson for assistance with the examination of the specimens from Greenwood Canyon. B. Slaugh- ter very kindly allowed me to study material from Butler Farm at the time unpublished, and provided casts. B. Patterson, Z. Kielan- Jaworowska and W. A. Clemens read draft manuscripts, and I am much indebted to them for helpful suggestions. LITERATURE CIVED BELIAJEVA, E. I., B. A. TROFIMOV, AND V. J. RESHETOV. 1974. (General Stages in evolution of late Mesozoic and early Tertiary mammalian fauna in Central Asia). Fauna i biostratigrafia Mesozoa 1 Kainozoa Mongolii. Trudy Sovm. Sov.-Mong. Pal. Exp. Moscow, 1: 19-45. BuTLeER, P. M. 1947. The evolution of carnassial dentitions in the Mammalia. Proc. Zool. Soc. London, 116, Part II: 198-220. BUTLER, P. M. 1972. The problem of insectivore classification. Jn Joysey, K. A. and T. S. Kemp (Eds.), Studies in Vertebrate Evolution. Edinburgh: Oliver & Boyd: 253-265. BUTLER, P. M. 1977. Evolutionary radiation of the cheek teeth of Cretaceous placentals. Acta palaeont. Polon., 22: 241-271. BuTLeR, P. M. AND Z. KIELAN-JAWOROWSKA. 1973. Is Deltatheridium a mar- supial? Nature, 254: 105-106. BUTLER, P. M. AND B. KREBS. 1973. A pantotherian milk dentition. Palaont. Zschr., 47: 256-258. CLEMENS, W. A. 1963. Wealden mammalian fossils. Palaeontology, 6: 55-69. CLEMENS, W. A. 1966. Fossil mammals of the type Lance Formation, Wyoming. Part II Marsupialia. Univ. Calif. Publ. Geol. Sci. 62: 1-122. CLEMENS, W. A. 1971. Mammalian evolution in the Cretaceous. Jn Kermack, D. M. and K. A. Kermack (Eds.), Early Mammals. Zool. J. Linn. Soc., 50, Suppl. 1: 165-180. CROMPTON, A. W. 1971. The origin of the tribosphenic molar. Jn Kermack, D. M. and K. A. Kermack (Eds.), Early Mammals. Zool. J. Linn. Soc., 50, Suppl. 1: 131-143. DASHZEVEG, D. 1975. New primitive therian from the early Cretaceous of Mon- golia. Nature, 256: 402-403. 1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN iy Fox, R. C. 1971. Marsupial mammals from the early Campanian Milk River Formation, Alberta, Canada. Jn Kermack, D. M. and K. A. Kermack (Eds.), Early Mammals. Zool. J. Linn. Soc., 50, Suppl. 1: 145-164. Fox, R. C. 1975. Molar structure and function in the early Cretaceous mammal Pappotherium: evolutionary implications for Mesozoic Theria. Canad. J. Earth Sci., 12: 412-442. HOFFSTETTER, R. 1970. L’histoire biogeographique des Marsupiaux et la dichot- omie Marsupiaux-Placentaires. C. R. Acad. Sci., Paris, Ser. D, 271: 388-391. HOFFSTETTER, R. 1972. Donnees et hypotheses concernant l’origine et l'histoire biogeographique des Marsupiaux. C. R. Acad. Sci., Paris, Ser. D, 274: 2635- 2638. KERMACK, K. A. 1967. The Aegialodontidae — a new family of Cretaceous mam- mals. Proc. geol. Soc. London, 1640: 146. KIELAN-JAWOROWSKA, Z. 1975. Evolution of the therian mammals in the late Cretaceous of Asia. Part I. Deltatheridiidae. Palaeont. Polon., 33: 103-132. Lepoux, J.-C., J.-L. HARTENBERGER, J. MICHAUX, J. SUDRE, AND L. THALER. 1966. Découverte d’un mammifere dans le Cretacé Supérieur a dinosaures de Champ-Garimond pres de Fons (Gard). C. R. Acad. Sci., Paris, Ser. D, 262: 1925-1928. LILLEGRAVEN, J. A. 1969. Latest Cretaceous mammals of upper part of Edmon- ton Formation of Alberta, Canada, and review of marsupial-placental dichot- omy in mammalian evolution. Univ. Kansas paleont. Contr., 50: 1-122. LILLEGRAVEN, J. A. 1974. Biogeographical considerations of the marsupial-pla- cental dichotomy. Ann. Rev. Ecol, Syst., 5: 263-283. MARTIN, R. 1973. Cretaceous — early Tertiary rift basin of Baffin Bay — con- tinental drift without sea-floor spreading. /n Pitcher, M. G. (Ed.), Arctic Geol- ogy. Amer. Assoc. Petrol. Geol. Mem. 19: 500-505. PATTERSON, B. 1956. Early Cretaceous mammals and the evolution of mamma- lian molar teeth. Fieldiana: Geol., 13: 1-105. SLAUGHTER, B. H. 1965. A therian from the lower Cretaceous.(Albian) of Texas. Postilla, 93: 1-18. SLAUGHTER, B. H. 1968a. Earliest known eutherian mammals and the evolution of premolar occlusion. Texas J. Sci., 20: 3-12. SLAUGHTER, B. H. 1968b. Earliest known marsupials. Science, N. Y., 162: 254- DSS: SLAUGHTER, B. H. 1968c.. Holoclemensia instead of Clemensia. Science, N. Y., 162: 1306. SLAUGHTER, B. H. 1971. Mid-Cretaceous (Albian) therians of the Butler Farm local fauna, Texas. Jn Kermack, D. M. and K. A. Kermack (Eds.), Early Mam- mals. Zool. J. Linn. Soc., 50, Suppl. 1: 131-143. TELFORD, R. H. 1974. Marsupials and the new paleogeography. Jn Ross, C. A. (Ed.), Paleogeographic Provinces and Provinciality. Soc. Economic Paleontol- ogists and Mineralogists, Sp. Publ. 21: 109-126. TURNBULL, W. D 1971. The Trinity therians: their bearing on evolution in mar- supials and other therians. /n Dahlberg, A. A. (Ed.), Dental Morphology and Evolution: 151-179. Chicago: Univ. Chicago Press. LILI CAA MAR 1 8 1985 BREVIORA Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, MASS. 20 DECEMBER 1978 NUMBER 447 FOOD SELECTION BY BEAVERS: SAMPLING BEHAVIOR STEPHEN H. JENKINS! ABSTRACT. Beavers in central Massachusetts frequently remove small pieces of bark (less than 25 cm?) from standing trees. After removing a sample, the beavers may abandon these trees. At two ponds studied by the author in fall 1973, there were significant differences in species composition of trees sampled, felled, and available near the ponds. If beavers are physiologically capable of measuring the nutritional value of small pieces of bark, then sampling may be an economical way for them to assess spatial and temporal variation in relative nutritional value of different tree species. INTRODUCTION During a recent study of food selection by beavers (Castor can- adensis) at two ponds in central Massachusetts (Jenkins, 1974), I observed numerous trees from which the beavers had removed small patches of bark (often less than 25 cm2). Without using them further, the beavers then abandoned these trees. During September and October 1973, beavers at Blue Heron Cove cut into or partially stripped the bark from the lower trunks of 46 trees without com- pletely felling them. At Tamplin Road Pond, there were 36 such trees. The beavers removed less than 25 cm? of bark from 25 of the 46 trees at Blue Heron Cove and the same amount from 17 of the 36 trees at Tamplin Road Pond. Chabreck (1958) reported that beavers in Louisiana stripped the bark from the lower trunks of many trees without felling them. Aldous (1938) and many others have noted that beavers may not use all the bark of the trees they fell. The present study is the first to describe and to consider the possible adaptive significance in the beavers’ pattern of removing small pieces of bark (less than 25 cm?) ‘Department of Biology, University of Nevada, Reno, Nevada 89557 2 BREVIORA No. 447 from standing trees. Secondly, the study compares the beavers’ patterns of selection of species for sampling with their patterns of selection of species for more complete utilization. For economy of expression, I will call the beavers’ incomplete use-of trees (described above) sampling. | define light sampling as use of less than 25 cm? of bark near the base of a tree without fell- ing it and heavy sampling as cutting into the wood of a tree and/or stripping more than 25 cm? of bark near its base without felling it. Several authors (Rozin, 1969; Freeland and Janzen, 1974; Westoby, 1974) have attached functional connotations to the word sampling in their discussions of optimal foraging. I wish to avoid a specific functional connotation in using the word. Instead, I present six alternative hypotheses which could account for beavers’ incomplete use of trees. The first of these hypotheses argues that beavers’ sampling might not be of direct adaptive value but is rather an incidental byproduct of the foraging process. The remaining five suggest five different ways in which sampling might function as an adaptive component of a beaver’s foraging strategy. My data are insufficient to discriminate completely among these hypotheses; I present them to establish that sampling by beavers is a phenomenon worth further study, especially in the context of recent developments in optimal foraging theory (Krebs et al., 1977; Pyke et al., 1977). RESULTS AND DISCUSSION Table | shows the generic distributions of trees sampled, felled, and available at the major tree cutting sites of Blue Heron Cove and Tamplin Road Pond between | September 1973 and 6 Novem- ber 1973 (Blue Heron Cove) or 2 November 1973 (Tamplin Road Pond). The following hypotheses might explain beavers’ tree sam- pling behavior. (1) A beaver may be interrupted while starting to fell a tree. Sampled trees simply represent those abandoned because of such interruption. If the probability of interruption is independent of tree genus, and if the time to remove less than 25 cm? of bark from a tree is less than the time to fell even a small tree (but one more than 2.5 cm diameter, the lower size limit considered in this study), then this hypothesis predicts that the generic distributions of lightly sampled and felled trees should be the same. (Heavily sampled 1978 FOOD SELECTION BY BEAVERS 3) TABLE | Generic Distributions of Trees Sampled, Felled, and Available at Blue Heron Cove and Tamplin Road Pond, Fall 1973 Lightly Heavily Sampled* Sampled** Felled Availablet Blue Heron Cove Oak (Quercus spp.) 11 (44%) 16 (76%) 31 (51%) 63 (25%) Maple (Acer spp.) 4 (16%) 1 ( 5%) 14 (23%) 97 (38%) Birch (Betula spp.) 9 (36%) 3 (14%) 1 ( 2%) 41 (16%) Others 1 ( 4%) 1 ( 5%) 15 (25%) 53 (21%) Totals 25 21 61 254 Tamplin Road Pond Oak (Quercus spp.) 6 (35%) 7 (37%) 2 ( 6%) 15 (12%) Maple (Acer spp.) 4 (24%) 9 (47%) 12 (34%) 59 (45%) Cherry (Prunus spp.) 5 (29%) 3 (16%) 15 (43%) 18 (14%) Others 2 (12%) 0 ( 0%) 6 (17%) 38 (29%) Motals~ 17 19 35 130 *Less than 25 cm? of bark removed. **Wood cut and/or more than 25 cm? of bark removed. +Based on randomly placed, 78.5 m?, circular quadrats. See Jenkins (1975) for details. trees are excluded because it may take less time to fell a small tree than to sample heavily a large tree, and a difference in the generic distributions of sampled and felled trees might arise from differ- ences in the available diameter distributions of various genera.) At Blue Heron Cove, generic distributions of lightly sampled and felled trees are significantly different (y° = 15.0, 2 df, p< 0.001, only oak, maple, and birch considered because few trees of other genera were sampled). At Tamplin Road Pond, these distributions are not quite significantly different (x’ = 5.2, 2 df, 0.05 < p < 0.10, only oak, maple, and cherry considered). This hypothesis may account for the existence of some sampled trees. However, it does not explain the marked difference in generic distributions of felled and sampled trees at Blue Hercn Cove (Table 1). 4 BREVIORA No. 447 (2) Beavers cannot identify trees taxonomically without tasting their bark, so sampling is necessary for selection of preferred tree genera. If this is true, the generic distribution of trees felled or sampled (the sum of columns I, 2, and 3 in Table 1) should match that of trees available. The significant differences between these distributions at both ponds (Blue Heron Cove: x” = 50.4, 3 df, p< 0.001; Tamplin Road Pond: y* = 31.3, 3 df, p< 0.001) cast doubt on this hypothesis. (3) Sampling is used to avoid ingestion of potential toxins and demonstrates cautiousness with novel food items (Freeland and Janzen, 1974). This hypothesis suggests that unfamiliar food types will be sampled more than familiar ones. Comparison of lightly sampled and felled trees at both Blue Heron Cove and Tamplin Road Pond refutes this hypothesis. In particular, oak is the genus most often sampled as well as felled at Blue Heron Cove, and cherry is most often felled and also frequently sampled at Tamplin Road Pond. Cautiousness in using new, potentially toxic foods may lead to sampling behavior in some species, but it does not explain these instances of sampling behavior by beavers. (4) Sampling satisfies a requirement for certain vitamins or minerals, needed in small quantities and available only in some tree species. This hypothesis fails to explain the extensive sampling of maples and oaks at Blue Heron Cove and maples and cherries at Tamplin Road Pond. Many trees of these species are felled. The felled trees should supply any micronutrients which may be uniquely available in these species, unless the felled trees have con- centrations of particular micronutrients several hundredfold less than sampled trees of the same species. (5) Sampling represents practice at tree selection by young ani- mals. I have no data on the amount of sampling done by individual members of the two colonies. However, young animals could prac- tice tree selection on branches and logs brought to the edge of the pond by adults with less risk of predation than if the young animals sampled standing trees at distances as great as 50 m from shore. Therefore, it is unlikely that sampling would evolve solely to serve this function. (6) Sampling is used by beavers to assess possible spatial and temporal variation in the nutritional quality of alternative foods. Unlike some of the others, this hypothesis cannot be tested by 1978 FOOD SELECTION BY BEAVERS 5 comparing generic distributions of trees sampled, felled, and avail- able. I present evidence elsewhere that beavers select different genera of trees for cutting at different sites around a pond (Jenkins, 1975), and different genera in different years at the same site (Jen- kins, in preparation). There is evidence of spatial (Cowan et al., 1950) and temporal (Kramer and Koslowski, 1960) variation in nutritional value of some tree species. It seems reasonable to sug- gest that the spatial and temporal variation in beavers’ tree selec- tion is related to spatial and temporal variation in relative nutri- tional value of different tree species. Sampling may be a mechanism for achieving an optimal diet in the face of such variation. CONCLUSIONS Two kinds of data, both difficult to collect, are needed to prove more conclusively which of the above hypotheses are incorrect. First, direct observations of beavers in the act of sampling would help decide the validity of hypotheses | and 5. Unfortunately, bea- vers are primarily nocturnal, so special methods (e.g., radio telem- etry, use of night vision instruments) would be needed to assist in making such observations. Data might be obtained on sampling during twilight hours without such assistance. However, such data probably would be inconclusive. For example, failure to observe sampling by young animals during twilight would certainly not prove that young animals don’t sample; they might in fact be active samplers, but only at night. Second, data on toxin and nutrient content of bark of individual trees would help decide the validity of hypotheses 3, 4, and 6. To get adequate data of this sort would require analyzing a large number of samples for numerous nutrients and possible toxins (since both intraspecific and interspecific, as well as spatial and temporal variations in these attributes of tree bark are important components of these hypotheses). In addition, some of the toxins probably are unidentified or incompletely char- acterized at the present time. In this note, I show that beavers are highly selective in sampling trees, but that they choose genera in different proportions than in felling trees. I suggest that sampling may be an economical way for beavers to assess spatial and temporal variation in relative nu- tritional value of different tree species, assuming they are capable of measuring the nutritional value of small pieces of bark. The 6 BREVIORA No. 447 physiological and behavioral mechanisms by which sampling func- tions are important subjects for future research. ACKNOWLEDGMENTS I thank R. Fagen, H. Hodgdon, and T. Schoener for critically reviewing this paper, and I thank K. Jenkins for help in collecting the data. LITERATURE CITED ALpbous, S. A. 1938. Beaver food utilization studies. J. Wildl. Manage., 2: 215- 222% CHABRECK, R. H. 1958. Beaver-forest relationships in St. Tammany Parish, Louisiana. J. Wildl. Manage., 22: 104-107. Cowan, I. McT., W. S. HOAR, AND J. HATTER. 1950. The effect of forest suc- cession upon the quantity and upon the nutritive values of woody plants used as food by moose. Can. J. Res., Sect. D, Zool. Sci., 28: 249-271. FREELAND, W. J., AND D. H. JANZEN. 1974. Strategies in herbivory by mammals: the role of plant secondary compounds. Amer. Nat., 108: 269-289. JENKINS, S. H. 1974. Food selection by beavers. Unpublished Ph.D. dissertation, Harvard Univ., Cambridge, Ma., 165 pp. 1975. Food selection by beavers: a multidimensional contingency table analysis. Oecologia (Berl.), 21: 157-173. 1978. Seasonal and year-to-year differences in food selection by beavers. In preparation. KRAMER, P. J., AND T. T. KoSLowSKI. 1960. Physiology of trees. McGraw-Hill, New York, 642 pp. Kress, J. R., T. ERICKSEN, M. I. WEBBER, AND E. L. CHARNOV. 1977. Optimal prey selection in the Great Tit (Parus major). Anim. Behav., 25: 30-38. Pyke, G. H., H. R. PULLIAM, AND E. L. CHARNOV. 1977. Optimal foraging: a selective review of theory and tests. Quart. Rev. Biol., 52: 137-154. Rozin, P. 1969. Adaptive food sampling patterns in vitamin deficient rats. J. Comp. Physiol. Psychol., 69: 126-132. Westosy, M. 1974. An analysis of diet selection by large generalist herbivores. Amer. Nat., 108: 290-304. BREVIO Rex Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, MASS. 20 DECEMBER 1978 NUMBER 448 SYSTEMATIC NOTES ON THE LOONS (GAVIIDAE: AVES) ROBERT W. STORER! ABSTRACT. The following discussion documents the treatment of the loons for the revision of Volume | of Peters’ Check-list of Birds of the World. The family is thought to be most closely related to the Charadriformes. The Cretaceous genera Enaliornis and Lonchodytes are thought to be convergent with rather than related to the loons (contra Brodkorb). Gavia immer and G. adamsii are considered full species but members of the same superspecies, while pacifica is considered conspecific with arctica. The subspecies Gavia stellata squamata, G. arctica suschkini, and G. immer elasson are not recognized. The sequence of forms followed 1s: stellata, arctica pacifica, a. arctica, a. viridigularis, immer, adamsii. I. INTRODUCTION The purposes of this paper are to present reasons for decisions made in preparing the section on the Gaviidae for the revision of Volume | of Peters’ Check-list of Birds of the World and to present some preliminary information gained while examining and measuring specimens of loons preparatory to making an analysis of geographic variation within the family. To date, I have examined over 2200 study skins of loons but still need to study material in several important collections before the final analysis can be made. For a group of only four species, there have been many diver- gences of opinion on such matters as the systematic position of the family, the sequence of species within the genus, the specific status of some of the forms, and the validity of several of the subspecies. 'Museum of Zoology The University of Michigan Ann Arbor, Michigan 48109 2 BREVIORA No. 448 Il. SYSTEMATIC POSITION OF THE LOONS There is general agreement that the living loons belong to the single genus Gavia. This, with one or two genera known only as fossils, constitutes the family Gaviidae. Several fossil genera re- ferred to families of uncertain systematic position have also been included in the order Gaviiformes, which otherwise contains only the Gaviidae. These fossils are discussed below. The varied opinions on the systematic position of the living loons within the class Aves have been well summarized by Sibley and Ahlquist (1972: 53-58). On the basis of fossil evidence (Storer, 1956) and studies of egg-white proteins (Sibley and Ahlquist, /oc. cit.), loons are believed most closely related to the gulls, auks, and shorebirds. Ideally, they should be placed next to or near the Charadriiformes, but this is impractical as only the first volume of Peters’ check-list is being revised at present. Itt, SEQUENCE OF SPECIES In presenting a linear sequence of species, it is customary to start with the most primitive and end with the most advanced. In the case of the loons, there is fossil evidence bearing on the matter of primitiveness. The known fossil record of these foot-propelled diving birds goes back to the Upper Eocene of England (Colymboides anglicus) and the Aquitanian deposits (late Oligocene or early Miocene) of France (C. minutus) and probably the Oligocene of North America (Gavi- ella pusilla). The Lower Cretaceous genus Enaliornis and the Upper Cretaceous genus Lonchodytes have been considered as families of the same order (Gaviiformes) as the loons by Brodkorb (1963a: 220-221: 1963b: 56-60). However, the material of these genera is fragmentary, and the pieces of leg bones resemble those of loons in showing adaptations for swimming and diving but in other respects are quite different from them. The distal end of the carpometa- carpus of Lonchodytes pterygius, as shown in Brodkorb’s figure (1963b: 59), differs so markedly in the relative sizes, angles, and positions of the distal articulations from those of the loons as to cast considerable doubt in my mind on the alleged phylogenetic relation- ship between Lonchodytes and Gavia. 1978 SYSTEMATIC NOTES ON THE LOONS 3 If, as I am convinced, these Cretaceous genera are convergent with the loons and if we accept Brodkorb’s comment that the position of the little-known Upper Paleocene genus Eupterornis is “tentative,” the earliest known loons (Colymboides and Gaviella) were smaller than later ones. The best-known species (C. minutus) was about the size of a teal, and, judging from the size of its wings, a strong flier (Storer, 1956: 423). The earlier species (C. anglicus) is poorly known but was somewhat larger and apparently had rela- tively shorter wings than C. minutus. It may have “represented a specialised offshoot from a more generalised gaviid stem, rather than a direct ancestral form of either the later Gavia species or C. minutus” (Harrison, 1976: 66). The Oligocene species, Gaviella pusilla, was also smaller than the species of Gavia. Known only from the proximal end of a carpometacarpus, Gaviella differs from other loons in several respects and is probably not closely related to them (Wetmore, 1940: 30). Thus the limited evidence from fossils indicates that Gavia evolved from smaller birds which were less well adapted for diving and had relatively larger wings. While the modern genus Gavia contains several Pliocene species, studies of these species have not as yet provided information regarding the relative antiquity or phylogenetic relationships among the Recent species. The Red-throated Loon, Gavia stellata, is the most distinct species in both its winter and nuptial plumages, and for this reason it should be placed at one end of the sequence. The square white spots on the upperparts and the well-developed stripes and iri- descence on the head and neck of the other three species can be considered advanced characters. G. stellata is the smallest of the living loons and also has the lowest wing loading. To estimate this, I divided the weight of the bird by the square of the length (arc) of the folded wing. The means of this index for ten specimens of each of the species are as follows: stellata 2.00, a. pacifica 2.25, immer 3.03, and adamsii 3.17 grams/cm?. Although detailed comparative stud- ies have not been made, ste//ata is said to be able to take off from water more easily than the other loons and alone among them can take off from land (Palmer, 1962: 59). As pointed out above, small size and greater flying ability may be considered primitive charac- ters in loons, so I have placed stellata first on the basis of its relatively simple nuptial plumage, small size and good flying ability. 4 BREVIORA No. 448 As members of a superspecies, adamsii and immer should be placed next to each other. Arctica has been known to hybridize with immer (Hunter and Dennis, 1972; Robertson and Fraker, 1974) and therefore these two species should be adjacent on the list. Assuming the largest species to be the most advanced and lacking other strong evidence to the contrary, I prefer the sequence arctica, immer, adamsii to the reverse for these three species. If one considers the forms of arctica to represent a broken rassenkreis, as I do, the sequence should be either pacifica, arctica, viridigularis or the reverse. I prefer starting with the smaller, better-known form, and thus the sequence I have adopted is: stellata, arctica pacifica, a. arctica, a. viridigularis, immer, adamsii. IV. SPECIES PROBLEMS THE COMMON AND YELLOW-BILLED LOONS, G. immer and G. adamsii. — The breeding ranges of these forms were presumably separated during the last glaciation — that of immer lying south of the glaciers and that of adamsii in Yukon-Bering Sea refugia to the north (Rand, 1948: 317-318). Of the principal differences between the species, the color of the bill and of the iridescence on the head and the head patterns may have been selected for as reproductive isolating mechanisms; all are located where major specific differ- ences among other groups of diving birds such as penguins and grebes are found. On the other hand, evolution in bill shape in adamsii was presumably related to feeding. The upturned mandibles of this species and of ste//ata appear to be adaptations for bottom foraging. When birds are moving close over the bottom, the bill must be held below the body to catch bottom-living prey. In this position of the head, the lower part of the upturned mandible would lie nearly parallel to the bottom, making capture of prey in this situation easier than if the bill were shaped like those of immer and arctica. The upturned bills of adamsii and stellata are probably an example of convergence, because there is no other reason to consider the former more like sfe//ata than it is like either of the other two species. The difference in size between immer and adamsii might be accounted for by Bergman’s rule, and like bill form, is probably not primarily related to reproductive isolating mechanisms. The effectiveness of the isolating mechanisms mentioned above is evident in the paucity or lack of known hybrids between the two 1978 SYSTEMATIC NOTES ON THE LOONS 5 species. There is an adult ”female” in the Royal Ontario Museum (No. 76,360) found dead at Pt. Credit, Ontario on December 7, 1957, which has been thought to be a hybrid between immer and adamsii (Godfrey, 1966: 11). In the recurved lower outline of the proximal segment of the mandibular rami (character 6 of Binford and Remsen, 1974: 115) this specimen resembles adamsii. The size of the remaining white squares on the back are within the range of those of adamsii and of males of immer, but not of females of immer. In size of wing, tarsus, and bill, the bird is well within the range of males of immer. In the other bill characters described by Binford and Remsen, the specimen is within the range of immer. While the possibility that this bird is an immer — adamsii hybrid cannot be completely discounted, I think it is more probably a male immer which was mis-sexed by the collector, who recorded nothing concerning the size or condition of the gonads. THE STATUS OF pacifica. — The three forms of the Arctic or Black-throated Loon (arctica) complex (arctica, viridigularis, and pacifica) replace one another around the arctic except in Iceland and Greenland, where none occurs. Thus they might be thought of as forming a broken rassenkreis. The nominate form, arctica, ranges from the British Isles eastward across the northern parts of Eurasia and intergrades with viridigularis in Siberia. The latter form is found in western Siberia, largely south of the range of pacifica, and in parts of western Alaska. The widespread North American form, pacifica, also breeds in northeastern Siberia. Portenko (1939) and Bailey (1948) have shown that the breeding ranges of pacifica and viridigularis overlap in Anadyrland and western Alaska and re- ported no interbreeding. They concluded that the two were, there- fore distinct species, an opinion followed by Vaurie (1965). There are several reasons for doubting the specific status of pacifica. In the first place, those who have discussed the problem in the past have ignored one pertinent question: would arctica and pacifica interbreed if their ranges were to expand and meet? I think it likely that they would because the throat color is the same and the difference in size is less than that between pacifica and viridigularis. Secondly, there has been no thorough, detailed field study of pacifica and viridigularis in the area of overlap, and thirdly, there is evidence of interbreeding between the two forms. Of the two forms, viridigularis is larger in all measurements, has a green (not purple) sheen on the throat, and a darker gray nape. 6 BREVIORA No. 448 Green- or blue-throated individuals, otherwise indistinguishable from typically purple-throated individuals of pacifica are known to occur (Bailey, 1948: 140; Palmer, 1962: 45; Vaurie, 1965: 5), but whether this is a result of introgression with viridigularis or part of the normal range of variation of pacifica is unclear. To date, I have measured and examined 12 examples of viridigularis and approxi- mately 270 of pacifica in breeding plummage. Of the latter, at least nine, or 3.3 percent have green or blue-green throats. All nine have measurements well within the range of pacifica. In addition, a male from Savoonga, St. Lawrence Island, Alaska (Colo. Mus. Nat. Hist. No. 26,769) labelled “viridigularis” has measurements well within the range of pacifica, although all above the means for males of that form. The nape is dark, like that of viridigularis, and it is either a very small example of that form or an intergrade. Two other green- throated birds may be intergrades: a male (Nat. Mus. Canada No. 8,816) from Barter Island, Alaska, has a nape intermediate in color between those of the two forms, a long wing (318 mm. or nearly 1.5 standard deviations above the mean for pacifica), a long tarsus (near the maximum for pacifica), but a small bill (below the mean for males of pacifica); and an unsexed bird (Amer. Mus. Nat. Hist. No. 348,959) from SE Victoria Island, Canada, has a light nape, long wing (321 mm.), a short tarsus (between the means for males and females of pacifica), a long bill (above the mean for males of pacifica) and a deeper bill than that of any pacifica I have measured. Two purple-throated males with long wings (323 and 325 mm.), long tarsi, and bills somewhat above the mean for males of pacifica, are also within the range of viridigu/aris in all these measurements and may be intergrades. While the sample of viridigularis which I have seen to date is too small to permit an accurate analysis of variation within that form, the presence of several probable intergrades suggests that interbreeding occurs between it and pacifica. V. SUBSPECIES PROBLEMS Gavia stellata squamata Portenko. — Vaurie (1965: 4) lists this subspecies as not well differentiated but warranting nomenclatural recognition, while Dement’ev and Gladkov (1968: 291) question its validity. I have examined 12 breeding adults from the range of this form from Franz Josef Land (3), Spitsbergen (7), and Bear Island (2), and I find that the plumage characters used to 1978 SYSTEMATIC NOTES ON THE LOONS 7 differentiate this form from nominate stellata (Vaurie, /oc. cit.) are not consistent within the population, and as Dement’ev and Glad- kov (/oc. cit.) pointed out, are found in some specimens from outside the range. On the evidence now available, I see no value in recognizing this subspecies. Gavia arctica suschkini Zarudny. — Although this race, based on migrant individuals from Russian Turkestan, was recognized by Peters (1931: 34), it is generally considered a synonym of the nominate race by recent authors (e.g. Dement’ev and Gladkov, 1968: 297; Vaurie, 1965: 5). I have not seen the material on which this race was based and follow the above authors in not recognizing suschkini. Gavia immer elasson Bishop. — This subspecies was also recog- nized by Peters (1931: 35), but not by Vaurie (1965: 7) and many other recent authors. This species varies greatly in size, wing lengths of adults ranging at least from 287 to 411 mm., but I have not collected sufficient data to work out the pattern of geographic variation, which appears to be largely clinal. While realizing the possibility that there may be recognizable subspecies within this species, I prefer to consider immer monotypic at least until a thorough revision can be made. LIRERATURE CITED BaiLey, A. M. 1948. Birds of Arctic Alaska. Colorado Mus. Nat. Hist., Popular Ser, 8., 317 pp: BINFORD, L. C., AND J. V. REMSEN, JR. 1974. Identification of the Yellow-billed Loon (Gavia adamsii). Western Birds, 5: 111-126. BRODKOoRB, P. 1963a. Catalogue of fossil birds. Bull. Florida State Museum, 7: 179-293. 1963b. Birds from the Upper Cretaceous of Wyoming. Proc. XIIIth Internat. Ornithol. Congress, pp. 55-70. DEMENTEV, G. P., AND N. A. GLADKOV. 1968. Birds of the Soviet Union, Vol. 2. Israel Program for Scientific Translations, Jerusalem. xi + 553 pp. GopFrREY, W.E. 1966. The birds of Canada. Nat. Mus. Canada Bull., 203. 428 pp. HARRISON, C. J.O. 1976. The wing proportions of the Eocene diver Colymboides anglicus. Bull. Brit. Ornithol. Club, 96: 64-65. Hunter, E. N., ANDR. H. DENNIS. 1972. Hybrid Great Northern Diver X Black- throated Diver in Wester Ross. Scottish Birds, 7: 89-91. PALMER, R. S.. 1962. Handbook of North American birds, Vol. 1. Yale Univ. Press. ix + 567 pp. PETERS, J. L. 1931. Check-list of birds of the world, Vol. 1. Harvard Univ. Press. xvill + 345 pp. 8 BREVIORA No. 448 PorTENKO, L. A. 1939. Inst. Polyar. Zemled. Zhivotn 6, Fauna Anadyr. Kraya, Ptitsy, 2, pp. 144-152. RAND, A. L. 1948. Glaciation, an isolating factor in speciation. Evolution, 2: 314-321. ROBERTSON, I., AND M. FRAKER. 1974. Apparent hybridization between a Com- mon Loon and an Arctic Loon. Canadian Field-Nat., 88: 367. SIBLEY, C. G., AND J. E. AHLQuIST. 1972. A comparative study of the egg white proteins of non-passerine birds. Peabody Mus. Nat. Hist. Bull. 39. vi + 276 pp., 37 figs. STORER, R. W. 1956. The fossil loon, Colymboides minutus. Condor, 58: 413-426. VAURIE, C. 1965. The birds of the Palearctic fauna. Non-Passeriformes. H. F. & G. Witherby. xx + 763 pp. WetTMorRE, A. 1940. Fossil bird remains from Tertiary deposits in the United States. J. Morph., 66: 25-37. tv] Us. COMP. ZOOL: LIBRARY MAR 1 8 1985 NARVARD BREVIO WA Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, MAss. 21 FEBRUARY 1979 NUMBER 449 SOUTH AMERICAN ANOLES: THE SPECIES GROUPS. 2. THE PROBOSCIS ANOLES (ANOLIS LAEVIS GROUP). ERNEST E. WILLIAMS! ABSTRACT. The Anolis laevis group is known from three species represented by only nine specimens, all males, from four localities. All are distinguished by a soft multi-scaled proboscis and form a graded series from Peruvian A. /aevis (proboscis minimally developed) through Brazilian A. phyllorhinus (proboscis of moderate size) to Ecuadorian A. proboscis (proboscis very long). Color in life and habitat are known only for A. phyllorhinus. The proboscis is interpreted as primarily an intraspecific social signal increasing the virtual size of the animals. The most readily recognizable — at least in males — of all South American Anolis are the three species that have a nasal appendage or proboscis. The group may be described as follows: Anolis laevis species group Type species: A. laevis Cope 1876. Definition: (Many possibly useful details are not determinable in A. laevis and hence are not mentioned here.) Alpha anoles of moderate size (60-97 mm snout-vent length) distinguished by a soft multi-scaled nasal appendage projecting forward above the rostral scale. Four to 10 scales across the snout between the second canthals. Supraorbital semicircles in contact or separated by as many as three scales. Loreal rows two to five. Interparietal of moderate to large size in contact with the surpaorbital semicircles or separated by as many as three scales. Supralabials in contact with suboculars. Middorsal scales uniform or the median row raised into a crest of triangular scales. Tail crest single or double. Lamellae IMuseum of Comparative Zoology, Harvard University, Cambridge, Massachu- setts 02138. 2 BREVIORA No. 449 under fourth toes ranging between at least 17 and 26. Dewlap and postanals large in males. Females unknown. Distribution: Widely disjunct: (1) trans-Andean Ecuador; (2) southern tributaries of the Amazon in central Amazonia; (3) western Amazonia. INCLUDED SPECIES Anolis laevis Cope 1876 (Figs 1) Scyvtomycterus laevis Cope, Jour. Acad. Nat. Sci. Phila., NS 8: 165. Holotype. ANSP 11368, collected by Prof. James Orton. Type locality. “Between Moyabamba and Balsa Puerto on the river Huallaga in eastern Peru.” Additional references: Boulenger, 1885: 56 (referred to the genus Anolis and placed between A. tigrinus and A. punctatus); Burt and Burt, 1933: 17 (listing only); Barbour, 1934; 154 (placed along with A. tigrinus as synonym of A. transversalis; both synonymies quite mistaken); Williams, 1965: 6-13 (discussed as member of the punctatus group sensu lato); Peters and Donoso-Barros, 1970: 57 (citation only); Malnate, 1971: 358 (listing of type in the Philadel- phia Academy). Anolis phyllorhinus Myers and Carvalho 1945 (Figs. 2 and 3) Anolis phyllorhinus Myers and Carvalho, Bull. Mus. Nac. NS No. 43: 2. Holotype. MN (Rio de Janeiro) 1804, adult male collected by Alexandre Parko on June 14, 1943. Type locality. “Borba, lower Rio Madeira, State of Amazonas, Brasil.” Additional references: Williams, 1965: 8-13 (discussed as member of the punctatus group sensu lato, first mention of the second known specimen MZUSP 7118, Jacareacanga, Rio Tapajoz, Para, Brasil'); Peters and Donoso Barros, 1970; 63 (citation only). Anolis proboscis Peters and Orces 1956 (Figs. 4, 5, and 6) Anolis proboscis Peters and Orcés, Breviora No. 62: 2. Note that the region between the Madeira and Tapajoz is one of Haffer’s (1974, p. 70) “distribution centers” for endemic Cis-Andean birds. 1979 ANOLIS LAEVIS SPECIES GROUP Anolis laevis, ANSP 11368. Lateral view of head of type. Figure 1. 4 BREVIORA No. 449 Holotype. MCZ (Cambridge) 54300, a mature male collected by Antonio Proano during April 27-29, 1953. Type locality. “Neighborhood of Cunuco, a small town at 1200 meters elevation, five kilometers northwest of Mindo, on the south bank of the Rio Mindo, a northern tributary of the upper Rio Blanco, in Pichincha Province, Ecuador.” (The five additional specimens — USNM 207671-73, in the National Museum of Natural History, Washington, D.C.; IPN 7611, 7612, in the collec- tion of Gustavo Orces V in Quito, Ecuador — are from Mindo or “region of Mindo.”) Additional references: Williams, 1965: 8-13 (discussed as member of the punctatus group sensu lato); Peters, 1967: 13, 17 (key and citation); Peters and Donoso Barros, 1970: 63 (citation only). Ecological information. None except for A. phyllorhinus Myers and Carvalho: “Mr. Parko collected the holotype of Anolis phyl- lorhinus in the capoeira (second growth or low jungle) that sur- rounds the town of Borba. This locality is in the Amazonian lowlands on the lower Rio Madeira, about 90 miles (airline) south- southeast of the city of Manaus. The lizard was caught ona low tree while Mr. Parko was collecting butterflies... .” Distinguishing characters of the species. A. laevis differs from the two other species in the small number of scales between the second canthals (4, rather than 9 or 10), the low number of loreal rows (2, rather than 3 to 5) and in having a very rudimentary proboscis. A. proboscis has a crested dorsum and tail and a very long proboscis, while A. phyllorhinus lacks such crests and in fact has two rows of scales dorsally on the tail and a shorter proboscis. Table | lists the standard scale characters for each species. Size. Snout-vent length: /aevis 60 mm, phyllorhinus 73: mm, proboscis 74 mm. Tail: /aevis 79 mm, phyllorhinus 153 mm, proboscis 97 mm. Hind limb length: /aevis 39 mm, phyllorhinus 49 mm, proboscis 42 mm. Color. Laevis: Cope reported the preserved specimen as follows: “Color above dark gray, below pigmented white (in spirits). The two colors are abruptly defined between the orbit and there are brown spots behind the axilla. Tail distinctly annulate.” Phyllorhinus — Myers and Carvalho: “Color in alcohol (formalin- fixed) plain, dull, bluish gray above, lighter beneath, with no sign of a color pattern anywhere save on the toes and undersides of the legs. 1979 ANOLIS LAEVIS Ges hs Soa SI TA? SPECIES GROUP Uy th “a pad 2 {y ae ae", FL oO ef aametraaed a se g Pap Cvceeeps obereees Bs Lateral view of head of Sao Anolis phyllorhinus, MZUSP 7118. 5 Figure Paulo specimen. 6 BREVIORA No. 449 Table 1. Scale counts and characters. laevis Phyllorhinus proboscis soft protuberance leaf-like leaf-like covered with ? laterally laterally small scales compressed with compressed with granular scales elongate scales N= 1 Nea N= 6 scales between second canthals 4 (Cope) 10 9-10 scales between semicircles 0 0) 1-3 scales between interparietal and semicircles 0 12 1-3 loreal rows 2 4-5 3-5 rows between suboculars and supralabials 0 0 0 labials to center of eye ? 6-7 9-12 sublabials in contact with infralabials ? 5-6 2-3 median gular scales in contact with mental between sublabials e 4 2 middorsal rows enlarged 0 0 l (crest of tri- angular scales) tail crest double row, double row, crest present no crest no crest lamellae 4th toe ? 25-26 =i 1979 ANOLIS LAEVIS SPECIES GROUP Syay tte FS ES SO = a ~~ ~ SS Ww LOS rae SS SSS eceras CIE aS m a = we ae ee wes LA" i, eae aa’ es : \% Cae 10 Cs UTA IY IO Ps he t i RAG iss Oo oo RSSSSSES z2S5: oma Bae Vth un Ge WASSER EE WENSS: oS cen tat A Ree ) Dorsal view of Sao Paulo Anolis phyllorhinus, MZUSP 7118. Figure 3. specimen, 8 BREVIORA No. 449 The undersurfaces of the legs and arms bear a fine, light mottling, perhaps better described as a bluish reticulation on a yellowish ground color. The undersurfaces of the digital expansions are dark. The superior surfaces of the toes and fingers (but not of the feet and hands) are light, with bold, blackish crossbands. The tail gradually becomes brownish toward the end. A color sketch made by Mr. Parko from life shows the creature bright blue-green, the dewlap yellowish, with traces of pink or red on the toes, the top of the snout, and the end of the rostral appendage.” Mr. Parko’s notes say the beast immediately changed from bright blue-green to brown when captured, but changed back to green on preservation. Proboscis — Peters and Orces: “Dorsal ground color (in alcohol) a dull gunmetal blue, with irregular, parallel, horizontal black streaks along vertebral margin. These streaks vaguely line up and form a pair of lateral bands between the limbs. Dark brownish- black spot at shoulder. Limbs and lateral surfaces spotted with light yellowish white; limbs vaguely barred with darker blue; skin in interstices between scales of dorsum of digits light, giving appear- ance of banding on fingers and toes. Dorsum of head unicolor, as back; temporal region and lips lighter, with faint reddish or purplish tinge and a marked light spot over ear opening. Ventral surfaces generally lighter, with vague reddish tints on chest and chin; belly stippled with light spots. Venter of limbs heavily spotted with white proximally, becoming totally white on the foot. Tail alternately barred with gunmetal blue and blackish bands approximately equal in width.” TAXONOMY AND NOMENCLATURE The single recognition character — the proboscis — differs considerably in the three known species, which are abundantly distinct. The extent of the differences might reasonably raise the issue of parallel evolution, but the proboscis is so singular a feature that it seems subjectively more probable that the known species are the relicts of a once much larger radiation. It is also very possible that there are more species still to be discovered, perhaps with quite restricted distributions. It is clear that the first adumbrations of a proboscis are found in A. punctatus and some of its relatives. In these the rostral scale and rostral area are produced to overhang the lower jaw. In /aevis there 1979 ANOLIS LAEVIS SPECIES GROUP LE AL tr Zs LE LLL a i. Lin CJ Oe: ee oe SOI Zl sae) roe) TS sine Canes, WSS aa oY EST 2 eee Ot ice: is oa ees = > aa See SS ai ) : iC \ Av ~ My SAD Ef ae Aen eens BP) 5 Nga ery SES Oiaeteas: Sane! ne Besepeste: beecteneregriyy iy i sasatteas a STR SRS SRN Sansa eats Roba ae, aetesay SERS (TRESS RII > YY Anolis proboscis, USNM 207672. Lateral view of head. Figure 4. 10 BREVIORA No. 449 appears to be a soft protuberance which involves the smaller post- rostral scales. In A. phyllorhinus and A. proboscis the rostral remains at the same level of development as in /aevis but the area above the rostral is expanded into an impressive nose leaf, longest in A. proboscis. The distinction between the proboscis and punctatus group 1s thus perhaps somewhat artificial. The morphological sequence is, however, clear and leads from proboscis at one extreme back to punctatus at the other. I find it convenient to make the distinction at the point at which the scales and tissues above the rostral are involved in the swelling. Since the relation of this area to the rostral is very similar in the three species, it provides greater plausibility for the contention that this is a single lineage. It is not, on the other hand, clear that A. punctatus is directly related to the species that mimic it by means of the presence of a projecting rostral in the male. However, any decision on this point is necessarily tentative, and I emphasize convenience rather than certainty of phyletic affinity as a basis for recognition of this group. Nothing is known of the function of the proboscis (see below) and all nine specimens of this group thus far collected have been males. This has raised (e.g. Peters and Orces, 1956) the issue that the proboscis anoles might be the males of species already described on the basis of females. However, there are few described species still known only from females. Also the other external characters of each of the proboscis anoles are such that it would require a degree of sexual dimorphism beyond anything known in lizards to make any of the three proboscis species possibly conspecific with any other described anoles. In the case of A. punctatus and the other species with projecting rostrals, this and the presence of a dewlap, enlarged postanal scales and sometimes minor differences in color distinguish the males. Scale counts and other aspects of morphology do not significantly differ between the sexes. Only A. proboscis is known from series (the type, three newly collected specimens given to the United States National Museum by Orcés, and two inthe Orcés collection in Quito). A. phyllorhinus is represented by only two specimens, the type and a badly preserved second specimen. A. /aevis is still known only from the type, which is noW in extremely poor condition. ANOLIS LAEVIS SPECIES GROUP 11 1979 eg nnn oeaeceeee “pray JO MOIA [BSIOG “ZL9LOT WNSN ‘Slosoqodd syoup ~¢ ainsi BM—e' SADIE Shs LES Oe Saere a a 12 BREVIORA No. 449 EVOLUTIONARY ASPECTS Three species, nine specimens, four localities and one note on habitat are the sum of our objective data on the proboscis anoles. They provide little opening for discussion or for evolutionary speculation. It is possible, however, to place this minimal amount of data against a background of the much greater knowledge of anoles as a whole. It is a peculiarity of the proboscis anoles that in no other way are they known to be different or indeed especially interesting. The single aspect that appears striking is the proboscis itself. Other bizarre structures are known in anoles. The dewlap would be bizarre were it less common and less familiar. High tail fins supported by heightened caudal neural spines occur in a number of distantly related species. Casque heads — heads enlarged by crests and ridges — occur in several Anolis and in the related anole genera Phenacosaurus and Chamaeleolis. All these structures — dewlap, tail fin, casque head and proboscis — have the effect of increasing the apparent size of the animals that bear them. In the case of the dewlap, there is evidence of use in agonistic encounters, and evidence (Rand 1967) that larger animals usually win in such encounters. Trivers (1976) has shown the importance of sexual selection for size in one species of Anolis; he has inferred the generality of the rule. There are, however, aspects of increase in real size that make the course of evolution complex. A real increase in size with its implied increase in weight can carry with it functional restrictions that compel changes in the life style of the species that exercises this option. Sometimes such changes are realized within the ontogeny of a single species. Sometimes we see them as the alternative strategies utilized by sympatric species (Schoener, 1970; Williams, 1972). Virtual changes in size — unreal but visually effective — are still another alternative. Virtual size has the advantage of introducing few if any functional problems. I have called attention to elongation of the head in many crown anoles (Williams, 1965). It is probable that this initially has a (still undefined) functional explanation. However, there are side effects: the change of head shape probably has its species-recognition aspects and the size of the head has its agonistic advantage aspects. Yet if these latter aspects have too much effect, the bony structure of loa) ROUP J ANOLIS LAEVIS SPECIES (¢ 1979 “M ol A [Rl 91e" ‘CLOLOT W NS { } S1ISOGO id syouy ‘g ound 14 BREVIORA No. 449 the head is elongated beyond its functional optimum. There will therefore again be a premium on virtual rather than real change of head shape. We see the beginning of such a phenomenon in Anolis punctatus and in some related species where (in males) only the rostral scale is enlarged. A. /aevis carries this process a slight step further. A. phyllorhinus and A. proboscis have gone much further. It is significant that the probosces in A. phyllorhinus and A. proboscis are not similar in detail. One would expect this if there were functional aspects to the structure per se. On the contrary, it appears a similar visual effect may readily be achieved by only broadly comparable means. This corresponds with species recog- nition devices and intra-species display where there is no require- ment that structures be similar; rather the requirement is that they be different. (In view of the wide allopatry of the known proboscis anoles, there could be no selection for striking difference now, but perhaps it has existed in the past.) Within the Iguanidae, the basiliscines would seem to be quite parallel cases. Head casques — very different in structure — are present in all three genera; in Basiliscus, dorsal and tail fins are present, different and wonderfully conspicuous in three species, very reduced in a fourth. (Once a bizarre structure has been adopted as an intra- and interspecies signal, an option open to a new member of a group is to abandon the structure: absence of a signal is itself a signal.) The absence of a dewlap is a strategy adopted by two species in the very complex anole fauna of Cuba; the dewlap is consistently retained in the simplest faunas and is reduced only in the fauna almost as locally complex as Cuba, that of Hispaniola (Williams and Rand, 1977). Outside the Iguanidae, a number of species in the closely related family Agamidae show parallels to the proboscis anoles: Cerato- phora (Ceylon, three species, see M. Smith, 1935 and Taylor, 1953); Harpesaurus (Java, Sumatra, Nias, five species fide Wermuth, 1967); Lyriocephalus (Ceylon, one species, Taylor, 1953) and Co- photis (Ceylon and Sumatra, De Roois, 1915). The probosces in these are all soft structures but considerably more variable in shape, number and squamation than those of the proboscis anoles. (Figs. 7-9 show the differences within the genus Ceratophora.) For cha- meleons Rand (1961) has already discussed the function of the ornamentation that is so remarkably developed in the montane 1979 ANOLIS LAEVIS SPECIES GROUP ee SSA Q\\ . Ne : SN Bil Pe SN = —- a8: fake SS ee os a. = aZ6 Za, SS os (EX SANS! Ret \ tf eats “a (\h y a AL | be RNG EEE LI Pity) A\veseanea: C ERO SRY | RO As pa Ra m4 “6 e te a aki Ae as 7. Ceratophora tennenti, MCZ 136187. Side view of head. Figure 16 BREVIORA No. 449 species of Africa. In the horned chameleons, however, the ornament is a solid structure. In this case, the ornament may serve a physical function in aggression, as the horns do. Still further afield, Hopson (1975) has discussed the singular crests of hadrosaurian dinosaurs as cranial display structures. His paper (see also Molnar, 1977) also cites the parallels with mam- malian horns and has references to the extensive literature. Of course, alternative explanations are available for structures such as probosces in Anolis, e.g. camouflage. However, such other functions, if they exist, do not disprove the display function of Anolis probosces or basiliscine crests. The value of a structure for more than one function is a familiar phenomenon and will, not surprisingly, be sustained by natural selection. ACKNOWLEDGMENTS The extant world material of proboscis anoles has been made available to me by the late James A. Peters, W. R. Heyer and Ronald Crombie of the National Museum of Natural History, by Gustavo Orces-V of the Escuela Polytecnica Nacional, Quito, Ecuador, by Edmond V. Malnate of the Academy of Natural Sciences, Philadelphia and by P. E. Vanzolini of the Museum of Zoology of the University of Sao Paulo. To all of them I am indebted for their help and kindness. | have discussed the function of the probosces with A. S. Rand. Robert Bleiweiss called my attention to the refugium between the Rios Tapajoz and Madeira cited by Haffer. Laszlo Meszoly has provided the illustrations of the proboscis anoles and of their counterparts in the agamid genus Ceratophora. This study has been supported in part by National Science Foundation grant GB 37731X. LITERATURE CITED BARBOUR, T. 1934. The Anoles II. The mainland species from Mexico southward. Bull. Mus. Comp. Zool. 77: 119-155. BOULENGER, G. A. 1885. Catalogue of the lizards in the British Museum (Natural History), 2nd ed., London, xin + 492 pp. Burt, C. E. AND M. D. Burt. 1933. A preliminary checklist of the lizards of South America. Trans. Acad. Sci. St. Louis 28: 1-104. Corr, E.D. 1876. Report on the reptiles brought by Prof. James Orton from the Middle and Upper Amazon and western Peru. Jour. Acad. Nat. Sci. Phila. (2) 8: 159-188. 1979 ANOLIS LAEVIS SPECIES GROUP sesteste —— & Saree Peo et) Loe 30 ww, Se - pee Se . MCZ 4139. Side view of head. aspera ophora Figure 8. Cerat 18 BREVIORA No. 449 De Roo, N. 1915. The reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. Brill, Leiden: i-xiv + 384 pp. HaAFFER, J. 1974. Avian speciation in tropical South America. Publ. Nuttall Ornith. Club No. 14, Cambridge, England, vill +389 pp. Hopson, J. A. 1975. The evolution of cranial display structures in hadrosaurian dinosaurs. Paleobiol. 1: 21-43. MALNATE,E. V. 1971. A catalog of primary types in the herpetological collections of the Academy of Natural Sciences, Philadelphia (ANSP). Proc. Acad. Nat. Sci. Phila. 123: 345-375. MOLNAR, R. E. 1977. Analogies in the evolution of combat and display structures in ornithopods and ungulates. Evol. Theory 3: 165—190. Myers, G. S. AND A. CARVALHO. 1945. A strange new leaf-nosed lizard of the genus Anolis from Amazonia. Bol. Mus. Nac. Rio de Janeiro N.S. 43: 1-14. Peters, J. A. 1967. The lizards of Ecuador, a checklist and key. Proc. U.S. Nat. Mus. 119: 1-49. Peters, J. A. AND R. DoNnoso-BArRRos. 1970. Catalogue of the Neotropical Squamata. Part II. Lizards and amphisbaenians. Bull. U.S. Nat. Mus. 297: viii + 293 pp. Peters, J. A. AND G. OrcEs-V. 1956. A third leaf-nosed species of the lizard genus Anolis from South America. Breviora No. 62: 1-8. RAND, A. S. 1961. A suggested function of the ornamentation of East African chameleons. Copeia 1961: 411-414. 1967. Ecology and social organization in the iguanid lizard, Anolis lineatopus. Proc. U. S. Nat. Mus. 122: 1-79. SCHOENER, T. W. 1970. Size patterns in West Indian Anolis lizards. Il. Correla- tions with the sizes of particular sympatric species — displacement and convergence. Amer. Natur. 104: 99-103. SmitH, M. 1935. The Fauna of British India, including Ceylon and Burma Reptilia and Amphibia. II. Sauria. Taylor and Francis, London: 1-xiii + 440 pp. Taytor, E. D. 1953. A review of the lizards of Ceylon. Univ. Kans. Sci. Bull. 35: 1525-1585. Trivers, R. L. 1976. Sexual selection and resource-accruing abilities in Anolis garmani. Evol. 30: 253-269. WeRMUTH, H. 1967. Liste der rezenten Amphibien und Reptilien. Agamidae. Das Tierreich 86: 1-xiv + 127 pp. WILLIAMS, E. E. 1965. South American anoles (Sauria, Iguanidae): two new species of the punctatus group. Breviora No. 235: 1-15. 1972. The origin of faunas. Evolution of lizard congeners in a complex island fauna: a trial analysis. Evol. Biol. 6: 47-90. WILLIAMS, E. E. AND A. S. RAND. 1977. Species recognition, dewlap function, and faunal size. Amer. Zool. 17: 261-270. ANOLIS LAEVIS SPECIES GROUP 19 1979 “SLE9IL ZOW ‘Véapppols psoydolpsd) ‘6 ons 183 on ad ten “ae 2 An Pie a FN : oe : \ : : \ MUS. COMP, ZOOL LIBRARY MAR 1 8 1985 BREVIO*RA Museum of Comparative Zoology ] BY US ISSN 0006-9698 CAMBRIDGE, MASS. 21 FEBRUARY 1979 NUMBER 450 A NEW SPECIES OF MOENKHAUSIA FROM THE MATO GROSSO REGION OF BRAZIL (PISCES: CHARACIDAE) WILLIAM L. FINK! ABSTRACT. A new species of the characid fish genus Moenkhausia is described from the Arinos River of the Mato Grosso region of Brazil. A brief discussion of some of the characters used in analyses of characid relationships is included. INTRODUCTION The fish described here as new presents some interesting problems regarding current classification criteria for characids. These prob- lems arise primarily because the system now in use, which dates from the work of Eigenmann (1917), is obsolescent and no longer able to incorporate the diversity of the fishes it was intended to classify. Most of the characters which are used to “define” genera and higher categories have been found to be inadequate to express hypotheses about relationships within the family. The bases of the subfamilies Cheirodontinae and Tetragonopterinae have been re- cently reviewed by several authors (see Fink and Weitzman, 1974) and this critical reappraisal is being carried on in other areas within the Characidae. The characters which cause problems in hypothesizing the rela- tions of the fish herein described are lateral line length and color pattern. The former character is of major importance in the “Eigenmann” system while the latter has been virtually ignored by characid systematists. Both of these characters are discussed below after the description. 'Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138. W BREVIORA No. 450 The methods of taking morphometric and meristic data used in this analysis are described in Fink and Weitzman (1974). Propor- tions are given as percent of standard length (SL). Paratypes have been deposited in the following institutions: Academy of Natural Sciences of Philadelphia (ANSP), British Museum of Natural History (BMNH), California Academy of Sciences, San Francisco (CAS), National Museum of Natural History, Washington, D.C. (USNM), Museum of Comparative Zoology, Cambridge, Mass. (MCZ), Museu de Zoologia, Universidade de Sao Paulo(MZUSP), and the Zoologisch Museum, Universiteit van Amsterdam (ZMA). Moenkhausia phaeonota, new species Figures eZ Holotype; MZUSP 13793; Brazil, Mato Grosso, from the shores of an island “below Rio dos Peixes,” Rio Arinos, Jurena-Tapajos drainage; collected by Harald Schultz, l6" June 1962: 32-6 mm: SL Paratypes: ANSP 135924, same data as holotype, one specimen 26.7 mim: SE: BMNH 1978.9.12.1, same data as holotype, one speci- men 28.8 mm SL. CAS 41707, same data as holotype, one specimen 29.] mim S_L: FMNH 83856, same data as holotype, one specimen 29.0 mmol: MCZ 54087, same data as holotype, three specimens 2/.5-=29.9 mm SL: MZUSP. 13794, same data as holotype, one specimen 195 mm SL. MZUSP. 13795, same data as holotype, one specimen 20.6 mm SL. USNM 218798, same data as holotype, twenty speci- mens 16.1-36.4 mm SL. ZMA 115.273, same data as holotype, one specimen 24.) “nom Se: Description See Tables | and 2 for variable meristic and morphometric data. Body elongate, compressed laterally. Predorsal body profile slightly NEW SPECIES OF MOENKHAUSIA 1979 “TS WW PTE “862817 WNSI | od Ayeied DJIOUO avyd DISNDY YuI0 PW ‘| ainsi 4 BREVIORA No. 450 convex with a slight concavity at nape. Dorsal body profile abruptly angles ventrally along dorsal-fin base, then becomes straight or slightly convex to adipose fin where profile dips ventrally a small amount, then continues as a nearly straight line to dorsal procurrent caudal-fin rays. Ventral profile gently rounded to anus; steepest inclination ventral to jaws. Ventral profile protrudes ventrally its greatest distance just anterior to pelvic-fin insettions. Body profile along anal fin-base nearly straight, or slightly convex; at anal-fin termination, profile nearly straight or slightly concave to procurrent caudal-fin rays. Maxillary with 3-7 teeth; in specimens below 21-25 mm SL, teeth mostly conical; specimens in that size range usually with dorsalmost tooth tricuspid (rarely with 4 cusps); specimens above that range usually with tricuspid maxillary teeth. Premaxillary outer row teeth, 2-5, larger fish usually with larger tooth number; in smaller specimens teeth usually conical; in speci- mens above about 24 mm SL, teeth usually tricuspid. Premaxillary inner row teeth, 5-6, each tooth usually smaller than tooth medial to it. Ontogenetic changes include replacement of earlier teeth by teeth with a greater number of cusps until adult condition reached. Smaller specimens (about 16-19 mm SL) usually have medial three teeth with 3 cusps, with lateral teeth having | or 2 cusps, giving a tooth cusp formula of 33311 or 33321. In the size range 22-25 mm SL there is a variable pattern: medial tooth 3-4 cusped, second tooth 3-5. cusped, third tooth 3-5 cusped, fourth tooth with 3 cusps, teeth lateral to those with |-2 cusps. In adults, inner row premaxillary tooth formula usually 45533 or 45433. Dentary teeth show ontogenetic trends similar to those exhibited by inner row premaxillary teeth, replacement teeth usually with more cusps. Small specimens usually have a formula 3331 +(6—-10 very small conical teeth); adults usually have tooth formulae ranging from 4333 +(6—-10) to 5353 +(6—-10); second tooth from symphysis is offset anteriorly in adults and rarely has more than 3 cusps. Fontanels moderate, that pari anterior to the epiphysedl bar slightly more than two-thirds as long as that part posterior to bar. Gill rakers moderate. Infraorbital bones well ossified, infraorbital 3 wide, with a narrow naked area behind and below. 1979 NEW SPECIES OF MOENKHAUSIA 5 Figure 2. Dentition of Moenkhausia phaeonota, paratype, USNM 218798, 28.2 mm SL. 6 BREVIORA No. 450 Scales moderately large, thin, cycloid with concentric circuli and about 2-7 radii on the exposed posterior field. Lateral line with a slight ventral curve along body, complete, with scales perforated to end of caudal base. Scales above lateral line 5; scales below lateral line 4. Scale sheath of about 4-7 scales along anterior anal-fin base. Scales extend over about 1/5 to 1/4 of caudal-fin rays. Small accessory scales present ventral to pelvic-fin insertion. Dorsal-fin origin anterior to anal-fin origin, posterior to pelvic fin insertion, nearer eye than caudal-fin base. Third or fourth ray of dorsal fin longest with posterior rays shorter, forming a straight or slightly convex posterior margin to fin. First unbranched ray of anal fin usually not visible externally; last unbranched anal-fin ray and first through fourth or fifth branched rays longer than posterior rays, forming a slightly concave fin margin. Pectoral fin low on body, pointed or slightly rounded, reaching to or beyond pelvic-fin insertion. Pelvic fin with distal tip reaching from just anterior to just posterior to anal-fin origin; 1 7 rays. Caudal fin with 10/9 principal caudal-fin rays; fin forked, not split to base. Dorsal and ventral caudal-fin lobes equal in size. Dorsal and ventral procurrent caudal- fin rays are equal in size and extend cephalad same distance. All fin rays without hooks. Color in alcohol. — Ground color light brown. Nape, top of head, and jaws covered with numerous small brown melanophores. Back with a reticulate pattern of dark brown melanophores. A broad stripe almost devoid of melanophores extends from just above and behind eye along side of body, terminating ventral to dorsal procurrent caudal-fin rays. Below pale stripe is a broad dark stripe consisting of numerous melanophores extending from cheek and opercle behind eye, across body side and onto caudal peduncle and posteriorly onto about six of middle caudal-fin rays. Immedi- ately posterior to opercle on second through fourth lateral scales, melanophores are extremely dense, forming a large humeral spot about two-thirds diameter of eye. Posteriorly, above anal fin and on caudal peduncle, melanophores very dense and tend to concentrate along junctions of myomeres. Immediately dorsal to anal fin, melanophores abruptly become less dense, leaving a pale stripe at anal-fin base which is continuous with pale color of belly. Dorsal fin with melanophores concentrated on interradial membrane between first and fifth branched rays. Pectoral and pelvic fins with a few 1979 NEW SPECIES OF MOENKHAUSIA 7 Table 1. Measurements of Moenkhausia phaeonota Character N Range x S Holotype Standard Length (SL) 31 16.1-36.4 25.8 5.670 32.6 % SL 4 Greatest Depth 29 25.6-35.5 30.2 2.302 29.1 Eye-Dorsal Fin Origin 31 34.7-38.2 36.7 0.890 36.2 Dorsal-Fin Origin— 31 50.5-56.2 53.3 1.307 52.5 Base of Caudal Peduncle Snout-Dorsal Fin Origin 31 50.3-53.4 SEG 0.737 50.6 Snout-Pectoral Fin Origin 31 24.7-29.8 26.4 1.017 DES Snout-Pelvic Fin Origin 31 42.4-47.2 45.3 1.164 44.8 Snout-Anal Fin Origin 3] 55.7-60.7 58.8 Ie ley7/ 59.2 Dorsal Fin Length 25 26.0-31.7 28.1 1.461 30.7 Pectoral Fin Length 30 18.5-24.6 DRS 1.448 19.3 Pelvic Fin Length 3] 10.8-16.4 14.6 TED 14.1 Head Length 31 19.4-28.6 252 2.141 24.5 Eye Diameter 31 9.6-13.8 Whites} 1.038 10.1 Snout Length 31 6.2— 7.8 6.9 0.384 6.4 Bony Interorbital 31 7.5— 9.0 8.3 0.354 8.0 Caudal Peduncle Depth 31] 8.3-10.7 9.5 0.659 9.8 Caudal Peduncle Length 31 10.4-14.3 12.6 1.011 IES, 8 BREVIORA No. 450 scattered melanophores on interradial membranes. Caudal fin with melanophores along edges of all rays, in addition to the extremely dark middle caudal rays mentioned above. Anal fin with melano- phores loosely scattered on interradial membranes, more concen- trated between anterior rays and especially along outer margin of fin. Adipose fin with many small melanophores. Etymology. — From phaios, meaning dusky brown and the suffix — nota, meaning having the attribute or quality of. The name refers to the color pattern of the fish. DISCUSSION This species, according to Eigenmann (1917), belongs in the genus Moenkhausia, which is diagnosed by the following combination of characters: complete lateral line, partially scaled caudal fin, five inner row premaxillary teeth and a naked area between the third infraorbital and the lower limb of the preopercle. However, in color pattern, preserved specimens are extremely similar to several species of Hyphessobrycon. A large number of Moenkhausia species were examined durng this study in an attempt to find a potential sister group for M. phaeonota (see Comparative Material Examined) but no currently recognized species was considered to have any derived characters in common with it. In Eigenmann’s (1917) key to the species of Moenkhausia, those that are included in the couplet “depth usually more than 2.75 in length” are most similar to M. Phaeonota in general body shape but they have marked color differences and in most cases meristic count differences. Specimens identified as M. dichroura (Kner) by Eigenmann (USNM 86792) approach M. phaeonota in both anal-fin ray number and lateral scale number, but are very different in color pattern, in being heavier bodied, and in having a larger third infraorbital. Of the more recently described species, M. takesi Gery is phenetically more similar to M. phaeonota than other members of the genus. Both are fairly similar in body and head shape, although M. takesi is “stouter.” Moenkhausia phaeonota has more predorsal scales (11-13 versus 9), more lateral scales (36-38, usually 36, versus 33), more anal-fin rays (ili, 22-25, usually 22 or 23, versus iv, 18-19) [The holotype of M. takesi has four unbranched anal-fin rays rather than three as reported by Gery (1964)], and fewer gill rakers (17-21, usually 18, versus 20-21). The color pattern of M. takesi in life and 1979 NEW SPECIES OF MOENKHAUSIA 9 Table 2. Meristic counts of Moenkhausia phaeonota Character N Range X S Holotype Branched Dorsal 8 “o Fin Rays 31 229 8.9 0.250 9 Branched Anal Mo ai aa OS Fin Rays 31 Wil 3 3} 23¥ 0.854 23 Branched Pectoral 10) WI Fin Rays 31 H 9 20 | 10.7 0.599 1] Branched Pelvic a7 Fin Rays 31 31 7.0 0.000 7 Lateral Sei) sb “sje Sxoy Sii/ Scales 11 2 ] 2 | 4 BS 1.635 Bi Perforated 34535) 36) 3) 38 Lateral Scales 9 2 | 2 3 l 36.0 1.414 37 Scales Around 12 13 14 Caudal Peduncle 8 l | 6 13.6 0.744 14 Predorsal LOR 23s Scales 26 eS ne 4 ee) HIE? 0.710 13 Gill ty dhe IO) AO pei Rakers 13 2 ao) l l 18.2 0.987 17 Precaudal 13 14 Vertebrae 3] [20 13.6 0.486 14 Caudal 23 24 25 Vertebrae 31 18 12 l 23.5 0.568 23 Total 37 38 Vertebrae 31 28 3 Bil 0.301 37 10 BREVIORA No. 450 in preserved specimens consists primarily of dense black melano- phores dorsal to a light lateral stripe and includes a dark area on the caudal peduncle and the area dorsal to the anal fin. The black band in M. phaeonota extends from the middle caudal-fin rays onto the base of the tail and continues to the posterior border of the opercle. In summary, M. phaeonota seems to share no unique characters with any known species of Moenkhausia and is rather different phenetically from the only Moenkhausia species which looks even slightly similar (M. takesi). Inasmuch as there is questionable phylogenetic reality to Moenkhausia, it is useful to look beyond the bounds of that genus to another group with which M. phaeonota seems to share a number of similarities. This leads us to a group of species within Hyphessobrycon, the members of which share a specialized color pattern. This group includes H. metae Eigenmann and Henn, H. loretoensis Ladiges, H. peruvianus Ladiges, and perhaps H. agulha Fowler and H. herbertaxelrodi Gery. All of these species have a color pattern in preserved specimens which includes a band of dark brown or black chromatophores which generally extends along the lateral body from the eye to the caudal fin base. The area of greatest concentration of chromatophores is usually dorsal to the anal-fin base, and there is often a secondary concentra- tion in the “humeral” area behind the opercular aperture. In life, the lateral band is dark brown to black, and dorsal to it is a bright lateral stripe, usually gold, goldish-red or creamy white. In addition, in live specimens there is generally some red coloration on the caudal and dorsal fins, and in some species, dorsally on the eye. Preserved specimens of Moenkhausia phaeonota have a color pattern nearly indistinguishable from H. metae of Colombia, and differ from that species primarily in meristic counts including: number of lateral scales (33-37, compared with about 33 in metae), number of branched anal-fin rays (22-25, compared with 17-19 in metae), and number of vertebrae (37-38, compared with 33 in metae). In view of this striking resemblance to these members of Hyphes- sobrycon, | would propose a relationship of M. phaeonota with those species. That group is currently being revised by myself and S. H. Weitzman and M. phaeonota will be considered in that study, after material available to us has been further examined and analyzed. In the interim, the species described herein will be referred 1979 NEW SPECIES OF MOENKHAUSIA 11 to the genus Moenkhausia, where it belongs according to the diagnostic characters of that series. A revision of this group of characids must include a redefinition of certain generic limits and a full review of these fishes, something which I am not yet prepared to present. COMPARATIVE MATERIAL EXAMINED Hyphessobrycon agulha Fowler, holotype, ANSP 39232, Brazil, Rio Madiera, about 200 miles E of Long W62° 20’. Coll. E. A. Smith, 1912. Hyphessobrycon herbertaxelrodi Gery, holotype, USNM 196089, Brazil, Mato Grosso, Rio Paraguay Basin, Coxim on the Rio Tacquary. Coll. K. H. Stegemann, 1960. Hyphessobrycon loretoensis Ladiges, MCZ 52154, Colombia, Rio Loreto Yacu, in small creeks about | km from river mouth (Rio Loreto Yacu is a trib. of Rio Maranon), approx. 03°30’S, 070° 10’W. Coll. Alfonso Doaz, Dec. 1976. Hyphessobrycon metae Eigenmann and Henn, holotype, CAS (IU) 13421, Colombia, Barizora, Rio Meta, Coll. M. Gonzales. Hyphessobrycon metae Eigenmann and Henn, paratypes, CAS (IU) 13422, same data as holotype. Moenkhausia browni Eigenmann, holotype, FMNH 52733, British Guiana: Arnataima. Coll. C. H. Eigenmann, 1908. Moenkhausia collettii Steindachner. USNM 66228, British Gui- ana: Erukin Creek, lower Potaro River. Coll. C. H. Eigen- mann, 1908. Identified by Eigenmann. Moenkhausia copei (Steindachner), USNM 66230, British Gui- ana: Rockstone, Essequibo River. Coll. C. H. Eigenmann, 1908. Identified by Eigenmann. Moenkhausia cotinho Eigenmann, USNM 66244, British Guiana: Tukeit, lower Potaro River. Coll. C. H. Eigenmann, 1908. Moenkhausia dichroura (Kner), USNM 86792, Bolivia: Rurrena- baque-Coll. N.. E. Pearson, Oct. 1921. Moenkhausia eigenmanni Gery, holotype, USNM 198640. Colom- bia: Rio. Marracacias into upper Rio Meta, at Restrepo. Coll. Ross Socolof, June, 1963. Moenkhausia lepidura gracilima Eigenmann, holotype, USNM 120279, Brazil: Serpa. Coll. Thayer Expd. 1866. 12 BREVIORA No. 450 Moenkhausia takesi Gery, holotype, USNM 198136, Brazil: Rio Guana basin, near Belem do Para. Coll. Takese, 1963. Moenkhausia robertsi Gery, holotype, USNM 200427, Peru: upper Amazon, near Iquitos. Coll. J. Roberts, 1963. ACKNOWLEDGEMENTS I wish to thank Sara Fink and Stanley H. Weitzman for editorial comments on the manuscript. James E. Bohlke and W. E. Esch- meyer generously loaned specimens of Hyvphessobrycon metae. LITERATURE CITED EIGENMANN, C. H. 1917, The American Characidae, 1. Mem. Mus. Comp. Zool., Harvard 43(1): 102 pp. FINK, W. L. AND S. H. WEITZMAN. 1974. The so-called cheirodontin characids of Central America, with descriptions of two new species (Pisces: Characidae). Smithsonian Contrib. Zool. 172: 46 pp. Gery, J. 1964. Two new tetras from the lower Amazon basin. Trop. Fish Hobb. 12(7): 13-15 and 59-69. MUS. fo BRE V1 O-R°A MAR 1 8 1985 Museum of Comparative v4Zeplogy UNIVER@ITY US ISSN 0006—9698 CAMBRIDGE, MASS. 21 FEBRUARY 1979 NUMBER 451 A NEW SPECIES OF CYBOTOID ANOLE (SAURIA, IGUANIDAE) FROM HISPANIOLA ALBERT SCHWARTZ! ABSTRACT. Anolis strahmi, a new species of clivicolous cybotoid anole from Hispaniola, occurs both north and south of the Sierra de Baoruco, the mountain range associated with the Dominican Peninsula de Barahona. Two subspecies are recognized. Additionally, a new subspecies of Anolis longitibialis Noble is named from the Peninsula; nominate 4. /. /ongitibialis occurs on Isla Beata off the southern tip of the peninsula. Details of distribution and known ecology of the two species are given, and comparisons of the two species (and their respective subspecies) are made. A hypothetical evolutionary and geographical sequence relating these two species to parent A. cybotes has been postulated. INTRODUCTION Anolis cybotes Cope is the first-named of a series of related taxa of Hispaniolan anoles. A. cybotes, sensu stricto, 1s a widespread mesophilic species of moderate size that occurs in wooded to rather Open situations throughout much of Hispaniola. Although the species prefers mesic habitats, it does not completely shun situations which are xeric. In extremely arid areas, the species is often confined to oases or other shaded enclaves. It reaches elevations in excess of about 1,525 m, depending upon which named upland populations one accepts as subspecies of A. cybotes (see Schwartz and Thomas, 1975:77). The species is also known from several of the Hispaniolan satellite islands: Ile de la Gonave, Isla Catalina, Isla Saona, Ile-a- Vache, Ile de la Tortue, and Ile Grande Cayemite. Only on the first of these has an endemic subspecies been named. The present paper 'Miami-Dade Community College North, Miami, Fla. 33167. 2 BREVIORA No. 451 does not attempt to deal with those named populations (armouri Cochran, doris Barbour, haetianus Garman) that are or have been associated with A. cybotes nomenclaturally. Three other species form the cybotes complex. A. shrevei (Coch- ran, 1939) is restricted to the uplands of the Dominican Cordillera Central; this has been recently affirmed as a distinct species by Williams (1975). A xerophilic ally of A. cybotes is A. whitemani Williams. This species centers in the Cul de Sac-Valle de Neiba plain, which today encompasses the area between the north and south paleoislands (Williams, 1961), east into the Llanos de Azua. A. whitemani and A. cybotes occur macrosympatrically in this region; the former occupies xeric situations whereas the latter occupies oases and riverine woods — shaded, mesic situations. There are also records of A. whitemani from the northwestern Haitian Presqwile du Nord Ouest (Mole St. Nicholas) and near Gonaives. It is likely that these “isolated records” are one end of a linear coastal range from the Cul de Sac Plain along the Golfe de la Gonave onto the xeric northwestern peninsula, but records are absent to affirm this continuity. A. whitemani is also known from the xeric Monte Cristi region in northwestern Republica Domini- cana. Most recently, Williams (1975) named A. marcanoi from the southern xeric slopes and associated plains of the Sierra de Ocoa, a southern affiliate of the Cordillera Central. A. marcanoi is sym- patric (and even rarely syntopic) with A. cybotes. Williams has given very pertinent details of this association, and Webster (1975) affirmed species-rank for A. marcanoi by electrophoresis. These two species differ slightly meristically, but in living animals the dewlap color and the head and body pattern are distinctive and dichoto- mous. Anolis longitibialis was described from Isla Beata, off the south- ern tip of the Peninsula de Barahona. Cochran (1934) first combined A. longitibialis with A. cybotes; the species are indeed very similar in general appearance and scutellation. It would seem quite logical that wide-ranging A. cybores would have a local insular population on Isla Beata, despite the absence of A. cybotes from the Peninsula de Barahona itself, at least as far as Cochran’s data (1941) suggested. In 1969, Richard Thomas collected a single large female cybotoid on the Peninsula de Barahona. This lizard was so much larger than 1979 NEW SPECIES OF CYBOTOID ANOLE 3 female A. cybotes from the adjacent foothills of the Sierra de Baoruco that suspicion was aroused that it might represent A. c. longitibialis. Comparisons were made, and it was determined this lizard was indeed much more similar in size and ventral scutellation to Beata lizards than to A. cybotes from the nearby mountains. In 1971, at a locality 9.6 km N of Pedernales, I collected cybotes and longitibialis at the same locality; the habitat was grassy and shrubby pasture adjacent to limestone cliffs and their talus. Since Thomas’s lizard was secured sleeping adjacent to a cave in limestone cliffs, it seemed a possibility that on the mainland /ongitibialis was limited to this habitat, whereas cybotes preferred more shaded and _ less extreme habitats. There was no doubt, however, that we were dealing with two distinct species, 4. cybotes and A. longitibialis, which have limited sympatry and syntopy on the Peninsula de Barahona, and of which one (A. /ongitibialis) occurs also on Isla Beata. In June 1974, Fred G. Thompson of the Florida State Museum collected in the Republica Dominicana on the north face of the Sierra de Baoruco near the Dominico-Haitian border, just northeast of the border post of El Aguacate. He courteously lent me his material from that expedition. Two anoles from 2 km NE of El Aguacate are extremely distinctive; both are males, one is very large, and each has its head more attenuate with a different profile and is much less “jowly” than A. cybotes. Most pertinently, the dewlaps were mustard-colored when I examined them 4 months later. Although this is not the dewlap color in life of this population, nevertheless, the retention of such deep and distinctive colors after preservation suggests the dewlaps are differently and brightly colored in life, in contrast to the pale colors of A. cybotes. In December 1974, Michael H. Strahm and I made an attempt to reach Thompson’s locality; however, the road to the north face of the Sierra de Baoruco from La Florida was in disrepair and our vehicle was unable to reach the high elevations (900 m) needed to secure the lizards. Dr. Thomson had advised me that the lizards were common on rocky and exposed roadcuts, but we never reached such a situation. In the summer of 1975 we were successful in reaching El] Agucate from the north, where the road has high to moderate artificial “cliffs” along its eastern margin as it ascends the mountains. We finally found a population of Thompson’s lizards 4 BREVIORA No. 451 and secured a series on three visits. My earlier contentions proved correct: 1) the dewlaps are very brightly colored, and 2) the lizards are large and are confined to creviced roadcuts in this area. Their status as a species distinct from A. cybotes was confirmed by taking typical A. cybotes on fenceposts across the road (Sm) from the cliffs which harbored the new lizards. Thus we seemed to have an easily solvable problem: a derivative of A. cybotes which 1s very special- ized in habitat (cliff faces) occurring on the north face of the Sierra de Baoruco at elevations of about 900 m. The two species differ grossly in adult size, head shape, and dewlap color as well as in habitat; there was no question that they are distinct and sympatric. Also during the summer of 1975, Ernest E. Williams and William E. Haas, while collecting in the Peninsula de Barahona lowlands, encountered large cybotoids in a shaded and fairly mesic ravine near sea level at a locality 5 km SE and 2.9-3.0 km N of Pedernales. Superficial examination of their specimens in the field, and subse- quent collection of a short series taken there by Thomas Wiewandt and Diderot Gicca, all served to confirm that these lizards were very comparable to the series from high elevations on the north face of this same mountain range. The habitat of the new population near Pedernales is very like that of the El Aguacate specimens — a limestone ravine or wash that is much creviced; however, vegeta- tionally this ravine is much more shaded and mesic than the bare and exposed road cut near El Aguacate. In 1975 Richard Thomas secured a fine series of A. /ongitibialis at a locality 17 km NW of Oviedo that represents the easternmost record for the species. This locality is a wooded but xeric ravine with creviced limestone walls, that lies on the eastern edge of the north- south limestone ridge which bisects the Peninsula de Barahona. There are specimens of A. /ongitibialis from west (Pedernales) and east (17 km NE of Oviedo) of the ravine which harbors the differently-colored dewlapped population discovered by Williams and Haas. Both species in this region show a predilection for limestone cliffs; they and the north slope lizards show remarkable agility and familiarity with their cliffs and are completely “at home” upon them. Other anoles (A. cybotes, A. brevirostris Bocourt) occur on cliffs in this general region but rarely seek refuge in crevices; rather, when pressed, they drop to the ground to hide. This is not true of either the new species or of A. Jongitibialis both of which 1979 NEW SPECIES OF CYBOTOID ANOLE 5 almost invariably seek refuge in crevices (see additional comments below). | have never taken the new species at night which suggests that the lizards sleep in crevices. A. /ongitibialis, however, has been secured while asleep on shrubs and low trees adjacent to cliffs. This implies that the two species differ in sleeping sites, although they may forage on the same vertical rocky faces. Williams (1975:7-8) made the following comment in his “‘justifi- cation” for the naming of A. marcanoi despite weak meristic differences between that species and A. cybotes: “It becomes more obvious that, in addition to those species in which museum tax- onomists rejoice because they are very distinct in terms of the characters conventionally studied, there are in many groups valid biological species only imperfectly separable on museum characters, if at all. This phenomenon is only interesting in terms of the history of museums, not of biology... . It will not be necessary in the near future to defend or specially comment on cases like that here described.” The same comments, to some extent, apply here. I have no doubt that the peculiarly isolated populations herein described as a new species differ biologically from 4. Jongitibialis. Except for some size and modal differences in scale counts, without preknowl- edge of the dewlap colors as a re-enforcing mechanism, one might be easily led into misinterpreting the situation. Verifying that A. longitibialis is distinct from A. cybotes required the chance en- counter of the two species syntopically after the lapse of two years. A. longitibialis geographically surrounds two of the three known populations of the new species. Differences between this new species and A. Jongitibialis are not subtle in life, but assigning long- preserved and discolored museum specimens to one or the other species is and will continue to be difficult. This fact, however, should not be a deterrent from naming a new taxon. Two biological entities (species) exist. It is the duty of the systematist to recognize this fact nomenclaturally and to present whatever data he may have to support his contention. Before proceeding to the new species, it is pertinent to examine variation in A. /ongitibialis. Noble’s (1923) description is detailed but lacks data on some features which are important when com- paring the Beata population with the Peninsula de Barahona population. These two samples differ from each other and from the new species about to be named in a number of ways; the differences 6 BREVIORA No. 451 in the former case are those which are accepted as being of subspecific rank. Accordingly, I first define A. /ongitibialis in those terms which will differentiate it from the new species, and then name a new mainland subspecies of A. /ongitibialis. Anolis longitibialis Noble Anolis longitibialis Noble, 1923, Amer. Mus. Novitates, 64:4. Type locality: Isla Beata, Republica Dominicana. Holotype: AMNH 24329. Definition. A cybotoid anole characterized by small size (males to 72 mm, females to 59 mm), supraorbital semicircles occasionally not in contact, modally |/1 scales between the supraorbital semicircles and the interparietal, smaller median dorsal scales (35 to 57 in snout- ear distance), subocular scales often in contact with supralabial scales, usually 4 or 5 postrostral scales, few fourth toe lamellae (15-21), dewlap pale yellow to pale orange, throats of adult males almost always immaculate and rarely streaked. Anolis longitibialis specuum, new subspecies Holotype: MCZ 132,370, adult male, 17 km NW of Oviedo Nuevo, Pedernales Province, Republica, 183 m, 7-8 August 1975, Richard Thomas, coll. Original number RT 3,461. Paratypes: All paratypes are from Republica Dominicana, Ped- ernales Province. ASFS V26,898-901, RT 3,462, LSUMZ 29,541- 47, same data as holotype; ASFS V16,728, 6.4 km SE Pedernales, 17 May 1969, J. B. Strong; ASFS V21,531,8 km N, 2 km E of Cabo Rojo, | August 1969, R. Thomas; ASFS V29,766-68, 7 km N, 20.0 km SE of Cabo Rojo, 183 m, 14 August 1971, A. Schwartz; ASFS V30,073-74, 7 km N, 20.0 km SE of Cabo Rojo, 183 m, 22 August 1971, D. C. Fowler; ASFS V42,235, 17 km NE of Oviedo Nuevo, 183 m, 12 August 1975, M. H. Strahm; ASFS V30,086-87, 7 km N, 17.6 km SE of Cabo Rojo, 152 m, 22 August 1971, A. Schwartz, B. R. Sheplan; ASFS V30,114, 9.6 km N of Pedernales, 244 m, 23 August 1971, A. Schwartz; ASFS V41,914, 7 km N, 2 km SE of Cabo Rojo, 29 July 1975, M. H. Strahm; UF/ FSM 21,567-68, | km SW of Las Mercedes, 380 m, 21 March 1974, R. Franz; MCZ 146,848, 1.2 km E of cave near intersection Cabo Rojo-Pedernales roads, 21 July 1975, W. E. Haas; MCZ 146,849, 7 km N, less than 1979 NEW SPECIES OF CYBOTOID ANOLE 7 1.2 km E of intersection of Cabo Rojo-Pedernales roads, 21 July 1975, W. E. Haas; MCZ 128,319, MCZ 128,342-43, 2 km E of turn tow Gabo: Rojo, 16 July 1971) T: P. Webster, R. B. Huey; MCZ 143,483, Cabo Rojo, behind police station, 5 July 1974, P. E. Hertz, R. B. Huey; MCZ 151,864-78, Cabo Rojo, behind laundry, 1-3 October 1976, W. E. Haas; MCZ 132,378-80, 7 km N, 1.2 km SE of Cabo Rojo, 2 October 1976, N. Atkins, W. E. Haas; MCZ 151,902, 7 km N, 1.2 km E of Cabo Rojo, 2 October 1976, W. E. Haas; MCZ 132,381-82,°17.6 km NW of Oviedo Nuevo, 2 October 1976, W. E. Haas; MCZ 151,828-48, 17.6 km NW of Oviedo Nuevo, 2-3 @ctober 1976; W.E. Haas: Associated specimen. MCZ 58,419, 30 km NW of Oviedo, Pedernales Province, Republica Dominicana. Definition. A subspecies characterized by the combination of large size (males to 72 mm, females to 59 mm, snout—vent length), larger dorsal scales (35-52), larger ventral scales (31-52), usually 4 or 5 postrostral scales, dorsum brown with 4 transverse dumbbells and often | pair of sacral blotches, throats of females almost always longitudinally streaked with dark brown, very rarely so in adult males; dewlaps very pale orange to very dull yellow or dirty yellow. Description. An adult male with a snout-vent length of 71 mm and femur length of 23 mm (measured as proposed by Ruibal and Williams, 1961:214): snout scales at second canthal scales 7, vertical loreal rows 4, supraorbital semicircles in contact, scales between semicircles and interparietal | / 1 (counted as proposed by Schwartz, 1968:260); subocular scales in contact with supralabial scales bilat- erally; 5 enlarged scales in supraocular discs, 7 postmentals, 4 postrostrals, 3/3 canthals, 19 fourth toe lamellae on phalanges II + III, 49 median dorsal scales and 43 median ventral scales in snout-ear distance. In life, dorsum brown with 4 darker brown dorsal dumbbells (narrow middorsally), the anterior pair the largest and forming a pair of blotches, somewhat faded along their anterior borders, the posterior 3 very fragmented and barely discernible as dumbbells; a dark but centrally very flattened postocular U, the arms very short and touching the upper eyelids; temples, neck, and remainder of dorsum with scattered longitudinal dark dashes or small spots; a pale narrow subocular crescent; lores and infraorbital area pale and without dark mottling; venter white, chin and throat immaculate: upper surfaces of fore- and hindlimbs concolor with 8 BREVIORA No. 451 dorsum and with a very few scattered dark brown dots and without defined crossbands; dewlap dull yellow. Variation. The series of 76 A. 1. specuum is composed of 38 males and 38 females. MCZ 151,828 is the largest male with a snout-vent length of 72 mm; the largest female (ASFS V21,531) has a snout-vent length of 59 mm. The smallest lizard is a female (RT 3,460) with a snout-vent length of 37 mm. Femora measurements (in mm.) of all males are 14.2-24.4 (x = 21.1) and are 11.8-19.2 (16.5) in all females. Snout scales between the second canthals vary between 6 and 10 (x = 7.4; M, = 7-44% of the specimens); loreal rows are 4 to 6(M, = 5-53%); supraorbital semicircles are in contact in 69 specimens and are separated by one row of small scales in five lizards. Scales between the supraorbital semicircles and the inter- parietal scale are modally 1/1 (40 specimens); other counts include 0/0 (2), 0/1 (), 1/2 (12), 2/2 (13). Dorsal scales m snout=ean distance are 35—52 (x = 44.3) and ventrals in the same distance are 31-52 (x = 38.5). The subocular scales may or may not be in contact with the supralabial scales; bilateral separation between these scales occurs in 44 specimens, unilateral contact occurs in six, and bilateral contact in 16; thus, there is contact at least unilaterally between the suboculars and supralabials in 33% of the lizards. Enlarged scales in the supraocular discs vary between five and 11 (x = 7.0; Mo =7 - 27%); this count 1s difficult to take since there may be many small (but not granular or tiny) scales along the periphery of the disc that might be counted if one desired. Postmental scales are 2 to 9 (x = 6.3; Mo — 41%); postrostral scales are 3 to 5 (x = 4.2; Mo = 4 or 5-36% in each category). Canthals are usually 3/3 but three lizards have 4/4 canthal scales and one has 3/4. Fourth toe lamellae on phalanges II + III are 15-21 (x = 17.6); femur/snout—vent length ratio X 100 is 28.7-36.3 (x = 32.5) in males, 28.8-35.2 (33.2) in females. Color notes in life on both males and females indicate that the dorsal ground color is brown with darker brown markings. There is often a white subocular semicircle and a gray lateral stripe which may be absent or may be suggested by the remnants of dark dorsal and ventral outlining. In males there are usually four dumbbell- shaped figures with their narrowest portions lying across the midline of the back and followed by a pair of dark sacral blotches; of the dumbbells, the first is the best defined and largest and 1s less clearly 1979 NEWSSREGIES, OFSCYBOTOID ANOLE 9 delimited anteriorly than posteriorly as in the holotype. Occasional adult males (MCZ 128,342) are virtually patternless above, without traces of the dumbbells and with only some longitudinal dashes and dark dots remaining. Another smaller male (ASFS V16,728) has only the finest transverse indications of the dumbbells, whereas ina comparably sized male (ASFS V30,074) the dumbbells are large and form prominent butterfly-shaped markings across the back. The truncate postocular U described for the holotype is present in most males, but may be very reduced, only indicated, or absent. A fine, vertically diagonal dark line often courses across the temporal region toward the dorsal midline and may even occasionally (MCZ 128,342) form a complete nuchal V or vague W. The throats of males are variable; young specimens have them streaked with dark brown while full adults have them most often immaculate. An exceptional male (RT 3,462) has streaking remnants on the throat at a snout-vent length of 71 mm. Females have the same dorsal pattern variation as do males. The dumbbells are, when present, conspicuous, dark, and well devel- oped. Some females (ASFS V30,086) have the dorsal pattern much reduced or even absent and the postocular truncate U is more often only indicated rather than present and well developed. One female (ASFS V41,914) was described in life as having an orangish middorsal streak with dark brown dorsal flecking between the dumbbells. The throats of females are variable — many are longitudinally streaked with dark brown whereas others have the throats immaculate white. Therefore there seems to be no correla- tion of size and throat streaking. Male dewlaps have been described as orange, pale orange, dirty yellow, or dull yellow (the latter two conditions at the type-locality). A male (ASFS V30,074, snout—vent length 54 mm) was recorded as having the dewlap orange (PI. 9K5; all color designations from Maerz and Paul, 1950); a topotypical male (ASFS V42,235; snout- vent length 67 mm) had the dewlap dull yellow (PI. 1216) shortly after death. The vestigial dewlaps in females were recorded as pale orange to orange. Comparisons. A. 1. longitibialis from Isla Beata (Schwartz and Thomas, 1975:89, incorrectly assigned A. /ongitibialis to Isla Alto Velo whence the species is unknown) differ from A. /. specuum in several ways. The Beata subspecies is smaller (males to 67 mm, 10 BREVIORA No. 451 females to 57 mm snout-vent lengths), the dorsal scales (39-57) and ventral scales (29-51) are smaller, and there are modally four postrostrals (46%). The series of A. / longitibialis consists of 30 males and 13 females. The dorsal coloration in life is grayish tan to gray-brown and the color closely matches the cliffs upon which the species lives on Isla Beata. In males, the lower sides are greenish and in this sex the pattern consists of a series of longitudinal fine dark brown lines, dashes, or dots; there is no indication of dark brown dumbbells. The truncate occipital U is barely indicated at best and is usually absent; the fine diagonal temporal line is present but faint or fragmented. All these features, plus dorsal ground color, differentiate A. / longitibialis from A. |. specuum. Throats of males are almost always immaculate; seven males show some vague streaking or dark scribbling on the throats. Females are much like the males dorsally, except that there may be a more clear indication of the dumbbells (AMNH 41,431) than in males. Female throats always show some indication of dark streaking or scribbling, but this is seldom bold and prominent. Noble (1923:4) noted that A. /ongitibialis from Isla Beata had the dorsum with “a fine penciling of dark brown.” He also commented that the color (in alcohol) was pale chocolate- brown, finely marked with a number of narrow lines of dark brown. Some of the lizards which were available to Noble have been examined by me and they still retain these features. One peculiarity of Noble’s description is his comments on dewlap color. He stated (1923:4) that the dewlap was “bright” and that it “oddly enough, sometimes retains its color in preservative.” This latter is a peculiarity of this species, of A. marcanoi, and of the one next to be described. Most Antillean anoles with orange, red, or yellow dewlaps lose the pigments shortly after preservation; but this seems not to be the case in these cybotoids. Despite Noble’s comments on dewlap color and its persistence in A. /ongitibialis, nowhere does he mention what are the characteristics of the bright dewlap color except to say that it is “yellowish.” Thomas’s (1964) field notes state that in males the dewlap is dirty yellow anteriorly to orange on the posterior three-quarters of the dewlap. It appears that the dewlap color in the two subspecies of A. /ongitibialis is fairly comparable. 1979 NEW SPECIES OF CYBOTOID ANOLE 1a Remarks. There is one specimen (MCZ 58,419, from 30 km NW of Oviedo, Pedernales Province) of whose status I am uncertain. This is an adult male 67 mm in snout-vent length. It differs from both A. /. specuum and the new species described below in that it has a complete dark nuchal band, three complete dark dorsal bands between the limbs, and a pair of sacral blotches that almost form a fourth complete band. There are no distinctive head markings and the specimen is somewhat faded (collected in 1958). If it is an A. /. specuum, it is remarkably well patterned dorsally; the throat is now immaculate. The small ventral scales assure that it is not A. cybotes. The locality places it within the distribution of the former taxon but I refrain from assigning it to A. /. specuum since it differs in pattern details as noted above. I suspect that it is an exceptionally well patterned male A. /. specuuwm, but the cautions noted in the introduction are re-enforced here: without knowledge of habitat or color and pattern in life, one is strongly handicapped in confidently assigning older specimens to any of these taxa. The name specuum is from the Latin for “of the crevices or caves” and alludes to the crevice-dwelling habits of this subspecies. The word specuum is genitive plural. Specimens examined. A. |. longitibialis, Republica Dominicana, Isla Beata, just E of Punta Beata (ASFS V17,215-19); no other locality (ASFS V2,772-80, USNM 83,878, USNM 83,880, AMNH 41.415, AMNH 41,422, AMNH 41,424-32,; AMNH 52,449-51, MCZ 17,686, MCZ 31,774, MCZ 37,480-82, UF/ FSM 21,572-78). The new species referred to in the introduction is composed of two populations which differ from each other. Michael H. Strahm collected most specimens of the northern population. Accordingly, I associate his name with this new species, which I call: Anolis strahmi, new species Definition. A cybotoid anole characterized by large size (males to 79 mm, females to 64 mm snout-vent lengths), supraorbital semi- circles always in contact, modally 2/2 scales between the supra- orbital semicircles and the interparietal, larger median dorsal scales (32 to 51 in snout-ear distance), subocular scales always separated from supralabial scales by one row of scales, usually three post- 2 BREVIORA No. 451 rostral scales, more fourth toe lamellae (17-24), dewlaps deep orange to deep orange-brown, throats of males marked, or without dark flecking or scribbling, by population. Head. Moderately massive, length from snout to posterior bor- der of eye much shorter than either femur or tibia. Head scales mostly smooth in males and very weakly carinate in females. Five to nine scales across snout at level of second canthal scales. A very shallow frontal depression and a deeper parietal depression. Nares in front of and above canthal ridge composed of three or four scales. Anterior nasal scale in contact with rostral. Snout distinctly pointed when viewed from above and with a distinctive profile due to the raised nares. Supraorbital semicircles in contact and are separated from the supraocular discs by 2 rows of granules. Supraocular discs consist of about 4 to II enlarged smooth to very weakly keeled (males) or keeled (females) scales separated by about 4 rows of scales and granules from the scales of the supraciliary rows. Two elongate supraciliaries are continued posteriorly by a double row of mod- erately enlarged scales. Canthus distinct, the first canthal scale the largest. Loreal rows five to nine, the lower rows larger and more regular. Supratemporal area scales are granular, grading rather abruptly into larger scales surrounding the interparietal depression. Interparietal is about the same size as ear opening and is separated from the supraorbital semicircles by one to three scales (modally two) or rarely unilaterally in contact. Subocular scales are almost always separated from supralabials by one row of scales (very rarely in contact, very rarely by two rows of scales), anteriorly grading into loreals, posteriorly grading into large scales at the corner of the mouth. Usually six supralabials to center of eye: Mentals are equally broad and long and almost equilateral, and are in contact posteriorly with one to ten small elongate postmental scales. Infralabials are broadly rectangular anteriorly, narrow pos- teriorly, in contact with three large tetrahedronal sublabials. Throat scales are small, granular, not keeled, and the anterior ones are elongate. Trunk. Middorsal scales are in two or four abruptly enlarged rows, about four times as large as flank scales and 32-51 in snout- ear distance. Ventrals are small, about 1.5 times as large as —— 1979 NEW SPECIES OF CYBOTOID ANOLE 13 middorsal series, cycloid, smooth, and 31 to 61 in snout-ear distance. Two postanal scales are enlarged in males. Gular fan. Very large; scales smooth, those along margin about twice as large as ventrals. Limbs and digits. Hand and foot scales are smooth. Between 14 and 24 scales are under phalanges II + III of fourth toe. Largest scales of arm are smooth to very weakly unicarinate, those of leg smooth and both those of arm and leg are larger than ventrals. Tail. Slightly laterally compressed, each verticil surmounted by three sharply keeled scales and ventrally by three pairs of unicari- nate slightly smaller scales. Anolis strahmi strahmi, new subspecies Holotype: MCZ 132,371, an adult male, from 3 km NE of El Aguacate, Independencia Province, 854 m, Republica Dominicana, taken on 19 July 1975 by Michael H. Strahm. Original number ASFS V41,729. Paratypes. All paratypes are from Independencia Province, Re- publica Dominicana. ASFS V41,730-34, same data as holotype; ASFS V41,284-94, same locality as holotype, 10 July 1975, M. H. Strahm; ASFS V28,453, ASFS V41,308—09, same locality as holo- type, 14 July 1975; ASFS V44,991-94, same locality as holotype, 22 December 1976, A. Schwzrtz, W. B. Southerland; UF/ FSM 21,565- 66, 2 km NE of El Aguacate, 900 m, 30 June 1974, F. G. Thompson. Definition. A subspecies of A. strahmi characterized by modally eight scales in the supraorbital discs, dorsum is pale gray and 1s at best flecked with dark gray and often unpatterned, throat is usually unpatterned in both sexes but occasionally has some vague scrib- bling, and dewlap is very deep orange to orange-brown. Description. An adult male with a snout-vent length of 78 mm and femur length of 25.5 mm; six snout scales at second canthal scales, seven vertical loreal rows, supraorbital semicircles in contact, scales between semicircles and interparietal 2/2; subocular scales separated by one row of scales from supralabials bilaterally: 11 scales in supraocular discs, six postmentals, three postrostrals, 3) 3 canthals, 20 fourth toe lamellae on phalanges II + III, 41 median dorsal scales and 48 median ventral scales in snout-ear distance. In life, dorsum is pale gray with a few scattered darker gray dots, most prominent above the forelimb insertions; remnants of a postocular 14 BREVIORA No. 451 truncate U are present, its very short arms abutting against the upper eyelids; supra- and infralabials contrastingly spotted with pale and dark gray; a very pale blue-gray subocular crescent; venter pale grayish, throat immaculate and without pattern; limbs and tail without any crossbands or other prominent markings; dewlap (shortly after death) deep orange (Pl. 4AI1). Variation. The series of 26 A. s. strahmi consists of nine males and 17 females. The holotype and another male (UF/ FSM 21,566) are the largest males with snout-vent lengths of 78 mm; the largest female (ASFS V41,288) has a snout-vent length of 64 mm. The smallest lizard (ASFS V44,994) is a female with a snout—vent length of 42 mm. Femora measurements (in mm.) in all males are 15.7-26.4 (x = 21.5) and are 13.0-21.5 (18.9) in all females. Snout scales between the second canthals vary between five and eight (x = 7.0; M.= 7-43% of the specimens); loreal rows are five to seven (M, = 5-52%); supraorbital semicircles are in contact in all specimens. Scales between the supraorbital semicircles and the interparietal scale are modally 2/2 (14 specimens), with other counts of 1/0 (1), 1/1 (2), 1/2 (4), and 2/3 (1). Dorsal scales in snout-ear distance are 34-48 (x = 39.3) and ventrals in the same distance are 32-61 (x = 41.2). The subocular scales are separated from the supralabial scales by one row of scales bilaterally in all but two lizards which have either two rows of scales or are in contact unilaterally. Enlarged scales in the supraocular discs vary between five and 12 (x = 8.0; M, = 8-36%); the same precautions made in the account of A. /. specuum apply here. Postmental scales are four to seven (x = 5.3; M, = 6-36%); postrostral scales are three to five (x = 3.5; M, = 3-58%). Canthal scales are always 3/3. Fourth toe lamellae on phalanges II + III are 16-21 (x = 18.6); femur/snout—vent length ratio X 100 1s 31.7-34.3 (x = 33.3) in males and 30.5=36:1' G29) females. Color notes on both males and females in life indicate the dorsal ground color is gray, remarkably similar to the color of the rocks on which the lizards live. Both sexes are patterned similarly dorsally; there are no dumbbells or flank stripes present (although the position of the latter is occasionally indicated by vague elongate dashes along its putative upper and lower margins). The dorsum 1s more or less randomly dotted or marked with scattered dashes aligned longitudinally; young males have vague dumbells which are 1979 NEW SPECIES OF CYBOTOID ANOLE 15 so reduced as to be only faint transverse lines hollowed laterally i.e., they are mere vestiges of what presumably are basically dumbbells. The general aspect of the males and females is of a faint and randomly dotted or lineate pattern, but many adults and subadults completely lack any sort of dark body pattern elements. The postocular truncate U is represented by its outlines and these may be fragmented and very obscure. The same is true of the diagonal temporal line. The venter is dark gray to yellowish gray in females. The throat is marked with some confused scribbling, oriented longitudinally in one adult male (ASFS V41,285) and in one subadult male (UF/FSM 21,565; snout-vent length 54 mm). Females generally have gray throats without markings, but one adult (ASFS V41,290; snout-vent length 62 mm) has some vague darker scribbling and two other females have remnants of similar markings only very faintly shown on the posterior portion of the throat. In life, the limbs are crossbanded with pale and dark gray, but these markings are not obvious on the preserved specimens. A distinctive feature of males (and somewhat less so of females) is the spotted supra- and infralabials. A very pale blue-gray subocular crescent is present and moderately conspicuous in life. The dewlaps in males are very deep orange to orange-brown and these colors are especially rich and almost velvety in texture. Color designations are Pl. 13J10 for one living male and PI. I1I11 and PI. 4Al1 for two recently dead males. Females have the same basic intensity of coloration on their vestigial dewlaps. Comparisons. Comparisons of A. s. strahmi with the subspecies on the south side of the Sierra de Baoruco as well as with A. /. specuum will be withheld until the former is described below. Remarks. 1 have commented previously on Dr. Thompson’s observations on the habitat of A. s. strahmi. Our own observations amply confirm his. On 10 July 1975 we searched several kilometers of the road below the post of El Aguacate and saw only very occasional lizards on the gray rocks of the roadcut and did not observe any on adjacent shrubs or herbs. In much of this distance, the roadcut is rather densely covered with vines, small shrubs, and herbs, but elsewhere it is almost bare with only scattered vegeta- tional cover. We finally encountered 4. s. strahmi in some abun- dance along an exposed section of roadcut which was about 30 m in length. The roadcut here was almost bare of plants with many 16 BREVIORA No. 451 crevices and solution holes. Between 1330 and 1600 hrs this section of roadcut was shaded (due to the direction of its face rather than to any arborescent or herbaceous cover) and the lizards were seen in their retreats, their heads or foreparts extended outside the crevices in an alert manner. They were not easily alarmed, relying upon the extremely cryptic coloration of their gray dorsa agreeing very closely with the hue of the roadcut face. Two lizards were seen “sunning” themselves (in the shade) vertically, head down, on completely exposed rocky surfaces; one was observed clinging to the roof of a small solution hole. In addition to the 11 specimens collected during this 2.5 hour period, at least eight others were seen but not collected. They allowed close (1 m or less) approach of the collector before becoming alarmed and retreating into crevices or solution holes. Recovery time was brief; on several occasions, as we walked slowly down this 30 m stretch of road, we noted refuges of specific lizards. Upon returning a maximum of 10 minutes later the lizards had already reappeared at the crevice mouths (which were their individual retreats) or on adjacent rock faces. On subsequent visits we observed lizards in the same precise places as previously. Their agility and assurance on the rock faces were impressive. On our third visit to the type locality on 19 July 1975, six lizards were collected between 1720 and 1830 hrs in the same stretch of roadcut. The adult male holotype was taken just below a slightly more covered section. About three or four other anoles were seen but escaped capture. Although the cliff face was completely in shade and the air was cool (but not cold) the rock faces were still warm to the touch. On 22 December 1976, W. B. Southerland and I visited the type locality once again. Our arrival was at 1200 hr and we saw very few lizards. The roadcut was in shade and the shade increased until 1500 hr so that maximum insolation had occurred in the early morning prior to our arrival. However, the rocks were still warm to the touch. As the afternoon progressed, more A. s. strahmi appeared, so that about 12 were seen. Some large males, once disturbed, did not reappear on the roadcut face. Four specimens were taken: a male on an open rock face, a female on a dead stem 8 cm from the base of the cliff and two other females in or near crevices or cavities in the roadcut face. The female noted here on a dead stem is the only individual we observed or collected not on the rocks. 1979 NEW SPECIES OF CYBOTOID ANOLE 17 Above and below the road at the type locality are steep slopes. An outstanding feature of the immediate region 1s the eroded limestone terraces and jumbled rocks and boulders, all sparsely covered with herbs and scattered shrubs and with only occasional large trees. The general aspect of the area is rather bleak. Two other observations at the type locality are pertinent. We encountered an active Uromacer catesbyi Schlegel foraging on the roadcut face-occupied by A. s. strahmi. These snakes eat lizards but are rarely seen in this habitat type and prefer trees and shrubs. Ina crevice at this site we also encountered a young Epicrates striatus Fischer; when young, these snakes eat lizards. It is not unlikely that both snakes were associated with this particular section of roadcut because of the ready supply of food (A. strahmi). Both snakes are diligent predators in that they search for prey by penetrating crevices, holes, cavities, and other possible hiding places. Elongate U. catesbyi is especially well adapted for this foraging pattern during the day and E. striatus forages similarly at night. Anolis strahmi abditus, new subspecies Holotype: MCZ 146,827, an adult male, from dirt road to Las Mercedes, 2.9 km from intersection (= 5 km SE, 2.9 km N of Pedernales), Pedernales Province, Republica Dominicana, 19-20 July 1975 by William E. Haas collector. Original number MCZ F- 29,006. Paratypes. All paratypes are from Pedernales Province, Repub- lica Dominicana. MCZ 146,920, MCZ 146,828-47, same data as holotype; ASFS V41,908-12, same locality as holotype, 29 July 1975, M. H. Strahm; UF/ FSM 34,423-27, 5 km SW, 2.5-3.0 km N of Pedernales, 8 August 1975, T. Wiewandt and D. Gicca; MCZ 151,879-901, 5 km SE, 2.9 km N of Pedernales, 30 September and | October 1976, W. E. Haas; MCZ 132,383, 5 km SE, 2.9 km N of Pedernales, 30 September 1976, N. Atkins and W. E. Haas; MCZ 151,849-54, MCZ 151,857-63, between 15 and 16 km N of Cabo Rojo, Alcoa road, 3 October 1976, W. E. Haas. Definition. A subspecies of A. strahmi characterized by modally six or seven scales in the supraorbital discs; dorsum tan to grayish tan, at times marked with small brown to reddish blotches; lateral flank stripe tan to whitish present, dorsum with transverse dumb- 18 BREVIORA No. 451 bells or hourglasses in both sexes; throat flecked or scribbled in both sexes with dark brown and dewlap deep orange. Description. An adult male with a snout-vent length of 78 mm and femur length of 26 mm; six snout scales at second canthals, supraorbital semicircles in contact, scales between semicircles and interparietal 2/2, subocular scales separated by one row of scales from supralabials bilaterally; seven scales in supraocular disc, six postmentals, three postrostrals, 3/3 canthals, 19 fourth toe lamellae on phalanges II + III, 35 median dorsal scales and 40 median ventral scales in snout-ear distance. As preserved, the dorsum is grayish brown, with four dark brown dumbbell remnants, all much hol- lowed or with their lateral expanded ends virtually missing and an additional fifth dumbbell remnant postsacrally; a dull interocular bar and a truncate postocular U, both hollowed; lores, supra-, infra-, and sublabials mottled dark and pale brown; sides of body with elongate dark brown linate fragments, with two of the lowermost fragment-series in part outlining the flank stripe; limbs marbled or mottled dark brown and brown, the only clear transverse dark bar on the crus; both fingers and toes with a moderately clear dark brown crossband; venter grayish to tan, chin and throat dotted with dark brown, sides of abdomen flecked with grayish brown; dewlap pinkish 6 months after preservation. Variation. The series of 68 A. s. abditus consists of 33 males and 35 females. The largest male (ASFS V41,911) has a snout-vent length of 79 mm and the largest female (MCZ 146,838) a snout-vent length of 63 mm. The smallest lizard (UF/FSM 34,425, a female) has a snout-vent length of 39 mm. Femora measurements (in mm.) in all males are 16.0—27.4 (x = 21.7) and are 13.2—20.8 (18.1) in all females. Snout scales between the second canthals vary between five and nine (x = 6.9; M, = 7-52% of the specimens); loreal rows are four to seven (M, = 5-62%); supraorbital semicircles are in contact in all specimens. Scales between the supraorbital semicircles and the interparietal scale are modally 2/2 (35 specimens), with other counts of 1/1 (9), 1/2 (5), 2/3 (6), and 3/3 (8). Dorsal scales in snout-ear distance are 32-51 (x = 39.8) and ventrals in the same distance are 31-53 (x = 40.2). The subocular scales are separated from the supralabial scales by one row of scales in all but two specimens; these lizards have these scales in contact either bilaterally or 1979 NEW SPECIES OF CYBOTOID ANOLE 19 unilaterally. Enlarged scales in the supraocular discs vary between four and 13 (x = 7.3; M, = 6 or 7-27% in each case). Postmental scales are four to 10 (x = 7.3; M. = 6—-33%); postrostral scales are three to five (x = 3.7; M, = 3-43%). Canthal scales are most often 3/3, but one lizard has 2/2 and another 3/4 scales in this position. Fourth toe lamellae on phalanges II + III are 17-24 (x = 19.0); femur/snout-vent length ratio X100 is 28.7-38.4 (x = 33.5) in males and 30.5—36.3 (33.4) in females. Color notes state that in both males and females in life the dorsum is tan to grayish tan with five narrow dumbbell or hourglass remnants; these dorsal pattern elements end above the flank stripe which is either tan or white. Despite its prominence in life, the flank stripe is outlined only by dark brown line fragments, including the remnants of the dorsal transverse dumbbells. A dark brown inter- ocular stripe and a postocular truncated U are both present and may be either prominent or vague; the U may be most conspicuous at its posterolateral “corners” on the superior temporal region. Some males (ASFS V41,910—-12, MCZ 146,829) show a progressive degen- eration of the dorsal pattern into a series of more or less random dumbbell-fragments and associated longitudinal lines or dots which are brown to reddish. In each of these lizards, the peculiar dorsal pattern is obviously a derivative of the customary dorsal dumbbells. Two females (ASFS V41,909, MCZ 146,838) are comparable to the above mentioned males in diminution of the dorsal pattern. In both sexes, the throat is marked with some sort of pattern; this varies from marbling or scribbling to longitudinal dark lines which is more common in females than males. The intensity of the throat markings is variable. The lores and supra-, infra-, and sublabials are contrast- ingly marked with dark and pale and the lip markings extend ventrally to give rise to the throat pattern. The male dewlaps are deep orange in life (Pl. 12F9); two dead males had comparable dewlaps (PI. 12F9, Pl.12H8). The deep orange color of the dewlap center is slightly richer along the margin. Comparisons. A. s. abditus requires comparison with the taxa A. s. strahmi and A. |. specuum. Anolis cybotes also occurs sympatri- cally with both subspecies of A. strahmi, but it is easily distinguish- able in dewlap color (pale yellow, pale pink, almost white), smaller snout-vent length, and much larger ventral scales. Anolis breviros- 20 BREVIORA No. 451 tris, a small gray dorsoventrally compressed species with a pale yellow dewlap and a paramedian double row of snout scales, also 1s syntopic with A. s. abditus. The two subspecies of A. strahmi are very similar, although they are altitudinally and geographically separated. The dewlaps in both are deep and rich orange (to orange-brown in A. s. strahmi) and reach the same size in both sexes. A. s. strahmi differs from A. s. abditus in that the former has modally eight scales in the supra- orbital discs versus six or seven in A. s. abditus. Dorsal coloration in A. s. strahmi is pale gray versus tan to grayish tan in A. s. abditus. The dorsum is at best flecked and often unpatterned in 4. s. strahmi males versus relatively prominent dumbbell or hourglass remnants in both sexes of A. s. abditus. Finally, throats in both sexes of A. s. strahmi are rarely marked versus throats in both sexes of A. s. abditus which are marked with flecking, dotting, scribbling, or lines. The dorsal color and pattern, as well as the throat markings, are the most distinctive characteristics separating the two subspecies. A. s. abditus has not as yet been taken sympatrically or syntopi- cally with A. 1. specuum (see Fig. 1); it is unlikely that these two species will be taken together since both are clivicolous. Though they seem to be ecological equivalents, it is possible that at some locality on the Peninsula de Barahona they occur together. The two species should be easily differentiable, if they are sympatric. The dewlap in A. s. abditus is deep orange, whereas that of A. /. specuum is dull or dirty yellow to pale orange. A. s. abditus reaches a larger size (males to 79 mm, females to 63 mm) than does 4. /. specuum (males to 72 mm, females to 59 mm). Although there are three to five postrostral scales in both taxa, the mode in A. s. abditus is three and in A. /. specuum is four or five. Two other scale relationships will aid in differentiating the two species. In A. /. specuum, there are more often 0/0—1/1 scales between the supraorbital semicircles and the interparietal (64% of the specimens) whereas in A. s. abditus there are more often |/2~-3/3 (79%) scales in this position. A. /. specuum more often (33%) has the subocular scales in contact with the supralabials than does A. s. abditus (3%). Although none of these scale counts or relationships is absolute, in combination they serve to differentiate the two species. There are some striking resemblances between A. /. specuum and A. s.abditus in color and pattern. The former is brown with darker 1979 NEW SPECIES OF CYBOTOID ANOLE DA a Mercedes Pedernales O 18°00 Figure 1. Map of the Peninsula de Barahona, Republica Dominicana, Hispan- iola, and Isla Beata. Pertinent population centers are indicated by labeled squares. Known distributions are shown for the anoles 4. /ongitibialis (triangles) and A. strahmi (circles). The distributions are further designated for 4. /. /ongitibialis (solid triangle), A. /. specuum (hollow triangles), A. s. strahmi (solid circles), and A. s. abditus (hollow circles). 22 BREVIORA No. 451 brown markings whereas the latter is tan to grayish tan with brown markings. Both are prominently marked dorsally with 4 dumbbells; however, in A. /. specuum, the first dumbbell is well developed laterally and open anteriorly to yield a large almost ocellus-like spot. The dewlap colors of the two species are extremely distinctive since no A. /. specuum is known that has the deep orange dewlap of A. s. abditus. One final feature serves to distinguish the two species in this area. A. /. specuum males most often have the throat immaculate as adults (one adult male is known with a streaked throat) whereas the throat is invariably patterned in A. s. abditus males. All A. s. abditus females have marked throats whereas the throats of A. /. specuum females are variable — from immaculate to streaked with dark brown. Remarks. There are only two localities for A. s. abditus. The first is a ravine (the type locality) between the Pedernales-Oviedo road and the village of Las Mercedes. At about 2.5 km N of the main road, the dirt road to Las Mercedes passes (for about | km) through a shaded and broad ravine with reddish rocky natural cliffs on both sides. These cliffs are creviced and have solution holes, but are separated from the road itself by low talus slopes which extend some five to ten meters between the cliffs and the roadway. The flora of the ravine stands in strong contrast to the Acacia-cactus desert which surrounds it; the ravine is distinctly more mesic, with trees and an understory of grasses, herbs and Brvophyllum. At the time of our 1975 visit the outside desert was exceptionally parched whereas the ravine was modestly luxuriant and moist. The ravine is almost at sealevel but at its northern end the road begins the ascent of the lower front ranges of the Sierra de Baoruco on which the village of Las Mercedes lies. The second locality is on the Alcoa road between Cabo Rojo and Aceitillar; I have not visited the site but William E. Haas (in /itt., 22 October 1976) wrote that it is “up a path just past km 16 on the Alcoa truck road, and the habitat is somewhat different [from the type locality] and the physiography a bit more so.” At the time of our 29 July 1975 visit to the type locality, the lizards were scarce at 1300 hr. The sun was almost directly overhead and the ravine was hot although fairly well shaded. As the afternoon progressed, the lengthening shadows of the western wall began to fill the ravine and by 1530 hr the lizards were moderately abundant. 1979 NEW SPECIES OF CYBOTOID ANOLE 23 They inhabited crevices and solution holes in the cliff faces (much after the fashion of A. s. strahmi). One male was seen foraging at 1630 hr on a shrub adjacent (I m) to the cliff and had a large caterpillar in its mouth when collected. Foraging thus involves securing food elsewhere than directly on the cliff face itself. A very small juvenile was dislodged from the basal rosette of a moderately sized dead sisal plant (Agave) on the cliff face but escaped without being collected. One behavorial incident is worthy of report: we pursued a large male along the cliff face and the lizard finally leaped to an isolated large (2 m diameter) boulder, ran across its top at full tilt and literally launched itself off the far side of the boulder onto the ground. Several minutes later the same lizard was collected within a few centimeters of the boulder’s base, immobile in the grassy and herbaceous ground cover. Whether its immobility was traumatic or whether it was “hiding” in a very unusual situation is unknown. This is one of only two instances when A. strahmi was secured in any situation other than a clivicolous one (the other exception is the female A. s. strahmi on the dead stem near the cliff base at El Aguacate). As with A. s. strahmi, A. s. abditus is impressively “at home” on its cliff faces. It is very agile, and its camouflage, tendency to remain immobile when approached — relying upon its cryptic coloration, and its prompt recovery time, all are extremely similar to those features in the behavior of A. s. strahmi. DISCUSSION It seems obvious that A. Jongitibialis and A. strahmi form a compact and closely related duo of cybotoid anoles whose distribu- tion centers upon the Peninsula de Barahona. Although | am convinced that they represent two separate species rather than subspecies, the latter interpretation is not to be lightly dismissed. There are no absolute meristic characters which separate the two, and they are similar in many features, not the least of which is their predilection for cliffs and vertical rocky surfaces to which they are unquestionably adapted. If, however, A. /ongitibialis and A. strahmi are conspecific, the peninsular distribution of the latter is peculiar in that it seems to be in enclaves surrounded on both east and west by (or interdigitating with) A. Jongitibialis. This arrangement can be 24 BREVIORA No. 451 interpreted as a mosaic, where one species (A. /ongitibialis) inhabits open and xeric cliff faces and the other (A. strahmi) inhabits shaded, more mesic, and less rigorous cliffs. The dewlap colors of the two species are very distinctive and since dewlap color in eye-minded anoles is sO important in species recognition and territorial defense, the very fact of the striking color differences of dewlaps in these two taxa strongly suggests that they are indeed species. The situation here is not so complex as that between 4. cybotes and A. marcanoi (Williams, 1975): in this case two unquestioned species without strong meristic differences but with distinctive and contrasting dewlap colors and different body patterns are sympatric and even rarely syntopic. In these two species, however, there appear to be subtle differences in habitat preference, and as Williams (1975:9) pointed out the “balance of power” between them in any particular area may be tenuous. The same situation may exist in the geo- graphical and ecological relationships between A. /ongitibialis and A. strahmi on the peninsula but presently the differences seem more overt. More puzzling is the occurrence of two subspecies of A. strahmi on the north and south slopes of the Sierra de Baoruco. This is coupled with the striking difference in elevations involved (near sea level, and between 854 and 900 m). The habitats of the two subspecies are comparable — generally creviced cliff faces or roadcuts which can be regarded as cliff-face surrogates. But the similarity ceases with this description. The ravine near Pedernales, although it transects an otherwise hostile and xeric area, has relatively luxuriant vegetation and 1s well shaded. The El Aguacate roadcut 1s open and exposed to direct sunlight (the same is true for localities where A. /ongitibialis has been taken on the peninsula itself) and the lizards seem to shun those sections of the roadcut that are heavily vegetated and overgrown. Even sections of the roadcut that are exposed and open often lack or have minimal populations of A. strahmi. The area is quite cool in the afternoon and cold at night. One explanation for the apparently precise niche occupied by A. s. strahmi may well be that, at such high elevations, insolation of cliff faces and lack of vegetational cover will determine the diurnal rhythm of these lizards. Cliffs, exposed or not, facing in directions Where the sunlight will not reach them until rather late in the morning are unsuitable because of lack of time exposed to the 1979 NEW SPECIES OF CYBOTOID ANOLE D5 warming effect of the sun. In contrast, vegetationally covered cliffs, even those oriented to achieve maximum early insolation, are too protected by vegetation to allow the lizards a sufficient period of activity during each day. The disjunct nature of the two subspecies of A. strahmi may be more an artifact than a reality. There are few suitable cliff faces readily attainable by road in this entire region. It seems likely that A. strahmi inhabits suitable habitats around the general periphery of the Sierra de Baoruco. Cliffs and roadcuts on the peninsula along the eastern edge of the mountains between Barahona and Enriquillo were searched casually. These cliffs are often at sea level but are well shaded and generally face east. We found both A. cybotes and A. brevirostris here but not A. strahmi or A. longitibialis. Such negative evidence is questionable at best, although we did not find A. cybotes syntopically on cliffs or roadcuts with either of the clivicolous species. However, A. cybotes does occur sympatrically (but not on cliffs) with A. / specuuwm and both subspecies of A. strahmi. It is interesting to note that in those places where A. cvbotes inhabits cliffs and roadcuts it is much less adapted for such habitats than are the two clivicolous species. When pressed A. cybotes is more likely to leave the cliff face and jump to the ground to seek refuge under adjacent ground cover than are A. /ongitibialis or A. strahmi which seek sanctuary in crevices or solution holes. Geographically it seems likely that there are more or less discon- tinuous populations of A. strahmi scattered around the periphery of the Sierra de Baoruco (but not across its summit) and the lower eastern regions of the adjacent Haitian Massif de la Selle. Popula- tions should be looked for on the bare roadcut-cliffs between Fond Parisien and Soliette and on the northern face of the La Selle in Haiti near the Dominico-Haitian border south of Fond Verrettes. Although these cliffs seem suitable (more for A. s. abditus than for A. s. strahmi), it may be that an appropriate habitat niche occurs in this region which will accommodate the latter subspecies. If A. s. strahmi did occur there at suitable elevations on the northern face of the Massif de la Selle, it may now be extirpated due to extreme modification of habitat (the region has been denuded of much vegetational cover). A. longitibialis, on the other hand, presents a more familiar pattern. It is a Barahona Entrapment xerophile derived from 26 BREVIORA No. 451 primarily mesophilic A. cybotes with which, under favored circum- stances, it is still sympatric. From the peninsula it has invaded Isla Beata and differentiated on the subspecific level. A phyletic se- quence between these three species might well be: widely distributed, ecologically tolerant but primarily mesophilic A. cybotes with a pale dewlap — xeric and cliff adapted A. /ongitibialis with a pale dewlap — clivicolous and more mesic adapted 4. strahmi with a vividly colored dewlap limited to disjunct enclaves where its ecological requirements are rather precisely met. Another possibility that should not be overlooked is that the four taxa involved (/ongitibialis, specuum, strahmi, abditus) should be regarded as separate species. Certainly they are isolated from each other (/ongitibialis is insular; specuum occurs on the Peninsula de Barahona; strahmi and abditus occur to the north and south of the Sierra de Baoruco massif, respectively). Regarding them as species has much to recommend it, but I have taken the more conservative course that no external characteristics are sufficiently obvious to make one seriously consider that there are four species involved rather than two. However, skeletal or electrophoretic evidence might well prove differently. ACKNOWLEDGMENTS I wish to thank the following curators for allowing me to study material in their collections: Richard G. Zweifel, American Museum of Natural History (AMNH), Douglas A. Rossman, Museum of Zoology, Louisiana State University (LSUMZ), Ernest E. Williams, Museum of Comparative Zoology, Harvard University (MCZ), Richard Thomas, University of Puerto Rico (RT), Walter A. Auffenberg, University of Florida, Florida State Museum (UF/ FSM), George R. Zug and Ronald I. Crombie, National Museum of Natural History (USNM). Specimens collected by myself and parties are in the Albert Schwartz Field Series (ASFS). Much pertinent material in the MCZ has been collected under National Science Foundation grant GB-37731X and preceding grants to Ernest E. Williams, and some specimens in the ASFS have been collected under National Science Foundation grants GB-7977 and B-023603. In the field I have had the competent assistance of the following men, without whose help much of the material for the present problem would not be available: Danny C. Fowler, David 1979 NEW SPECIES OF CYBOTOID ANOLE ay C. Leber, James A. Rodgers, Jr., W. Barry Southerland, Michael H. Strahm, James B. Strong, Bruce R. Sheplan, and Richard Thomas. Most especially I am grateful to Fred G. Thompson for allowing me to examine the first specimens of A. strahmi. LITERATURE GiiED COCHRAN, DM. 1934. Herpetological collections made in Hispaniola by the Utowana expedition, 1934. Occ. Papers Boston Soc. Nat. Hist. 8: 163-188. 1939. Diagnoses of three new lizards and a frog from the Dominican Republic. Proc. New England Zool. Club 18: 1-3. 1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus. 177: vii + 398 pp. MAERZ, A., AND M. R. PAuL. 1950. A dictionary of color. McGraw-Hill Book Co. Inc., N.Y., vii + pp. 1-23 and 137-208, 56 pls. Nosie, G. K. 1923. Four new lizards from Beata Island, Dominican Republic. Amer. Mus. Novitates 64: I-S. RuIBAL, R., AND E. E. WILLIAMS. 1961. The taxonomy of the Anolis homolechis complex of Cuba. Bull. Mus. Comp. Zool. 125(8): 211-246. SCHWARTZ, A. 1968. Geographic variation in Anolis distichus Cope (Lacertilia, Iguanidae) in the Bahama Islands and Hispaniola. Bull. Mus. Comp. Zool. 137(2): 255-309. , AND R. THOMAS. 1975. A check-list of West Indian amphibians and reptiles. Carnegie Mus. Natur. Hist. Spec. Publ. !: 216 pp. WEBSTER, T. P. 1975. An electrophoretic comparison of the Hispaniolan lizards Anolis cybotes and A. marcanoi. Breviora, No. 431: 1-8. WILLIAMS, E. E. 1961. Notes on Hispaniolan herpetology. 3. The evolution and relationships of the Anolis semilineatus group. Breviora, No. 138: 1-8. 1975. Anolis marcanoi new species: sibling to Anolis cybotes: description and field evidence. Breviora No. 430: 1-9. . Pa MUS. COMP, ZOOL LIBRARY MAR 1 8 1985 BRE VIO Ryde Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, MASS. 21 FEBRUARY 1979 NUMBER 452 LIZARDS OF THE SCELOPORUS ORCUTTI COMPLEX OF THE CAPE REGION OF BAJA CALIFORNIA WILLIAM P.HALL!:2 AND HOBART M. SMITH! ABSTRACT. Sceloporus orcutti licki as hitherto defined is shown to embrace two cryptic species, neither of which is subspecific in relation to the allopatric S. orcutti: an arboreal species, S. /icki, and a petricole, S. hunsakeri sp.n., from 3 mi. E of San Bartolo (holotype MVZ 73,570); both are restricted to the Cape region of Baja California. All three species, S. orcutti, S. licki and S. hunsakeri have identical 2n = 34 (XY 4 ) karyotype patterns, but differ in numerous other respects. In these three species, the most succinctly diagnostic character states are: presence of a dorsolateral light stripe in S. licki only, 59 or fewer dorsal scales plus femoral pores in most (96% of 27) S. orcutti, and 61 or more in most S. hunsakeri (98% of 61). INTRODUCTION Field work in 1969 indicated that the Cape Region of Baya California (the area south of the Isthmus of La Paz) is inhabited by three distinct species of Sceloporus which are broadly sympatric and frequently syntopic with one another (Figs. |, 2). Two of the three species, Sceloporus licki Van Denburgh and a species we describe here, are sufficiently similar to one another and to S. orcutti Stejneger that Smith (1939) considered them variants of a single race, S. orcutti licki. However, Hall’s discovery in 1969 that the licki variant was arboreal and the other petricolous prompted reexam- ination of material from Baja California Sur. Specimens of the unnamed taxon have been available in museums since 1890. ‘Department of Environmental, Population and Organismic Biology, University of Colorado Boulder, Colorado 80309 *Current Address: Department of Genetics University of Melbourne, Parkville, Victoria 3052, Australia No. 452 BREVIORA : ia Tel SAN DIEGO-- : } ° UY S ENSENADA M,-~4 aloye I FOC MIC Ocean oO Sce/oporus orcutti cs) Sceloporus n.sp. ras Sceloporus /icki ie) 150 Kr -—L—__1_ paeae Distribution of the three species of the Sceloporus orcutti complex in Fig. 1. Baja California. 1979 LIZARDS OF THE SCELOPORUS ORCUTTI 3 EL TRIUNFO Z\ an SAN BARTOLO °. cer ‘ e DG e g f #_acua e CALIENTE PoKearie A Ocean SAN JOSE DEL CABO Sceloporus orcutti a CABO DE SAN LUCAS e Sceloporus n.sp celoporus /icki WY 0 30 Km ——1__1___j Fig. 2. Distribution of Sceloporus hunsakeri, licki and orcutti on or near the Cape region of baja California. 4 BREVIORA No. 452 Comparative material relevant to this problem has been exam- ined from most of the major museums in the country. We are indebted to the curators and directors of these institutions. Cited specimens are referenced to collections by the following acronyms: CAS, California Academy of Sciences; CUM, University of Colo- rado Museum; LACM, Los Angeles County Museum; LMK, L. M. Klauber pers. coll.; LSJU, Leland Stanford Jr. Univ.; MCZ, Museum of Comparative Zoology, Harvard University; MVZ, Museum of Vertebrate Zoology, University of California, Berkeley; SDSNH, San Diego Society of Natural History Museum; UMMZ, University of Michigan Museum of Zoology; and USNM, U. S. National Museum of Natural History. We name the newly recognized species for Dr. Don Hunsaker II, in recognition of his pioneering studies in the comparative ethology of the genus Sce/oporus and for stimulating karyological study of its species by Hall. Sceloporus hunsakeri, new species Holotype: MVZ 73,570, adult male, tail complete, intact although fractured in two places in distal half, not regenerated, from 3 mi. E of San Bartolo +500 ft., Feb. 17, 1960, R. G. Crippen, field no. 277. Paratyvpes: Thirty-three, including eight topotypes, two of which were taken on the same date as the holotype (MVZ 73,572, 73,575) and six on March 14, 1960 (MVZ 73,579-84); two from 5 mi. SW of San Jose del Cabo, July 6, 1970, Richard L. Holland (CUM 45,880-1); two from Migrino, 33 mi. SSE of Todos Santos, July 5, 1969, RLH (CUM 40,779-80); one from '4 mi. N of Migrino, July 8, 1970, RLH (CUM 45,882); one from 14 mi. N of Migrino, July 9, 1970, RLH (CUM 45,883); two from 7 km WNW of Buenavista, 100 Km 105.32 on Mex. Hwy. 1, Aug. 3, 1969, W. P. Hall (MGZ R122,242); one from Cabo San Lucas, Aug. 4, 1919, J. R. Slevin (MCZ 15,565); one from 2 mi. NW of Cabo San Lucas, P. Elias, M. Merrill, and J. Lazell, Jan. 29, 1972 (MCZ R131,704); one from Cabo San Lucas, Elias, Merrill and Lazell, Jan. 29> 1972\(MiGz R131,706); 10 from 6.4 mi. FE of San Lucas, Playa Publica, T. Paul Maslin, Apr. 8, 1974 (CUM 51,401-10); and three from 6.5 mi. E of San Lucas, 300 ft., T. Paul Maslin, Apr. 9, 1974 (data from last 13 not included in tabulations herein). All of the type material is from 1979 LIZARDS OF THE SCELOPORUS ORCUTTI 5 the Cape Region south and east of the Isthmus of La Paz in the territory of Baja California Sur, Mexico. Other specimens examined. A total of 67, all from the Cape Region of Baja Califernia Sur, Mexico, was examined only by Hall: Agua Caliente, 800 ft. (CAS 46,790-1, MVZ 11,709-11); 2.7 mi. W of Agua Caliente, El Chorro (LACM 18,880-2); Rancho Buena Vista (LACM 18,883); 4 mi. SE of Rancho Buena Vista, Cerro Agua Amargo, 23°36’ N, 109°37’ W (MVZ 50,070); Boca de la Sierra (CAS 91,457, LACM 18,879, SDSNH 30,188-9): Cabo San Lucas (CAS 16,532-3, 46,800-6, 46,808, SDSNH 32,903); 0.3 mi. SW of Cabo San Lucas (CAS 91,446); 7.7 mi. NE of Cabo San Lucas (CAS 91,319-20); El Triunfo (CAS 46,779, 46,823, 46,826-8): Poe imi Ne of El Erunto (CAS 911,476): La Paz (CAS: 46.775. SDSNH 17,696-8); 10 mi S of La Paz (SDSNH 30,191-3); 12.4 mi. E of La Paz on road to Las Cruces (CAS 91,255): 12.9 mi. E of La Raz Arroyo de los Pozos (CAS 91367): 16 mi E of La Paz (SDSNH 30,186); 19 mi. N of La Paz[N of Pichilinque Bay] (CAS F095): 19°59 mi. SE 7. -= > : ae 4 ” = —— al RIV GW ds) TOKEN Se bi vaey DPT: 42 te ’ Vik Oza - Oa ay id Te = meee = a 2 _ _ = = 7 > aa = a - 7 ae a 2 ee ' : _ _ a : a eee = = @- bales i, < a) i - | ee if; er eas rk) - = SS = 2 = , = fe : - sia ~ a 7 _ | | | . | pS SY ee MAR 1 8 1985 imanrkVAKD UNIVERSITY. BREVIORA Museum of Comparative Loology US ISSN 0006-9698 CAMBRIDGE, MASS. 21 FEBRUARY 1979 NUMBER 453 THE LARGE PALAEOTRAGINE GIRAFFID, PALAEOTRAGUS GERMAINI, FROM LATE MIOCENE DEPOSITS OF LOTHAGAM HILL, KENYA C. S. CHURCHER! ABSTRACT. An isolated and slightly damaged left M! constitutes the first record of the species P. germaini in East Africa. This specimen matches in size and develop- ment of the buccal ribs and styles the upper molars of P. germaini from Oued el Hamman, Algeria. An isolated and damaged left upper molar, probably M! (KNM- LT 414, field no. 138/67K), was collected by Dr. V. J. Maglio from Lothagam Hill, Kenya, in 1967. The tooth was found at Lothagam-! near the base of Member B (Patterson et a/. 1970), and is deposited in the collections of the International Louis Leakey Memorial Institute for African Prehistory, Kenya National Museums, Nairobi. The specimen (Fig. 1) is very slightly worn on the occlusal surface and derives from an immature individual. Minor damage has removed the mesial face of the crown, including the mesial half of the protocone, but the paracone Is essentially entire. Slight damage to the distobuccal corner of the metacone has not destroyed the shape of the cusp or of the distobuccal wing of the hypoloph. Minor fragments of enamel are absent from the buccal surfaces of the apices of the paracone, metacone, and metastyle. The surface of the enamel is typically giraffid in its overall rugosity and thickness. The crown is tilted lingual on its roots in the anteroposterior (parasagit- tal) plane, and the roots tip distad. The mesial pair of roots is more damaged than the distal pair, all four roots are separate and are ‘Royal Ontario Museum and Department of Zoology, University of Toronto, Toronto, Ontario MSS IAI, Canada. No. 453 BREVIORA ‘SWID UT *] X 2[BIS “9[BOS SpIVMO] SUMOIO PUB sI9}}a] ISAIvIU S}001 YIM poiuso [Te oe q pue ‘d ‘OD ‘Vv ‘wedse eoong — q ‘joadse yenduyy — q ‘oadse [eisip — > ‘ayeos SPIPMO}J JOKJINS [BOING PUL Y SPIBMO} [RISA YUM JOOdse [esn[990 — g ‘joadse [eIsam — wy ‘eAUdy ‘[-WeIZeYIOT Woy IejowW puoosas 10 sayy taddn yay ‘1uipw4as snspsjoanjog —*| oin3i4 S341 3WILN39 oe S O 1979 PALAEOTRAGUS GERMAINI 3 round or oval in section. The exception is the distobuccal or metacone root which is grooved on its distal surface and crescentic in section. The cusps are typically giraffid with tapered protocone and hypocone set buccally from the lingual cingulum. The paracone and metacone similarly are set lingually towards the mid-line of the crown. The buccolingual diameter over the mesial cusps (paracone and protocohe) is greater than that over the distal cusps (metacone and hypocone). There is a well-developed median rib on the buccal surface of the metacone and a small pli is present distally in the postfossette on the wall of the hypoloph. A similarly located and formed pli, which is broken away, appears to have been present on the protoloph. The specimen is attributed to Palaeotragus germaini Arambourg 1959 because of its similar size (Table |) and because it matches well with Arambourg’s (1959, Pls. XIV, XV) illustrations of first and second upper molars of that species (Fig. 2, A & B) from Oued el Hammam. The specimen resembles P. germaini in 1) the presence of strongly developed parastyles and mesostyles and 2) a strong rib on the buccal surface of the metacone of M’, in the feeble junction of the paraloph and metaloph, and in the overlap of the mesostyle mesiobuccally to the distal end of the paraloph. Arambourg’s specimens possess interlophar endostyles or pillars between the bases of the protocones and hypocones of the first and second molars. The Lothagam-1! specimen lacks any such endostyle. How- ever, this structure is not always present in modern Giraffa camelopardalis (Fig. 2, F), Okapia johnstoni, or in fossil P. primaevus (Fig. 2, C; Churcher 1970, 1977). Therefore it cannot be considered a reliable diagnostic character. The specimen resembles P. primaevus in the presence of the metacone rib, and in the presence of plis on the buccal walis of the protocone and metacone. The specimen differs in that the ribs are more developed and the plis finer and directed mesiad rather than buccad as in P. germaini (Fig. Dea. SC) The Lothagam tooth expresses the other characters ascribed to P. germaini by Arambourg (1959, 100-101), but many of these char- acters (e.g.° strong cingula or deep fossettes) are common to Giraffidae and are probably not diagnostic for species or genus. The dimensions of the Lothagam P. germaini tooth fall within the ranges reported by Arambourg (1959) for M!’s of P. germaini from No. 453 BREVIORA O¢ (KE *«tP *«tP #8E «alt tp 8s «Sh JO OF «x7 JO Ep Or bE «€b 10 Op +0P 10 6€ 7W IW 7W IW (66 4 ‘6S6l Gils feu «Gg —d ‘Jinoquiviy ) ‘9Z6l “UNYoR) “BUIy) “BUIy) MADUINAU ASUBUIS UNTIYIOWUDSY LUNTMIYIOUDY OVC-LIZ CCCP 8 *0E x0 LC *0E +S LT CSCC Cte Ll 6l Chek CESS LE 0'8C- 97 O'ST-9'8I Shomts (4s LVSE=66 7W IW 7wW IW CHIT “QPL *€7 ‘d (ep “Sy :66 “d “OL61 “494younyD) ‘6S61 “dinoquiriy ) “eAUdYy “RLIasy ‘uBUIO] OY ‘WRUWeRH [9 pang snaanuid IUIDLUAID SNBDALOAD] Dg SNBDALOID] Dd SC LEG 99C Salts (PIP LT-WN®) eAuay ‘[-ueseylo 7] (1eamM Aq paonpal) 1ysloy-uMOID [RooNg auosodAy-auoorjaw Jo R[nsulo 19,0 JojawWeIp [ensulpooong auo0d0dAy-au0d0101d 1340 J3]9WIPIP [PISIPOISaP au0d0101d-au0dRiRed Jo e[nsulo 1340 Jojaweip [ensulooong auoodRjoW-auodeied 13A0 JOIWPIP [PISIPOISaP ‘WW Ul O1B S}UDWIINSPSU ITV “S¥Siaise Aq paieoipul are suUONeISN]]! WO] PaAlap sjusMaINseayy “[-WWeseYIO] Wo ejOW Joddn ay) Jo pur LtoUmau “§ puR asuauts UNIIYIOUDY ‘snAapouiid “q ‘IUIDWUdad SNBdAJOaD[D_ JO Siejow Joddn puodas pur isilj sy JO syudsWdINSvaW aANeIedWIOD ‘T 31921 1979 PALAEOTRAGUS GERMAINI 5 Oued el Hammam and marginally on or below the ranges for M?’s (Table I). In all instances, the measurements of the Lothagam tooth are greater than any reported for P. primaevus from Fort Ternan (Churcher, 1970), or for Samotherium sinense (Fig. 2, E) or S. neumayri reported by Bohlin (1926) and Arambourg (1959) respec- tively. (S. sinense is probably a junior synonym of S. neumayri according to Erdbrink, 1976). The Lothagam specimen is assigned to Palaeotragus germaini on the bases of both size and morphology. Palaeotragus germaini was originally described by Arambourg (1959) from Oued el Hammam or Bou Hanifa, Algeria. Churcher (1978) assigned the large palaeotragines from Beni Mallal, Morocco (Lavocat 1961), Bled ed Douarah, Tunisia (Robinson and Black 1974), and possibly Smendou, Algeria (Joleaud 1937) and Douaria, Tunisia (Roman and Solignac 1934) also to this taxon. These North African sites range from middle to late Miocene in age, with Smendou and Douaria extending into early Pliocene times. Arambourg (1959) described P. germaini as a large giraffid with elongate neck and legs, and with the forelimb slightly longer than the hind. The radius is longer than the tibia and the femur is short, the metapodials are subequal, long, slender and laterally compressed with subrectangular sections. Arambourg considered that P. ger- maini’s upper molars were characterized by strong buccal walls, interlophar endostyles, metacones separated from paracones, and dep fossettes that are usually open lingually. He considered the teeth to be strongly brachydont, but the tooth from Lothagam is not more brachydont than teeth of similar size from modern G. camelopardalis nor do his illustrations show teeth that are more brachydont than that from Lothagam. The ossicones were consid- ered to be long, supraorbitally placed, subtriangular in section and oriented posterodorsally over the middle region of the skull. The single ossicone known from Oued el Hamman is proportionately smaller than those of Samotherium and is similar to those of other species of Palaeotragus. Arambourg described the Oued el Hammal skeletal material as larger than that of other Pa/aeotragus species, similar in robustness to that of Samotherium, and intermediate in size between that of Samotherium and Giraffa. He considered that P. germaini exhibited a parallel evolution separate from Giraffa or Samotherium, and its lineage would thus be separate from those of the Giraffinae or No. 453 VIORA BRE “SIOJOWINUID Ul a[TeIS “BpPPUP “OJUOIO] “WiNasn, OLPIUG [PAO Y 9y) Ul UdUIOA|dS JUadA1 B WoL] UMBIP ‘syDpaDvdojaiuD? D{fD41D Jo _ W PI 4 pue (“sy c TIA JEL (9761) UNYOY Wosy UMBIPSI ‘asUoUIS UNLaYJOUIDgY JO “passaAdd *. ISU q(¢ ‘sy ‘77z “d ‘p6]) Aoyeay pue oiuinsy WO1J UMPIP “BAUDY “RYN WoO “ds wnMayIOWUDG JO ‘passaaai *_ 10 W 1ysu C ‘00L7 79/ LA usumoads wnasny jeuoneN PAUudy W YP SD) (el 231g jo yseo pue (¢ “BY “gy “d “Q26]) JoyoINYD Wooly UMPIPSI ‘PAUDY ‘URUIO] OY WOIy snaapiuid snspsjoan]Dg JO . t-1 “AIX ‘Id “6S61) SAINOGUIRIY, WoOl) UMPIPO “BLdIaa]y “WPLUWPH [9 pang WoO] IUIDLUAIS SNBDALOAD] Dd lore IW Vo] q seAUdy “UPB PIO ] WOl, IMIDLUAID SNBDALOIV]Dd JO Wi Vel V “oepysesly payoajas jo sivjow taddn puoosas pue isi jo susayed [esnpooQ °Z analy 1979 PALAEOTRAGUS GERMAINI 7 Sivatheriinae and would represent the more progressive and larger Palaeotraginae (Churcher 1978, Fig. 9). Arambourg assigned the Qued el Hammam giraffid to Palaeotragus on the proportions of the limbs, especially the elements of the hind limb. However, the characters of the molar teeth also place the taxon within the genus Palaeotragus rather than any other genus of the Giraffidae. P. germaini was known previously only from North African sites of middle to late Miocene age, or possibly early Pliocene age. The Lothagam-| late Miocene tooth represents the first East African record of the species. Some confusion may exist between some of the reported occurrences of Samotherium in Africa and P. germaini, especially when dental material is absent. The cervical vertebrae and two crushed ossicones from Fort Ternan that were assigned to Samotherium africanum by Churcher (1970) may in fact represent P. germaini or a form ancestral to it, since these ossicones are not excessively large and the single ossicone from Oued el Hammam cannot be considered necessarily typical. The identification of Samotherium by Aguirre and Leakey (1974) from the Nakali Tuffs of the Ngorora Formation, Kenya, suggests that both genera were present in East Africa during the late early to late Miocene. Their determination is founded on an isloated first or second upper molar (Fig. 2, D) which possesses a strong parastyle and mesostyle, a strong paracone rib, and reduced cingulum and metacone rib, which are all features of Samotherium. ACKNOWLEDGMENTS I wish to thank Charles L. Smart for sending the tooth to me and suggesting that I publish a description of it; Professor Bryan Patterson for allowing me to include mention of the specimen in my 1978 review of the African Giraffidae and for suggestions about this report; and Mary Lynn Richardson for advice and constructive criticism. The work was supported by grant A-1704 from the National Research Council of Canada. The Museum’s expeditions to Kenya were supported by National Science Foundation grants GP-1188 and GA-425. REFERENCES AGUIRRE, E. AND P. LEAKEY. 1974. Nakali, nueva fauna de Hipparion del Rift Valley de Kenya. Estud. Geol. 30: 219-227. 8 BREVIORA No. 453 ARAMBOURG. C. 1959. Vertebres continentaux de Miocene supérieur de l'Afrique du Nord. Mem. Serv. Carte Geol. Algerie, n.s., No. 4: 1-161. BOHLIN, B. 1926. Die Familie Giraffidae. Palaeont. Sinica, Ser. C, 4 (1): 1-178. CHURCHER, C. S. 1970. Two new Upper Miocene giraffids from Fort Ternan, Kenya, East Africa: Palaeotragus primaevus n. sp. and Samotherium africanum n. sp. Fossil Vert. Africa 2: 1-109. —____. 1978. Giraffidae, Chapter 25. /n Evolution of Mammals in Africa, V. J. Maglio and H. B. S. Cooke (eds.), Harvard Univ. Press; Cambridge, Mass., pp. 509-536. ERDBRINK, D. P. B. 1977. On the distribution in time and space of three giraffid genera with Turolian representatives at Maragheh in N.W. Iran. Proc. Koning. Nederl. Akad. Wetensch., Ser. B, 80 (5): 337-355. JOLEAUD, L. 1937. Remarques sur les giraffides fossiles d’Afrique. Mammalia 1: 85-96. LavocatT, R. 1961. Le gisement de vertebrés de Beni Mellal (Maroc). Etude systematique de la faune de mammiferes et conclusions geneérales. Notes Mém. Serv. Geol. Maroc 155: 29-94, 109-145. PATTERSON, B., A. K. BEHRENSMEYER AND W. D. SILL. 1970. Geology and fauna of a new Pliocene locality in northwest Kenya. Nature 226: 918-921. ROBINSON, P. AND C. C. BLACK. 1969. Note preliminaire sur les vertebres fossiles du Vindobonien (formation Beglia), du Bled Dourah, Gouvernorat de Gafsa, Tunisia. Notes Serv. Géol. Tunisia 31: 67-70. ROMAN, F. AND M. SOLIGNAC. 1934. Decouverte d’un gisement des mammiferes pontiens a Douaria (Tunisia Septentrionale). C. R. Acad. Sci. Paris 199: 1649 1650. | Th Pe MAR 1 8 1985 BRE V kv@noR A Museum of Comparative ZLoology US ISSN 0006-9698 CAMBRIDGE. Mass. JULY 31, 1979 NUMBER 454 EVOLUTION OF LIFE HISTORIES: A COMPARISON OF ANOLIS LIZARDS FROM MATCHED ISLAND AND MAINLAND HABITATS ROBIN M. ANDREWS! ABSTRACT. Faunal comparisons between islands in the West Indies and the South and Central American mainland suggest that Anolis lizards play very different trophic roles in these two regions. Island anoles, with few predators, appear to be food limited relative to the heavily preyed upon mainland species. Theoretical considerations of species living under different competitive regimes suggest that island and mainland anoles should have evolved major differences in their life histories. In order to evaluate the evolution of life history in the genus Anolis, | consider the degree of food limitation for anoles in comparable island and mainland habitats, the life history consequences of different selective regimes, and the correlation between observed life histories and theoretical expectations. Food availability on island and mainland sites was assessed from determinations of the standing crop of arthropods relative to the standing crop of anoles. A more direct assessment was accomplished by evaluating growth rates of Anolis juveniles. Both comparisons indicate that island anoles are food limited relative to mainland counterparts. Island anoles exhibit delayed maturity, long adult survivorship, large adult size, high sexual dimorphism ratios, and probably low reproductive effort in comparison with mainland anoles of similar size. For the most part, observed life histories fit the predictions based on current life history theory. However, for a given population, maximum body size, sexual dimorphism ratios, and offspring size are relatively invariant in expression. On the other hand, growth rate, rate of egg production, and age at sexual maturity vary as a function of food availability. INTRODUCTION In this paper I evaluate the effects of the competitive milieu on evolution of life history in the lizard genus Anolis. Various observa- tions suggest the island and mainland members of this large group ‘Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061. 2 BREVIORA No. 454 differ in the way that their populations are regulated. Anoles of the South and Central American mainland have numerous competitors for their insect prey and also have many predators; these anoles play minor roles in the middle levels of their complex food webs (Rand and Humphrey, 1968). On the other hand, anoles of the West Indies have few competitors or predators. There are fewer species of birds on these islands than in comparable mainland habitats and many types of insectivorous and lizard-eating birds are totally absent. These missing species include motmots, antbirds, woodhewers, ovenbirds, and puffbirds (Bond, 1971). Moreover, West Indian bird faunas do not exhibit density increases as a result of fewer species; density of insectivorous birds is lower in the West Indies than in comparable mainland habitats (Recher, 1970; Terborgh, pers. comm.). At a given West Indian site there tend to be fewer species of frogs (possible competitors for small insect prey) and snakes (possible predators on lizards) than at a comparable mainland site. The number of lizard species is also often lower than on the mainland (Rand, 1961; Schwartz and Thomas, 1975). Army ants which compete with anoles on the mainland have never reached the West Indies. It is this poverty of potential competitors that allows Williams (1972) to conclude that competition among West Indian anoles is, for all practical purposes, intrageneric. Trophic level differences suggest that island anoles, with few predators, may be food limited relative to the heavily preyed upon mainland species (Slobodkin et al. 1967; Wiegert and Owen, 1971). Recent theory predicts that such differences in mode of population regulation might have profound effects on Anolis life histories (MacArthur and Wilson, 1967; Gadgil and Bossert, 1970; Gadgil and Solbrig, 1972; Pianka, 1970, 1972). The effect of the competitive milieu on the allocation and utilization of energy result in the predictions listed in Table I. Heightened competition in saturated environments will either 1) reduce the overall amount of energy that an organism can gather per unit time or 2) force changes in the allocation of time and energy devoted to conflicting demands, or both will occur (Pianka, 1972). The consequences of this are important. The absolute amount of energy that can be used for reproduction is reduced. Thus a relatively low reproductive effort is predicted for populations in saturated environments. Moreover, selection will usually favor 1979 A COMPARISON OF ANOLIS LIZARDS 3 Table |. Predictions of relative differences in life history between island and mainland Anolis lizards. ISLAND ANOLES: MAINLAND ANOLES: Food Limited Not Food Limited 1. Reproductive effort Low High 2. Sexual maturity Late Early 3. Relative size of offspring Large Small 4. Size Large Small 5. Sexual dimorphism in size High Low individuals with attributes that make them successful in competitive situations. In Anolis, selection for large size can be expected to be particularly efficacious because larger individuals are usually domi- nant over smaller individuals in competition for space, which provides an advantage for obtaining food and mates (Rand, 1967: Trivers, 1976). Sexual dimorphism in size is a means by which competition for food between male and female Anolis is reduced (Schoener, 1967). High sexual dimorphism ratios are expected when competition is intense. Since juveniles fare poorly in competition with established adults, selection should favor long adult life with reproduction occurring in more than one breeding season (Murphy, 1968). Tinkle et al. (1970) predict that lizards living in saturated environments will have relatively large eggs and hatchlings because of the relationship between size and competitive ability. In sum, island anoles should be K-selected in the sense of MacArthur and Wilson (1967). On the other hand, if mainland anoles are generally limited by their predators, they should be r-selected relative to island anoles that are limited by food resources (Pianka, 1970, 1972). Because the risk of death is high for all age classes of mainland anoles, fitness is maximized by high reproductive effort. Attributes that maximize present reproduction will be favored by selection because the chance of future reproduction is small. Because age at first reproduction and size are closely and positively related (Fenchel, 1974), selection for early reproduction (which is the most effective way to increase r) will paralle] selection for small adult body size. In order to evaluate the evolution of life history in the genus Anolis, | have considered three main questions: Are island anoles food limited relative to mainland anoles? If so, what have been the 4 BREVIORA No. 454 life history consequences of the different selective regimes? Do the observed life histories conform to theoretical expectations? To evaluate the hypothesis that island anoles are food limited relative to mainland anoles, I first compare the standing crop biomass of Anolis predators and their arthropod prey in com- parable island and mainland habitats. A lower ratio of prey to predator biomass on islands than on the mainland would provide indirect evidence supporting the hypothesis. More direct evidence is sought in comparisons which involve the relative allocation of energy to feeding and social interactions as opposed to growth and reproductive effort. The hypothesis will also be tested with data on population turnover. Higher individual persistence in island versus mainland populations would provide evidence that predation pres- sure is lower in island than in mainland habitats. Next, various life history attributes of island and mainland anoles are contrasted to determine if island and mainland anoles differ in body size, age at sexual maturity, sexual dimorphism ratios, reproductive effort, and size of offspring. Finally, I discuss the evolution of life histories in Anolis. MATERIALS AND METHODS Comparisons of island and mainland Anolis are based on data that I collected from cacao plantations on Dominica (an island) during August 1971, March 1972, and July 1972 and in Costa Rica (mainland) during April and May 1971, August 1971, and February 1972. Cacao plantations were chosen because different localities have similar climates; cacao requires rainforest climates with high, evenly distributed rainfall. In such environments seasonal differ- ences in lizard reproduction are minimal (Licht and Gorman, 1970; Sexton et al., 1971) facilitating comparisons of different sites. Moreover, the closed canopy of a mature cacao plantation and the shade required by the trees themselves make the understory of a cacao plantation similar to that of an undisturbed rainforest (Leston, 1970). Sites and anoles studied The island site was located on the windward side of Dominica at Londonderry Estate 2 km N of the Melville Hall Airport at an elevation of about IS m. Weather stations in this area frequently 1979 A COMPARISON OF ANOLIS LIZARDS 5 average over 2500 mm of rain a year and even during the relatively dry period, mid-January to mid-March, rainfall is usually more than 100 mm a month (Beard, 1949; Hodge, 1954). Rainfall at Melville Hall Estate for August 1971 was 268 mm, for March 1972 was 265 mm, and for July 1972 was 108 mm (Christopher Maximae, pers. comm.). Dominica has a single species of anole, Anolis oculatus. On the basis of geographical variation in size and color pattern four subspecies have been described (Lazell, 1972). A. oculatus winstoni is found in lowland habitats on the windward side of the island. Subsequent references to A. ocu/atus will apply to this subspecies. Anolis oculatus are common in such diverse habitats as beach strand and undisturbed forest but appear to be most abundant in man-made “forests” such as cacao and orange groves. In cacao plantations A. oculatus occupy all structural components of the habitat, i.e. individuals are seen on the ground, on trunks, and in the canopy. The mainland study site was located in the Atlantic lowlands of Costa Rica about 45 km W of Limon (Limon Prov.) on Finca La Lola, the cacao experimental station of the Instituto Interamericano de Ciencias Agricolas (IICA, Turrilaba) at an elevation of 33 m. Average annual rainfall at La Lola for the years 1949-1967 was 3501 mm (Soria et al., 1969). There are two relatively dry periods a year (February-March and September), but rainfall rarely falls below 100 mm a month. Rainfall at La Lola for April 1971 was 266 mm, for May 1971 was 111 mm, for August 1971 was 106 mm, and for February 1972 was 332 mm (Alfredo Paredes, pers. comm.). Although seven species of anoles are recorded from nearby forests, only two are commonly found in the cacao plantations (Talbot, 1977). Of these, A. /imifrons is by far the most abundant, being found in all localities where shade is present. In the cacao, individuals are usually found less than 2 m above the ground but seldom on the ground itself. In contrast, the terrestrial A. humilis 1s restricted to older cacao stands where understory vegetation is sparse. Mark-recapture studies Mark-recapture studies, conducted during three visits over about a year, were made in mature cacao plantations characterized by a continuous canopy and sparse understory. In Dominica, the 625 m? 6 BREVIORA No. 454 study site included 43 regularly spaced cacao trees and six large breadfruit trees (Fig. 1). In Costa Rica, the 652 m2 study site (in Experimental Section 19 of Finca La Lola) included 45 regularly spaced cacao trees and two large Hura crepitans (Fig. 2). Insecti- cides or other chemical treatments were not used on the study sites during this study or for many years previously. During census periods, anoles were captured by hand and information on size, sex, individual identification number, and location were recorded in the field. Anoles were released shortly after capture where they were caught. Snout-vent length (SVL) was measured to the nearest mm for each capture and during some census periods weight (W) was determined to the nearest one-tenth of a gram (Pesola scales). Sex of adults and all but the smallest juveniles was readily distinguishable by the presence of the large dewlap in males only. Additionally, male A. oculatus of all sizes have enlarged post-anal scales which females lack. Individuals were identified by clipping the terminal phalange of toes in unique combinations involving not more than one toe. Population size was estimated from mark-recapture data. For A. oculatus, | estimated population size with the Peterson method using a normal approximation to place 95% confidence intervals around the mean (Seber, 1973). Census data for these estimates were collected on 17-20 March and 30 March-! April, 1972. For A. limifrons, population size was estimated with a regression procedure (Marten, 1970) using data of five census periods during April and May 1971]. Adult survivorship on the study sites was determined from the proportion of marked adults collected during the two site visits after the initial marking period. Diet analyses Anoles were collected? for stomach-content analysis in cacao plantations adjacent to and similar in vegetation structure to the mark-recapture sites. On Dominica, this collection was made on two days in August 1971, and in Costa Rica, on six days in early May 1971. Collections were made in late afternoon on rainless days so that all individuals had an equal time to feed. Individuals were ?These collections have been deposited in the Museum of Comparative Zoology, Harvard University. A COMPARISON OF ANOLIS LIZARDS 1979 No. 454 BREVIORA ‘ROLY BISOD Ul UONRURId OOP. UI ats Apnig 7 34 1979 A COMPARISON OF ANOLIS LIZARDS 9 collected by size in proportion to their occurrence in the popula- tions. Specimens were killed within an hour of capture and pre- served in alcohol. Stomach contents were analyzed using the following techniques. The length of each item found in the stomach proper was measured and the average width and depth were estimated to the nearest 0.1 mm with an ocular micrometer. The volume of each arthropod was determined by multiplying these three dimensions. Arthropods were assigned to one of 15 taxonomic categories (Table 2). Anolis body size and reproductive condition For each lizard in the above collections SVL and head length (HL) were measured. Head length was measured as the distance Table 2. Composition of arthropod collections made by sweeping understory vegetation on the Dominica and Costa Rica cacao sites. Values are percentages of total numbers in each collection and do not include collembola and mites. DOMINICA COSTA RICA cacao forest cacao forest foliage foliage foliage foliage Lepidoptera larvae 0.2 0.0 1.9 0.3 Lepidoptera adults 0.3 yal) 5.8 0.7 Orthoptera 1.9 0.5 D4) 4.] Blattidae 0.1 0.0 0.0 1.1 Hymenoptera: Formicidae 40.8 25) 10.3 21.4 Hymenoptera: Other 10.7 3P 18.5 12.1 Homoptera 5.5 5.8 10.6 6.4 Hemiptera 15 0.0 1.8 0.7 Coleoptera adults 12 23 133 1k Diptera adults 323 20.6 D3 Moll Holometabolous larvae 0.1 0.0 0.1 0.1 (except Lepidoptera) Araneida 4.5 11.8 565) 6.3 Isopoda 0.0 0.0 0.1 2.4 Isoptera 0.3 0.0 0.0 9.6 Miscellaneous 0.5 0.7 0.5 129 Sweeps 700 450 250 350 Total arthropods 3522 3180 1952 1219 No. 2 5 mm/1000 sweeps 243 44 972 526 10 BREVIORA No. 454 from the anterior edge of the ear opening to the tip of the snout. The sex of each specimen was confirmed by dissection and the repro- ductive condition of females was indexed by the number of oviducal eggs and the diameter of the largest ovarian follicle. The relation of head length and weight to SVL was determined by a least squares analysis. A single regression equation was appropri- ate for each of the three species because separate analyses for males and females did not alter the equation significantly (p > 0.05). Lizards were classified as adult male, subadult male, adult female, and juvenile. Females were considered adult (sexually mature) at SVLs not less than that of the smallest female with oviducal eggs. For species in which males are considerably larger than females, subadult males are equal in range of SVL to adult females (after Schoener, 1968). Subadult males are small, sexually mature males (Licht and Gorman, 1970; Sexton et al., 1971) that appear to be excluded from the breeding population by the larger adult males (Rand, 1967; Andrews, 197la) and thus form a separate class. Juveniles, defined as immature individuals, were sometimes divided into two groups: juvenile I and juvenile II, with smaller and larger individuals, respectively. The total live weight of anoles on each of the study areas was estimated using the following technique. | divided the A. limifrons collected in May 1971 and the A. oculatus captured in March 1972 into SVL categories separated by | mm increments. The appropriate regression equation was used to convert SVL to weight. Weight was then multiplied by the number of individuals in each SVL category and the results were summed within classes to give the total weight per class. These values were adjusted for the estimated number of individuals per class on the study areas. Behavioral studies Information on foraging behavior was collected through a series of standard 30-minute observation periods (e.g. focal-animal sam- pling, Altmann, 1975). Foraging behavior was typified by adult females because foraging is more clearly defined in females than in (adult) males (Andrews, 1971a). On Dominica, observations were made between 6 and 18 August 1971 on the capture-recapture area. On the Costa Rica site, observations were made between 29 April 1979 A COMPARISON OF ANOLIS LIZARDS 11 and 10 May 1971 on both the capture-recapture area and the collection area. Characteristics of arthropod communities Arthropod abundance, size distribution and taxonomic composi- tion on the cacao study sites and also in undisturbed forests nearby were determined from sweep samples of understory foliage. Sweep samples previde representative collections of the kinds of prey eaten by anoles that forage in this stratum (Andrews, 1971b). The limita- tions and assets of sweep sampling as an ecological technique are discussed in previous literature (Southwood, 1966; Janzen and Schoener, 1968; Janzen, 1973a). On the cacao sites, sweep collections for arthropods were made within two days of the collections of anoles and were taken on the same areas as the anole collections. The forest site on Dominica (Palmist Ridge) was located 5 km SW of the cacao site at an elevation of about 275 m. Rainfall is approximately 4000 mm/ y. This ridge top forest is extensive, with a high (max. 44 m) and fairly open canopy. The understory is sparse enough to walk through easily at most points. This site is further described by Soriano-Ressy et al. (1970). Sweep samples were taken in this forest in March 1972. The forest site in Costa Rica was located 2 km from Finca La Lola near the road between Limon and Siquirres and at a similar elevation to the cacao site. The forest area was relatively small (perhaps less than | km?) and was surrounded by cacao and areas recently clear cut. Trees had been selectively removed from the forest and therefore in some places the understory vegetation was dense enough to prohibit easy access. Sweep samples were taken in this forest on 29 February 1972. Methods used for sweep sampling are described by Janzen (1973a). The total number of sweeps on each site ranged from 250 to 700 with sets of 50 separated for statistical purposes. Arthropods were manually sorted from the leaves and detritus immediately after collection and then preserved in alcohol. Sampling was conducted only during sunny or intermittently cloudy conditions and at the same time of day (1000-1200 hours) on both sites. The same methods of taxonomic categorization and measurement of length were used for these arthropods as for the arthropods taken from the Anolis stomachs. However, dry weight in each length class was 12 BREVIORA No. 454 estimated using regression equations (Sage, 1974, Appendix Table 2) rather than from a volumetric determination. The following relationship for adult insects was used for most length classes: Ln Y = —7.761 + 0.3498X — 0.003931 X2 where Y = dry weight in grams and X = length in mm. For Lepidoptera larvae and Orthoptera of 15 mm and greater, I used equations specific for these taxa. RESULTS: AND DISCUSSION Prey Availability: Size, Abundance, and Taxa Size-frequency distributions of arthropods collected in cacao plantations show that large individuals were poorly represented in the island sample (Fig. 3). Only 4.8% of all individuals were 5 mm or greater in length on the Dominica site in contrast with 12.8% on the Costa Rica site (p < .001, Kolmogorov-Smirnov two-tailed test). This contrast between sites is accentuated when dry weight in each arthropod length class is plotted (Fig. 4) and further illustrated by summing total dry weights per 1000 sweeps, 6.4 g for Dominica and 17.8 g for Costa Rica. Moreover, significantly fewer arthropods were collected on the Dominica site than on the Costa Rica site (Table 2, p = 0.02, Mann-Whitney two-tailed test). The Dominica site averaged 247 arthropods per 50 sweeps and the Costa Rica site, 391 arthropods per 50 sweeps. Thus the low abundance of arthro- pods on the Dominica site 1s related to, in part, the low numbers of large individuals. The collection of arthropods from forest understories on Domin- ica had significantly fewer large arthropods than did the collection from the Costa Rica site (Table 2, p< .001, Kolmogorov-Smirnov two-tailed test). The island site also had a lower dry weight biomass of arthropods per 1000 sweeps than did the mainland site, 6.5 g as opposed to 8.1 g, respectively. However, the Dominica forest collection had significantly more individuals per 50 sweeps than did the Costa Rica forest collection (p = .02, Mann-Whitney U test, two-tailed). This is the result of large numbers of one very small myrmicine ant in the Dominica collection. The taxonomic composition of the arthropod collections on the two sites is contrasted in Table 2. Formicidae dominate the island collections (41-53%) with the one myrmicine species contributing respectively 27.4 and 46.7% of all arthropods collected from the 1979 A COMPARISON OF ANOLIS LIZARDS 13 4 a ! oe ] 3 A 5004 ie) a Sears | ww ‘ 1 8 a Lea, & 100 Sj £ '— (ret = 1 | Pg 50 | > ie 7) fae | ~ ! 2 aaa er i o = ' 2 t S , s 10 ' ' I 5 sg 2n IM a (im in ae | cy tt tie 't Ra Ween Lt ‘4 | ee rt iytine fer) | Ou 0 1 1 4 8 12 16 20 24 28 Arthropod Length (mm) Fig. 3. Size-frequency distributions for arthropods collected in understory foli- age in cacao plantations in Dominica (—) and La Lola, Costa Rica (----). Arthro- pods of 28 mm or more in length are not shown (2 in Dominica and | in Costa Rica). cacao and forest foliage. In contrast, Formicidae contributed only 10-21% of all arthropods from the mainland site. In general, large arthropods are well represented in sweep sample collections from mainland rainforests. On the Costa Rica site, 526 arthropods 5 mm or greater in length were collected per 1000 sweeps and this comprised 15.1% of all arthropods. Comparable figures are 14 BREVIORA No. 454 Dry Weight Arthropods / 1000 Sweeps 4 8 12 16 20 24 28 Arthropod Length(mm) Fig. 4. Dry-weight — frequency distributions for arthropods collected in under- story foliage in cacao plantations. Conventions as in the preceding figure. available for two other lowland rainforests in Costa Rica. In wet and dry season collections, | (Andrews, 1971b) found 510-525 arthropods 5 mm or greater in length per 1000 sweeps and these comprised 22.4—31.6% of all arthropods. In a dry season collection, Janzen and Schoener (1968) found 201 arthropods 5 mm or greater in length per 1000 sweeps and this comprised 25.2% of all arthro- pods. In comparison, very few large arthropods were collected on Dominica, the one island locality for which comparable data are available. I collected only 44 individuals 5 mm or more in length per 1000 sweeps and only 0.6% of all arthropods collected were in this size category (Table 2). 1979 A COMPARISON OF ANOLIS LIZARDS 15 More extensive comparisons of island and mainland arthropod communities have been made in second growth habitats. Allen et al. (1973) and Janzen (1973a, b) sweep sampled comparable sites on Puerto Rico and nearby smaller islands and on the mainland in Costa Rica. Both studies found a significantly lower abundance of arthropods in the island than in the mainland collections. Population Densities Anoles were two to three times more abundant on the island than on the mainland study site (Table 3). The total population size of A. oculatus was estimated at 299-363 individuals and of A. limifrons, 123 individuals. I assume that gains and losses to the adult populations during the census intervals were small because of their fidelity to established territories (Rand, 1967; Ruibal and Phili- bosian, 1974a; Andrews and Rand, unpub. mainland data) and because mortality would be slight during the census periods. The Peterson estimate for A. oculatus juveniles is probably reasonable because the census period spanned only two weeks. However, the regression technique used for A. limifrons adults gave unrealistically high values for 4. /imifrons juveniles. This was the result of the large Table 3. Population composition, density, and biomass of A. oculatus on the Dominica study site and A. /imifrons on the Costa Rica study site. Estimates for each class are followed by 95% confidence limits. Asterisked values for juveniles indicate the total number captured during the census period. Two totals for A. oculatus are given, one for each estimate of juvenile numbers. A. oculatus A. limifrons Site area (m2) 625 652 Number of Individuals Juveniles 170 (138-241) — 106* 46* Subadult males 44 (22-90) — Adult females 111 (80-198) — Adult males 38 (28-65) — Total adults — 77 (72-82) Total numbers 299-363 123 Total biomass (g) 1056-1121 96 16 BREVIORA No. 454 number of unmarked individuals that were captured during the seven week census period. The census period was long enough to allow recruitment from eggs. Moreover, because of a lower site fidelity of juveniles than adults (Rand, 1967), appreciable numbers of unmarked individuals may have moved into the study area and marked individuals moved away from it. A conservative estimate of juvenile population size is the total number of juveniles captured on each site during the census period (Table 3). Only 22 A. humilis were encountered on the mark-recapture area; the inclusion of its num- bers would increase the number of anoles on the mainland site very little. For similar habitats, the commonest species on islands character- istically have higher population densities than the commonest mainland species (Table 4). In non-forested and disturbed habitats, some island species become extremely abundant. For example, densities of 1400 and 2000 individuals per 1000 m2 are recorded for the grass anoles, A. poncensis and A. pulchellus, on Puerto Rico. Population densities of more than 400 individuals per 1000 m2 are typical of trunk-ground anoles in disturbed habitats (e.g. A. ocu- latus, A. lineatopus, A. acutus, A. sagrei). The island species listed with relatively low densities share habitats with species of equal or greater abundance. On Bimini, although A. distichus, A. carolinen- sis, and A. angusticeps are generally uncommon compared to A. sagrei, the range of combined densities on five sites where all four species were present was 197-935 individuals per 1000 m2? (Schoener and Schoener, unpub. data). Anolis lineatopus and A. opalinus had a combined density of 312 individuals per 1000 m? on Jamaica. In contrast to the island examples, the highest densities recorded for anoles in disturbed mainland habitats are all less than 200 individuals per 1000 m2. On Barro Colorado Island in Panama, A. auratus, a mainland counterpart of the West Indian grass anoles, had an estimated population density of 137 individuals per 1000 m2 (Andrews, unpub. data) even when population densities were unusually high (A. S. Rand, pers. comm.). Anole densities are lower in forests than in more open and disturbed habitats on both islands and on the mainland. On Abaco, Schoener and Schoener (unpub. data) estimate population densities for A. sagrei of 167-235 individuals/ 1000 m2 on forest sites and 439-969 individuals/ 1000 m2 on secondary sites over a two year period. Fleming (pers. comm.) found only six A. /imifrons per 1000 1979 A COMPARISON OF ANOLIS LIZARDS 17 m2 in forest plots in the Atlantic lowlands of Costa Rica, in contrast to my estimate of 189 individuals/ 1000 m2 in cacao plantations. Greater densities of some anoles in disturbed habitats compared with forested habitats has also been noted in Mexico by Henderson and Fitch (1975). Anolis density estimates on islands and on the mainland may be even more disparate than indicated in Table 4. Sparse populations are usually“not chosen for ecological study because of difficulty in obtaining sufficient data to accurately estimate population size. Since Anolis densities are generally lower on the mainland, main- land site selection is more likely to be biased towards high density situations. Population densities have not been determined for any South American anoles. Work by Rand and Humphrey (1968), Fitch et al. (1976) and Duellman (in press) indicate that the most conspicuous characteristic of Ano/is communities in this region is the extremely low densities of all species. An on-going study (Miyata, 1977) is being conducted, in part, in disturbed habitats where some species are encountered fairly frequently. It is obvious that low densities have generally discouraged ecological research on anoles in South America. Diets The island cacao site is characterized by low arthropod abun- dance, arthropod size distributions skewed towards small sizes, high frequencies of ants and a high density of Ano/is lizards relative to the mainland site. These differences in the prey community should be reflected in the diets of the island and mainland cacao anoles. Daily food intake of the cacao anoles was assessed from the quantities of food material found in their stomachs (Fig. 5). The two mainland species consistently had greater volumes of prey in their stomachs per gram body weight than did the island species. The differences were the greatest for the adult females of A. limifrons and A. humilis which had about twice the prey volume as did the similarly sized juveniles of A. oculatus. The greatest similarities were for adult males of the two mainland species which had mean prey volumes comparable to those of similarly sized A. oculatus. Metabolic rate, and thus energy requirements, are a function of temperature in lizards (Bennett and Dawson, 1976). Since A. oculatus and A. limifrons seem to have similar thermal regimes, No. 454 BREVIORA 18 OLI (€Z6] Usssuaf) eoreWeL ‘at[IAopuR|, qnios-ainjsed sndojvauy ‘y Trl (€L6| Uassusf) ROreWeL ‘a[IADpURY qnios-ainised snuyodo “py [EGeSS (SW ‘laus0yoISg pue Jous0YOS) lWIWIg SOUS SNOLIBA sdaosnsuv ab 69-81 (SW ‘au90Y49g puke JaUd0Y9S) TuUIWIg SOUS SNOLIBA SISUIUIJOADI “YY 601-6 (SW ‘1au30y9g puke Jaud0 Yas) TuTWIg SOUS SNOLIBA SNYIUSIP “Y S89-LL (SW ‘J9us0Y49g pur Jaua0YyoS) tulWIg Says SNOUPA 10ABDS “YY 696-L9I (SW ‘1ou20Y9S puke JIaUd0YIS) OoRGy SaUS SNOLIRA 10A3DS Orr (WpL6] UPISOqIIY puke [eqiny) xXIOID 1S yied SNINID “py 0007 (LL6| POOMIRH puke URWION) OoTY OVIING purysseis snyjayojnd “py 087-007 (OL61 ISI puke Jain] ) OdlYy olan, ISOIOJ 1YIDIPUNS “py 86E1 (196| puey) Oory olang yoied snjoro sisuaquod ‘py Srl (1961 Pury) OOrY Ong yied SNIJAIDISIAD 1601 (L96| Puey) eorewer deay ysniq sndojpauy “pv €LS—-99b (€ Iqe]) PoruUIWOg ororo SnIDINIO ‘py cW QO0T /'ON Al[e90 7] 1PVIQeH satoads SANV ISI “7 OOO] Jad Jaquinu 0} paliaAuod San[eA [PUIBIIO “SprezZi] SOUP PuRlUIeW PUR pUrISsI Jo sanisuap UoNRINdog “p aqry 19 A COMPARISON OF ANOLIS LIZARDS 1979 OOl-€1 OOI- eT 6 Stl 16l-v81 IpI-Ls SS 2 Il 681 (€L61 [leqdwe)) eureurg (€L6l [eqdwe)) eweurd (€L6] UOSyORL) SeINpUO H (ejep ‘qndun ‘smoipuy) ewReuRg (PEL61 YOU) BOY BISOD (REL61 YOU) BY PISOD (SL6] JOYOOH pure Burma) voy eISOD (eyep ‘qndun ‘surmoy4) Bory eBIsOD (eyep ‘qndun ‘surwoy.) voy eBISOD (L96] UOIxas “dad ul ‘sMoIpuy) PUeURg (elep ‘qndun ‘smoipuy) Bory B1ISOD (OL6| UASsUdL) ODTXI|J (€ 91GB L) BY BISOD CNV INIVIW adpo weds aspo weds purlpoom oud purysseid ysaioy Alaypes JUIARI PAPOOM S910} S910] S910] 1S910J S210] pur[poom yeo ovoRd sndopioaod ° snjouol * snjouopidod) SNIDAND ° snaddny °* snaddno * snaadno * SUNY * SUOA{ILUT] * SuOdfILUT] * sidajAjod ° snsojnqau ° Ssuodfiual] * TUX RG NG NG ONS 0.05). For A. limifrons, the classes listed in order of decreasing prey size are: adult female, juvenile I], adult males and juvenile I (p< 0.05 for all pairwise comparisons, K-S two-tailed tests). Thus, males took relatively small prey for their size. For A. humilis, males and females took prey of similar sizes but significantly larger prey than did the juvenile classes. Although qualitatively similar, the prey size utilization curves of the island and mainland anoles differ markedly when absolute prey sizes are considered. The sampled population of A. ocul/atus took significantly smaller prey than did the A. /imifrons population (p< .001, K-S two-tailed test). This difference is also illustrated from comparisons of lizard size classes (Fig. 9). The largest males were 22 BREVIORA No. 454 ADULT MALE (N= 28) Prey vol (mm) /individual = 2288 ~ Prey no /individual = 490 02 Ol aw O05 SUBADULT MALE (N=10) O04 , 68 Prey vol (mm) /individual = 141 5 Prey no /individual = 616 O3 WY a 02 WwW @ > Ol =) Zz > WwW a rat 04 ADULT FEMALE (N= 32) a M 76 Prey vol (mm3)/individual =1143 ro) O23 Prey no /individual = 604 WW = = 02 zi) SS Fel a BESS 6 =) 4 re O5 | JUVENILE Il (N=28) oO Prey vol (mm?) /individual = 49.2 = 04 Prey no /individual = 526 wL o O03 Zz La o2 f= a (S) on] Qa {e) a Se Pa eu 06 05 JUVENILE | (N= 20) ; \ Prey vol (mm?) /individual = 12.8 : Prey no /individual = 318 04 03 02 Ol O 4 8 \2 I6 20 PREY LENGTH (mm) Fig. 6. Prey utilization curves by the fraction of prey numbers in each prey- length class (----) and the fraction of prey volume in each prey-length class (—) for A. oculatus. Arrows indicate mean prey sizes for the volume distribution. 1979 PROPORTION OF TOTAL PREY VOLUME OR PREY NUMBERS A COMPARISON OF ANOLIS LIZARDS 23 025 I\ 020i 8.2 ADULT MALE (N=49) aes | Prey vol (mm*)/individual =22.9 O15 i \ Prey no /individual = 3.8 \ ADULT FEMALE (N=65) I Prey vol (mm®) /individual = 58.| i! Prey no./ individual = 4.0 I | i JUVENILE I! (N=61) O2O0r , \ Prey vol (mm®*)/individual = 20.2 yo 30 Prey no. /individual = 4.0 ei Nee O.5O- !1 JUVENILE | (N=33) ee 1 ae Prey vol (mm3) individual =6.5 (as) Prey no /individual =5.4 O20)- 7 eA \ O15 | O10F \ O05 Se NS Zz Or = 1 i = 4 8 12 16 20 24 PREY LENGTH (mm) Fig. 7. Prey utilization curves for A. limifrons. Conventions the same as for the preceding figure. 24 PROPORTION OF TOTAL PREY VOLUME OR PREY NUMBERS Fig. 8. BREVIORA No. 454 ADULT MALE (N=18) Prey vol (mm?>)/individual = 300 Prey no /individual=29 ADULT FEMALE (N=19) Prey vol (mm?) / individual =66 4 Prey no /individual = 75 JUVENILE Il (N=12) Prey vol (mm®*)/individual =165 Prey no /individual =63 JUVENILE | (N=24) Prey vol (mm®)/individual=7.8 Prey no /individual =55 8 16 20 24 PREY LENGTH (mm) Prey utilization curves for A. humilis. Conventions the same as Figure 6. 1979 A COMPARISON OF ANOLIS LIZARDS DS; the only A. oculatus that averaged larger prey than any A. /imifrons and this was a result of three large orthopterans eaten by one individual which strongly influenced the mean. Mean prey sizes for A. humilis are not plotted but are very similar to those of A. limifrons (compare class means in Figs. 7 and 8). However, in the comparison of the prey size distributions for the entire collection of both species, A. /imifrons was found to take significantly larger prey than did A. humilis (p< .001, K-S two-tailed test). Associated with the differences in prey sizes were the numbers of prey found per anole individual — 32-62 for A. oculatus as opposed to 4-S for A. limifrons and 3-8 for A. humilis. The three species also differed in the taxonomic composition of their diets. For A. oculatus, the most frequently eaten arthropods cl 5 4 ss al E i2f : emi Limifrons 55 © ? © lOk a ae | De rae j ye moll ae g ae ae 3 2 A= 5G um (On OMI MIZAIS ra SISNIGHI7aISMlIN2O HEAD LENGTH (mm) Fig. 9. Mean prey length for the volume distribution (see Fig. 6) as a function of head length for A. oculatus and A. limifrons. 26 BREVIORA No. 454 were Formicidae (ants) which contributed 44-57% of all items (Table 5). Formicidae, Isoptera, and miscellaneous (mostly psocids, collembola, and mites) arthropods were the most important con- tributors to prey volume although decreasingly so for the larger anoles with contributions of 77, 61, 58, and 39% for juveniles, adult females, subadult males, and adult males, respectively. For A. limifrons, nO One taxon was outstanding in frequency but Lepidop- tera larvae were the most important contributor to volume for all classes (27-44%) while Homoptera, spiders, and Orthoptera were of lesser importance (Table 6). For A. humilis, frequency of various taxa was more consistent, with holometaboulous larvae (mostly Diptera and Coleoptera), spiders, miscellaneous, and Isopoda being consistently highly ranked (Table 7). Again the most important contributor to volume was Lepidoptera larvae (25-28%). Stomach content analyses show that the diet of the island A. oculatus is made up of numerous small insects, primarily ants, and Table 5. Dietary composition of A. oculatus on the Dominica site in August 1971. Values shown are the decimal fraction of total numbers and volume (mm?) for each class. Adult Subadult Adult Juvenile Female Male Male No. Vol. No. Vol. No. Vol. No. Vol. Lepidoptera L. OO! O11 003 =.002 002 = .005 012 .088 Lepidoptera Ad. — — 001 .006 — — 001 001 Orthoptera .002 020 004 = .035 006 .025 005 = .300 Blattidae .002 .023 00S .041 002 .006 001 .002 Hymenoptera: Formicidae 460 .256 502 .244 570 .224 444 ~=.206 Hymenoptera: Other 036 =.011 008 ~=.001 010 = .005 009.042 Homoptera .006 = .006 006 = .010 005 ~=.007 005 001 Hemiptera 006 .004 004 .004 002 001 001 .001 Coleoptera Ad. 084 = .039 067 = .058 063 = .009 S91 02 Diptera Ad. 052.080 01S .144 014 .356 008 .067 Holometabolous L. .012 .020 19.006 013.005 005 .002 Araneida 033.019 012 .087 014 .005 0157 4005 Isopoda —_ — — — — —_— — — Isoptera OV 377 3° 2210 119.274 180 = .126 Misc. 229s ai35: 242) FAlid3 181 .078 124 056 Total N 2110 1934 616 1373 Total Volume 1633.8 3656.0 1415.2 6407.6 1979 A COMPARISON OF ANOLIS LIZARDS HY Table 6. Dietary composition of A. /imifrons on Costa Rica site in May 1971. Values shown are the decimal fraction of total numbers and volume (mm3) for each class. Adult Adult Juvenile Female Male No. Vol. No. Vol. No. Vol. Lepidoptera L. a3; 437 1237-273 lls) Silt} Lepidoptera Ad. 014 017 054 =.097 043.038 Orthoptera 002 .004 042 125 027.039 Blattidae 019 .066 034 =.120 005.001 Hymenoptera: Formicidae .074 = .008 .069 = .008 a09> 2033 Hymenoptera: Other 108 =.030 061 O11 038 .023 Homoptera 108 = .129 088 .076 147.090 Hemiptera 026 =.005 019 = =.004 005 =.003 Coleoptera Ad. 053.030 161 .078 125.084 Diptera Ad. 110 = =.056 050 = .024 125 .049 Holometabolous L. .026 = .030 004 =.001 Araneida 165 .099 M9 2056 .087 = .230 Isopoda 045.071 142 .076 O71 .064 Isoptera = = = = == = Misc. 096 017 034.051.038.028 Total N 418 258 184 Total Volume 1445.4 3776.1 1122.0 of the mainland 4. /imifrons of a few large insects, primarily Lepidopteran larvae. This dichotomy seems typical of most island and mainland anoles so far studied (Rand, 1967; Schoener, 1967, 1968; Schoener and Gorman, 1968; Andrews, 197la, b; Sexton et al., 1972; Lister, 1976; Scott et al., 1976). If the frequency of prey by length is considered, it seems generally true that the majority of the prey of island anoles (population or class) are less than 5 mm in length and the majority of the prey of mainland anoles are 5 mm in length or more. Ants are conspicuous in the diets of island anoles only. A “giant” anole, A. cuvieri of Puerto Rico, seems to be an exception (Rand and Andrews, 1975), but generalizations discussed here may not apply to such a large species. This problem is discussed in the Overview Section. Although intraspecific comparisons indicate that head and body dimensions of anoles are related to prey size (Schoener, 1968: Schoener and Gorman, 1968), comparisons between species, and 28 BREVIORA No. 454 Table 7. Dietary composition of A. humilis on the Costa Rica site in May 1971. Values shown are the decimal fraction of total numbers and volume (mm!) for each class. Adult Adult Juvenile Female Male No. Vol. No. Vol. No. Vol. Lepidoptera L. 091 = 281 085 = .276 320254 Lepidoptera Ad. 014 = .009 028 .090 019.070 Orthoptera 039.136 021 = .030 076 = .189 Blattidae 005.021 007 = .007 019 = .036 Hymenoptera: Formicidae 082 = .032 042 .008 094.039 Hymenoptera: Other 039.012 — —_— — — Homoptera 029.036 042 .087 057 = .037 Hemiptera 048. .014 —_—_ — _—->_ =— Coleoptera Ad. 048 = .028 014 .006 132 .042 Diptera Ad. 043.020 063," “al35 019 .001 Holometabolous L. 73" 2207 507.196 132.058 Araneida 149.039 092.026 094.089 Isopoda 101.142 070 = .137 3 eal27, Isoptera — — —_ _ _— — Mise: 139 028 = 028 001 3 ens Total N 208 142 53 Total Volume 386.2 1261.6 540.8 particularly between species that occupy different habitats, show that lizard size alone is a poor indicator of prey size. For example, most sizes of A. /imifrons eat larger prey than do A. oculatus even though A. oculatus is by far the larger species. Moreover, A. oculatus has a relatively longer head and greater weight per unit SVL than A. limifrons. There is apparently no direct functional relationship between the size and body proportions of anoles and the size of prey eaten. For A. oculatus, the linear relationship between lizard body size and mean prey size may result from interference competition where larger individuals monopolize the largest prey through social dominance over smaller individuals. Anolis activity patterns Both diets and population densities suggest that island anoles may have to spend more time and energy foraging and interacting with other individuals than do mainland anoles. If turnover rates and caloric values of stomach contents are comparable, then for 1979 A COMPARISON OF ANOLIS LIZARDS 29 similar energy requirements, the utilization of small prey will necessitate more active foraging by island than by mainland anoles. This prediction is borne out by results of observations on adult females on the cacao sites (Table 8). A. ocu/atus individuals were considerably more active than A. /imifrons during foraging as judged by the number of shifts in position and the number of movements that were directed towards potential prey. A. oculatus made about twice the number of major shifts in position (greater than one body length) than did A. /imifrons. A. oculatus was observed to attack prey about five times more frequently. The greater activity of A. oculatus was associated with a relatively shorter prey attack distance with 70% of the 60 prey capture attempts made at 10 cm or less as opposed to only 25% of the 8 such attempts for A. limifrons. Although A. oculatus females fed more on the ground than did A. /imifrons females (85% versus 38% of attacks), the success rate of both species was similar; about 80% of prey capture attempts resulted in capture. Where anole densities on islands are greater than on the main- land, the activities and associated energetic costs of social interac- tions will be greater for island than for mainland anoles. Social interactions in reptile populations appear to increase as a function of population density (Brattstrom, 1974). Presumably, increased density is associated with increased levels of interactions between males and females as well as increased levels of aggression between individuals of the same sex. Ruibal and Philibosian (1974b) found a linear increase in aggressive encounters with density for both male and female A. acutus. Brad Lister (pers. comm.) recorded the Table 8. Indices of foraging activity of adult female Anolis on the island and mainland cacao sites. A. oculatus A. limifrons n 18 12 Hours of observation 9 6 Position shifts/h De S Prey capture attempts/h 6.7 3 % successful attempts 82 1 Mean prey distance (cm) 19 30 % captures on ground 85 38 30 BREVIORA No. 454 activities of adult male A. sagrei in areas with densities of 13 and 50 individuals per 231 m? on Abaco Island in the Bahamas. He found that this approximately four-fold increase in male density was associated with a 15-20 fold increase per anole in such socially relevant activities as chases and displays (headbobs and dewlaps). High activity rates of island anoles may only be possible because of the relaxation of predation in island environments. With low risk of predation, island anoles maximize fitness by moving frequently to capture prey and to engage in social interactions with neighbors. In contrast, mainland anoles live in environments where a moving lizard has a relatively high probability of attracting the attention of a predator (Andrews, 1979). Mainland anoles maximize fitness by feeding infrequently on large, high yield items. Observations by Toft (1976) address this point. Diurnal members of frog communities in tropical rainforests exhibit one of two foraging patterns. The poisonous dendrobatids and some bufonids are active foragers, moving almost incessantly and eating very small items such as ants. In contrast, the palatable frogs (mostly lepto- dactylids) are sit-and-wait predators on relatively large arthropods. Their foraging behavior is much like that of the arboreal sit-and- wait anoles. The conspicuous behavior of dendrobatids and bufo- nids may be possible only because they are distasteful to most predators. Survivorship Survivorship of adult A. ocu/atus on the island site was consider- ably greater than of adult A. /imifrons on the mainland site (Table 9). Nine months after the original census of the 4. limifrons population, none of the males that were present on the site in May were recaptured and only one (6.6% of females captured) female was recaptured. In contrast, a high proportion of the originally marked A. oculatus were recaptured in following censuses. After 7 and 11 months respectively, 32.6% and 25.0% of males and 62.9% and 53.5% of females were recaptured. Although there are few long-term population studies on anoles, the evidence at hand suggests that adult survivorship is greater for island than for mainland anoles. Annual adult survivorship of A. acutus on St. Croix ranged from 27-40% for females and 0-19% for males (Ruibal and Philibosian, 1974a). Schoener and Schoener (1978) found annual survivorships up to 20-50% for some classes of 1979 A COMPARISON OF ANOLIS LIZARDS 31 A. sagrei, A. distichus, and A. angusticeps on Bimini and Andros. Several populations of A. opalinus also had many adult individuals that lived for more than one year (T. A. Jenssen, pers. comm.). On the other hand, both capture-recapture data and seasonal changes in population composition suggest complete annual turnover of adults for mainland A. polylepis (Andrews, 1971b), A. limifrons in Costa Rica (Table 9; Fitch, 1973b), A. limifrons in Panama (Andrews and Rand, in prep.; Sexton et al., 1963), A. nebulosus (Jenssen, 1970), A. cupreus (Fitch, 1973a; Fleming and Hooker, 1975), and other mainland anoles (Fitch, 1973b). Size. Relationships The island and mainland cacao anoles are markedly different in size (Table 10). A. oculatus is large and exhibits a pronounced sexual dimorphism in adult maximum size. Males reached 79 mm SVL or about 11 g and females 63 mm SVL or about 5 g. In contrast, A. limifrons and A. humilis are small with slight sexual dimorphism in size. Female A. /imifrons reached 41 mm SVL and female A. humilis 40 mm SVL at weights of about |.3 g and males were a few mm less in SVL. This difference in size and in degree of sexual dimorphism is typical of the differences between island and mainland anoles. Sexual dimorphism ratios (SDR’s) of Greater Antillean and mainland species are contrasted in Figure 10. Species size is indexed by maximum SVL of males and females (Williams, 1972). In this comparison, the number of sympatric species (as many as 6-7) are similar in both the island and mainland faunas. This comparison is Table 9. Survivorship of adult anoles on cacao sites. For A. oculatus this included all individuals > 48 mm SVL. A. oculatus A. limifrons No. adults initially marked 444 23 ¢ 85 9 32 9 Time elapsed 7 months 3 months Total captured (% marked) 46 4 (32.6) 18 ¢ (55.6) 62 9 (62.9) 17 9 (47.1) Time elapsed 11 months 9 months Total captured (% marked) 44 3 (25.0) 18 4 (0.0) 71 9 (53.5) 1S 9 (6.6) No. 454 BREVIORA (66°) TAS Ip@ + IST IT = 1H ce OF Le STM AY, (76°) --2TAS 870000° = A (L6°) TAS EZ + 996° = TH ce Ip 6€ suodfiuu “y (86°) ¢¢1AS ~Z0000' = AA (66°) TAS 77 + 097° =H 8P £9 6L S717 0119 OY suonenby uolssaisay ‘WY *x9aS 5 2) TASS} TAS wnuixepy sa1oads ‘(sasayjuaied ul suonenba smoyoj 71) stskteue sarenbs Isea] Aq paulwaiap suoNenbs uolssaisay ‘sued ul AY pue WU UI TAS pue TH yum (TAS) YISUs] JUSA-]nNOUS Jo UOTIOUNY eB SB UDATS o1e (AY) 1YSIOM pue (TH) YSU] peay ‘Apnjs sty ut suonejndod syoup ayy Jo sdiysuoneal azis Apog ‘QT aqey 1979 A COMPARISON OF ANOLIS LIZARDS 33 18 14 WV oo ® 10 rot (Te) = ) fe} Z 6 2 1.0 1:2 1.4 1.6 18 Sexual Dimorphism Ratio Fig. 10. Sexual dimorphism (male maximum SVL/female maximum SVL) ratios for Greater Antillean (solid line) and mainland (South and Central America) (dashed line) Anolis. Data for the Greater Antillean species are from Schoener (unpublished measurement tables, only species with a sample size of 15 or more individuals were used). Data for mainland species are given in Table 12. also a conservative test of the prediction that island anoles will have higher SDR’s than mainland anoles (Table 1) because of lower SDR’s in the Greater Antilles than on the one- and two-species islands of the Lesser Antilles (Schoener, 1977). Island species are found to have significantly greater sexual dimorphism ratios than do mainland species (p < .001, 2 X 2 Chi- squared test, Siegal 1956). Fitch (1976) has also measured SDR’s of mainland Anolis. Excluding those species for which I present data, the range of SDR’s (0.80-1.36) and mean SDR (1.07) for the tropical anoles in his data set are very similar to mine (Fig. 10) and further support the prediction of relatively high sexual dimorphism ratios for island anoles. 34 BREVIORA No. 454 An even more relevant analysis is suggested by Schoener’s (unpublished MS) demonstration that ecomorphs, which are species occupying similar structural habitats, are also similar in their sexual dimorphism ratios. Although 6-7 ecomorphs have been described (Rand and Williams, 1969), the present discussion is restricted to species whose structural habitat includes shrub level vegetation. Thus plotting the SDR’s of bush-grass and trunk-ground anoles as a function of maximum male SVL shows that island and mainland species form exclusive groups (Fig. 11). For any given male size, the island bush-grass species have higher sexual dimorphism ratios than mainland counterparts. Furthermore, the island trunk-ground spe- cies have greater SDR’s than all but one mainland trunk-ground species. Thus, even when similar ecomorphs are compared, the island anoles have greater SDR’s than mainland species. Comparisons of size are difficult since there is a wide range of size of different species as indexed by maximum male size both on islands and on the mainland. In the Greater Antilles the range in species size increases with the number of species per island, and on Cuba, the most extreme case, the range is from 38-191 mm (Williams, 1972). Although Cuba has by far the largest Anolis, the range of species sizes in the Greater Antilles is generally similar to that for mainland species where the range is from 38 to about 150 mm (Williams, pers. comm.). On the one- and two-species islands of the Lesser Antilles the range in species sizes 1s smaller, from 47-128 mm; extreme sizes are absent. Thus the range in species size 1s related to species richness; the greater range for Greater Antillean than for mainland anoles is possibly due to ecological opportunities that are not present on the mainland (Williams, 1972). To avoid this problem I compare the relative abundance of various sized species within Anolis communities on islands and on the mainland. Relatively large species are common and frequently dominant members of anole communities in the West Indies. On single species islands in the Lesser Antilles maximum male SVL’s tend to range between 65 and 95 mm (Williams, 1972). On islands with two species, one tends to be larger and the other smaller than the average for solitary anoles (Williams, 1972). However, both members of these species pairs are frequently co-dominants in the habitats where they occur together (G. Gorman, pers. comm.). There are many A COMPARISON OF ANOLIS LIZARDS 1979 “‘suoleoo] pasodxa pue ysty ATaanejas ul yosod AypeodAy yorym safoue punoss-yuNI) Jsow uvy) snonoidsuoo ssay yonwi st jy “(e1ep ‘qndun ‘smarpuy ‘‘wuW0d ‘siad ‘YydI14) $3011 JO aseq ay} 1B Sassada1 Yep Ul saydsad AJUOWWOD II ‘aJouR puNnos3-yuNI & st satoads sty] YSNoyY ** UB AQ pajoU SI snULNWa] “PY ‘OT “BI Ul SB sadinos ee ‘suoneindod ajdwes om) pry yotyM saroads purjulew InoJ ay} 10J payoyd s19M Sones wWstydiowlp [enxas yIog ‘sydrowrosa punoi3-yuni) pue sseid-ysng 10J TAS WNwWIxeW aJeW JO UONOUN] B sB passoidxa sonnei Wstydiowip [enxag “[] “Sly (ww) yyBuay] yuaA-yNous ajDW WNnuIxDW OSI ool 08 09 4 01 Za la Va 4 fs L is 4 Daly ace ee 5 [ 9 3 © r al Ee x L =| r.°] el punoi6-yuny &W \ @ Ze ; o is y sspi6 -ysng Vv Me oP 9 Ll VOIWIWV HLINOS 8 IWyLNIO =e punoib- yuna @ ssoiB-ysng © SITNLNY Yd1Vvado : 81 36 BREVIORA No. 454 species in the Greater Antilles with maximum male sizes of 55 mm SVL or less (Schoener, unpub. measurement tables). Despite this, small bush-grass anoles and large trunk-ground anoles that are sympatric may have comparable densities (Schoener and Schoener, 1971la, b; Jenssen, 1973) or large trunk-ground anoles may have greater densities than sympatric bush-grass anoles (Turner and Gist, 1970; Rand, 1967). In contrast to the island situation, Ano/is communities on the mainland are either dominated by small species or are comprised entirely of species with low population densities. According to Fitch et al. (1976): “Throughout Mexico and Central America one or another of the seven species mentioned [A. humilis, A. limifrons, A. cupreus, A. sericeus, A. nebulosus, A. subocularis, and A. tropi- donotus] tend to dominate habitats suitable for anoles”. To his list I would add A. po/lylepis, the dominant in southwestern Costa Rica. All of these species are small, with males not exceeding 55 mm in maximum SVL. Furthermore, all species occupy shrub-level struc- tural habitats. Large congeners are rarely encountered relative to the small dominants. Examples of large species with low population densities are A. frenatus in Panama (Andrews and Rand, unpub- lished data), A. capito in Costa Rica (Andrews, 1971b), A. attenu- atus and A. woodi in Costa Rica (Fitch et al., 1976), and A. biporcatus and A. pentaprion in Costa Rica and Panama (Andrews, unpublished data; Fitch, unpublished data). In South America, the species most frequently encountered may be relatively large in size. It appears that the structure of Anolis communities in South America may be quite different than in Central America and Mexico, but because all species are infre- quently encountered, much more information is necessary before such a generalization is possible. Because of the high abundance of large species on islands, comparisons of population density (Table 4) provide a minimal contrast between Ano/is community structure on islands and on the mainland. For example, the total live weight of A. oculatus on the island study was estimated at 1094 g (including 173 juveniles) and the total live weight of A. /imifrons on the mainland site at 96 g (Table 3). The 2~3 fold difference in numbers becomes a ten-fold difference in biomass. 1979 A COMPARISON OF ANOLIS LIZARDS 37 Reproduction (females) For lowland Anolis species, reproductive activity is associated with rainfall (Licht and Gorman, 1970). Judging from studies on sites with rainfall regimes similar to those of this study, reproduc- tion probably continues year round for A. oculatus and A. limifrons. Possibly some dimunition occurs in the number of females with oviducal eggs during January-March (Licht and Gorman, 1970; Fitch, 1973c). Data on reproductive condition was collected during times of the year that both A. oculatus and A. limifrons females should have been producing eggs (August 1971 and April-May 1971, respectively). The majority of adult females on the cacao study sites were reproductive as measured by the presence of oviducal eggs (Table 11). A greater proportion of female A. /imifrons were considered reproductive by this criterion than female A. oculatus (p < .01, 2 X 2 Chi-squared test, Siegal, 1956). Using reproductive condition as an index of the rate of egg production (Licht and Gorman, 1970), the data of Table 11 suggest that the rate of egg production by A. /imifrons females might be greater than for the A. oculatus females. A general comparison of reproductive effort or proportion of energy budgets allocated to reproduction (Hirshfield and Tinkle, Table 11. Reproductive condition of female Anolis in study populations expressed as the number of individuals with oviducal eggs. Repro. class Number of % females (no. of eggs) Females with eggs A. oculatus 0 13 Aug. 1971 l D2 63 Dominica 2 0 A. limifrons 0 6 May 1971 l 53 91 Costa Rica 2 6 A. humilis 0 4 May 1971 l 13 80 Costa Rica 2 2 38 BREVIORA No. 454 1975) for island and mainland anoles is not now possible. Such comparisons require information on the rate at which eggs are produced during the breeding season as well as their caloric worth. Rates of egg production under field conditions are known for only two tropical species. A. aeneus on Grenada laid eggs at intervals of about 12 days (Stamps, 1975) and A. /imifrons (both Panamanian and Costa Rican populations) at intervals of 7-8 days (Andrews and Rand, 1974). Although reproductive condition through the annual cycle has been determined for many anoles (Licht and Gorman, 1970; Sexton et al., 1971; Andrews, 1971b; Gorman and Licht, 1974; Sexton and Brown, 1977), such data cannot be used to index the rate at which eggs are laid unless environmental conditions are com- parable. One confounding factor is that anoles retain eggs under dry conditions (Stamps, 1976). Thus, two oviducal eggs may indicate retention rather than high rates of production. A second influence is the year to year variation among species in fat accumulation during dry periods and its utilization for egg production during the breeding season (Licht and Gorman, 1970; Fleming and Hooker, 1975). Fat storage would facilitate high rates of egg production over short periods. Both of the above factors make the interpretation of reproductive condition data difficult. Associated energy costs of re- production, such as territorial defence or preparing egg laying sites, are even more poorly known than the direct costs of eggs per se. OVERVIEW We are now in a position to evaluate the evolution of life histories in island and mainland Anolis. A critical portion of the information that has been reviewed concerns present-day conditions in these two environments. The expectation that island and mainland anoles will differ in life histories is based on supposed differences in the way that their populations are regulated. Therefore, I first consider the evidence that island anoles are food limited relative to mainland anoles. Determinations of arthropod and Anolis abundance suggest that food availability is less on islands than on the mainland. My Dominica site had a ten-fold greater live weight of anoles but about one-third the dry weight of arthropods than the Costa Rican site had. Although I measured standing crop, measurements of site 1979 A COMPARISON OF ANOLIS LIZARDS 39 productivity for arthropods would presumably enhance the differ- ence between sites. Assuming the rate of population turnover is roughly a function of individual size, production of small arthro- pods would be roughly comparable on both sites due to the similar representation of arthropods < 8 mm in length (Figs. 3 and 4). Production of large arthropods would be greater on the mainland site because such individuals were almost absent in the island collection. Limited arthropod data from other islands support my findings (Janzen, 1973b). More conclusively, the evidence for high anole densities on islands is well documented (Table 4) and would be strengthened by use of biomass rather than numbers of individu- als because of the relatively large size of common island anoles. The sizes and kinds of arthropods in anole diets suggest that island anoles eat more non-preferred items than do mainland anoles. Like many generalized predators, anoles prefer large items to small ones (Andrews, 1971b, unpub. data; Schoener, 1969a); the number of large prey eaten should be some function of their availability to a feeding lizard. Small items, particularly ants, are relatively more abundant in island than mainland habitats and more abundant in the diets of island than mainland anoles as well. The large numbers of prey in the stomachs of island anoles is a necessary consequence of small average prey sizes. Thus, a comparison of diets suggest that while mainland anoles have the “option” of feeding on large, high yield prey items, island anoles do not. The above comparisons constitute indirect evidence that anoles on islands are food limited relative to anoles on the mainland. In addition, comparisons of processes that are potentially energy limited would provide more direct evidence in support of this hypothesis. The growth of juveniles is such a process and 1s, in fact, considerably slower for island than mainland species. For com- parable lowland habitats, island juveniles grew at one-half to one- third the rate of mainland juveniles (Andrews, 1976). The low growth rates of island anoles may reflect low food availability or a relatively high allocation of energy to foraging and social activities or both. The data at hand suggest that island anoles are food limited relative to mainland anoles. The data on survivorship are in accord. Mainland anoles have greater population turnover than island anoles, presumably reflecting the greater predation intensity in 40 BREVIORA No. 454 mainland than island habitats. The hypothesized dichotomy in selective regimes for island and mainland anoles is strongly sup- ported. The consequences of such differences are predictable, at least in theory. I now review the life histories of island and mainland Anolis. First, is age at maturity later for island than mainland anoles? Island females were estimated to reach sexual maturity in 5-9 months as opposed to 2-4 months for mainland females (Andrews, 1976). Comparing species with similar female size, A. lineatopus (Jamaica) becomes sexually mature in 156 days and A. polylepis (Costa Rica) in 100 days. However, in Anolis, size at sexual maturity is a linear function of maximum size (Andrews and Rand, 1974). Thus, the later maturity of island than mainland anoles must be a function of their lower growth rates. Moreover the low growth rates of island anoles are facultative; with augmented food, rates of growth may be more than doubled (Andrews, 1976). This means that age of maturity in female Anolis on islands will vary with food availability. Two observations support the prediction that reproductive effort is lower for island than for mainland anoles. First, less energy 1s available for the growth of juveniles on islands than on the mainland. Energy may likewise limit reproduction for island fe- males. On the other hand, island females have the potential to lay eggs at rates comparable to those of mainland females (Andrews and Rand, 1974). Second, hatchling size is primarily a function of female size (Andrews and Rand, 1974). Island females do not have larger hatchlings than mainland females which may be related to constraints of the arboreal life styles of anoles. However, the slope of the linear function relating hatchling weight to female weight is considerably less than one. Therefore, proportionally, the caloric investment per offspring (ignoring associated costs) will be consid- erably less for large than for small species. For comparisons in which island species have larger females than mainland species, relative investment per offspring would be lower for island females even if rates of egg production in the two groups were the same. High SDR’s are characteristic of West Indian Anolis. The male is larger in all cases. However, if sexual dimorphism is solely a function of the need to partition food by size (Schoener, 1967, 1968; Schoener and Gorman, 1968), it should not matter which sex is the 1979 A COMPARISON OF ANOLIS LIZARDS 4] larger. On the mainland, the situation is quite different. For 21 of the 59 populations measured by Fitch (1976) and myself, females were larger than males. One explanation for the larger size of males than females is sexual selection. Trivers (1972) points out that when females are clumped in space but dispersed in time by asynchronous breeding that one male can monopolize breeding. The more females potentially monopolized, the more intense will be male-male competition for this resource. These assertions are particularly applicable to Anolis because: 1) adult males are often associated with more than one female (Rand, 1967; Andrews 1971a), 2) females potentially mate just prior to the ovulation of each egg (Crews, 1973), and 3) male size is positively related to reproductive success (Trivers, 1972). Because of the potential for males to monopolize many females, sexual selection should be particularly effective in insular situations where Anolis densities may be high. In addition to sexual selection, factors that promote fitness in females may also explain the larger size of males than females in the West Indies. Because size at first reproduction is linearly related to maximum size (Andrews and Rand, 1974) for the same growth rates, the smaller the species size the sooner females can lay eggs. If we ignore other factors, early maturity should be advantageous because age at first maturity is such an important contributor to Inax (Cole, 1954). Furthermore, decrease in female size lowers metabolic requirements faster than it lowers offspring size. For example, reduction of female weight from 4 g to 2 g reduces standard energy requirements by about 40% but offspring weight by only about 20%. In highly competitive situations, it may be advantageous for females to have low energy demands and yet be able to produce relatively large offspring. Thus female size may be a compromise between low energy demands and early reproduction and the ability to produce competitive offspring. This might explain Schoener’s (1969b) obser- vation that there is less variation in median female size than in median male size between islands differing in number of congeners. The prediction that mainland anoles will be smaller in size than island anoles is supported by community level comparisons; in mainland habitats common species are generally small and in island habitats common species are both small and large. I have argued that small size is one of a suite of r-selected life history attributes 42 BREVIORA No. 454 adaptive in mainland Anolis habitats where predator pressure is high. However, large species do occur in these habitats. The large mainland anoles appear to fall into one of two ecological categories, each minimizing the risk of predation in different ways. The very low densities of many large species such as A. capito and A. frenatus are associated with dietary specialization on large prey items. The mean length of 62 prey items eaten by 23 adult A. capito was 16 mm when calculated by prey numbers or 30 mm when calculated by prey volume (Andrews, 1971b). Adult 4. frenatus eat similarly sized prey (Scott et al., 1976; Andrews, unpub. data). Individuals of 15 mm in length or more constitute less than 0.4% of arthropods taken by sweep sampling at lowland rainforest sites in Central America (Andrews, 1971b; Janzen and Schoener, 1968, Guapiles site). Therefore large anoles that specialize on rare re- sources must themselves be rare. Although small arthropods are much more abundant than large ones, large anoles are probably large prey specialists because the low foraging rates are adaptive in their rainforest habitats (Andrews, 1979). The tendency of the large trunk-ground species such as A. capito, A. frenatus, and A. woodi to “freeze” rather than to flee when disturbed (Fitch, pers. comm.; Andrews, 1971b) also suggests that they avoid detection by preda- tors by long periods of immobility and highly cryptic coloration. The mainland anoles that are both large and relatively abundant seem to be adapted to specialized habitats where predator pressure may be relatively low. The “aquatic” anoles have maximum male sizes ranging from 70-100 mm SVL (Table 12, Fitch 1976). Camp- bell (1973) found densities of A. poecilopus and A. lionotus up to 100 individuals per 1000 m? in Panama. Escape from avian pred- ators may be relatively easy in the boulder strewn, stream side habitats where these anoles live. The “rock” anoles, with maximum male SVLs of about 80 mm, are locally abundant in Mexico (Fitch and Henderson, 1976). Here also, predator pressure may be low. In contrast, the overriding determinant of Anolis life histories on islands is competition for food. Large size is one of the K-selected attributes which maximize fitness in situations where resources are limited. The contention by Case (1978) that the increased size of territorial lizards on islands is a function of a greater availability of food resources is not supported by the Anolis data. Both insect 1979 A COMPARISON OF ANOLIS LIZARDS 43 standing crops and the growth rates of juveniles in comparable island and mainland habitats indicate that the ratio of resource supply to resource demand by anole populations is relatively low on islands. Although food availability may provide one constraint to the maximization of body size in island habitats, it does not explain the relative increase in size of island anoles. The dichotomy in the life histories exhibited by island and mainland anoles generally conforms to the initial predictions (Table 1). However, life history theory implies that life history attributes are invariant expressions of the genome. It is obvious from exam- ination of Anolis life history attributes that while some attributes are relatively constant in expression, others are extremely plastic. In Anolis, attributes such as maximum size, sexual dimorphism ratios, and offspring size have easily definable values for a given popula- tion. These indicate selection for fitness under environmental conditions that are “constant” across many generations. On the other hand, growth rate, age at sexual maturity, and rate of egg production are highly variable under experimental conditions and presumably in the field as well. There has been a recent recognition that such variability may be highly adaptive in heterogeneous or fluctuating environments (Hickman, 1975; Hirshfield and Tinkle, 1975: Nichols et al., 1976). The ability of the food limited island anoles to exploit temporally or spatially abundant food by increas- ing rates of growth or egg production is not surprising considering the importance of early maturity and reproductive effort to individ- ual fitness. The life history model proposed here must be qualified in one important respect. It was developed by focusing on the major selective forces acting on island and mainland Anolis that live in lowland tropical environments. Although the available data support the predicted island-mainland dichotomy, only two anole eco- morphs, the bush-grass and the trunk-ground species, have con- tributed to these data. The anoles we know the least about, very large species, canopy species, and species that live in extreme environments, are also the most likely to prove exceptions to the model (Andrews, 1976). The trophic position of the very large anole species may very well differ from small species since many are frugivorous (Brach, 1976; Schoener, unpub. data) and some include No. 454 aspo weoi}s-onenbe (eueued) €/6] []eqdweD 60'1 OL 9L snjouol ‘py punoi3-ysnq (Bory BISOD) Z IIQVL 6 Ip 6E punoi3-ysnq (eueued) [6] ‘1B 19 UOIKIS 001 OS OS suodfiuy *Y (oolxayw “eiep ‘qndun) yoy °S “H 00°1 6L 6L sSnuLinue “Vy punoi3-yunJ} (edly BISOD ‘eIep ‘qndun) yoy ‘Ss “HY L6 09 8s 7SNuLuNniua] “YY punoi3-ysnq (Bory BISOD ‘eIep ‘qndun) yoy ‘S ‘Hy 00'I vs vs snipausajul ‘py punoid (BOY BISOD) Z FIQVL 6 OP 8E SUIuINY “VY puno3-ysnq (Jopensg “wwod ‘siad) uewT[ang ‘qA 16 Lv €p snipanvoosn{ ‘py punoi3-yunJ} (eureued) 9/6] [B19 09S SZ OZI OSI snipuadf s 80° OP Os ° punois-ysnq (Bory BISOD) Z7L6I ‘TP 19 YOU aia Lv Ss \snaddno “y > 83° 08 0L Z punoi3 (WY *S) 061 SWRI Pue tulfozURA £01 L9 69 \S1dajosdayo “py a = (eoty BISOD) IL6T TP 19 A]aYoq t6 bP Iv lualuadivo “y punoi3-yun4} (Bory BISOD ‘BIep ‘[qndun) smoipuy 16 L6 88 old) "py Adoues (eweueg ‘“WWwOd ‘siad) puey 06° L6 L8 snjp2410diq "vy a3pa Weoaijs-oNenbe (ODIXIP) 8961 JAIN 67 I BL 101 lUaydDg “Y ssei3 (eueued) [6] [B19 UOIKIS 00'I IS IS SNIDIND ‘py fo) 2 yeIQqeY [eINjoONIyS 301n0S yds TAS wnuixey, saioads ‘SyOUp UROLIIWY [e1UID Puke YINOS Jo sones WstydIoWIp [eNxas puke sazIg “Z] IQR 44 45 ‘sojoue UBILIOWY [ejuag Jo AWOUOXe} dy} JO 9}R]S PasNJUOD PUR S9dUdIIJJIP IZIS dy JO asnedaq saloads JOUNSIP PalapIsUOD 1B SnULINUAa, OMI BY Iz ‘UDAIS DIV SYS 9UII9x9 10} asoy} A[UO ‘suoNRindod AuewW 10j JIQRI[TeAe SUONRUIWIII9P YS, A COMPARISON OF ANOLIS LIZARDS 1979 (Bory BISOD) 1L61 SIOAW 86° LS 9S SUDLAfIION “Y puno3-ysnq (1openog “wuw0d ‘siad) uewTfang “A AM 16° 9S IS DUAIPAYIDAL “YY punoi3-ysnq (seinpuoH) €/6] UOSyoerr 90'I TAS 6S snjouopidoa] “y (Bory BISOD) 7L61 YOU cO'l 8S 6S sidajopidodl “y punoi3-ysnq (eueUued) [6] ‘|B 19 UOIXIS 00'I IS IS dajspsopidod] “py ({) yunn (oorxaW “eiep ‘qndun) yoily °S “H v0'l 8P OS SNaIdas “py youriq-31M} (Jopenog “wwod ‘siod) uewyjang “qi 90'I LL Z8 snjpjound ‘py punod-ysnq (Roly BISOD) BIL6] SMaIpUYy Ol 8h €¢ sidajdjod "y a3pa weais-onenbe (eureued) €/6] [][aqdwed 80'1 99 IL sndojivaod ‘py ysnq (aopenog “wwood ‘siod) uewyjang “qs 96° 0S 8p 1u0}40 ‘fy punoi3-ysnq : (ejanzaua,A “WUO0d ‘siad) puey ‘Ss ‘Vv O€' | L9 L8 DIUO 'Y puno.3-ysnq (OSIX9W) OL6] Uassuar L0'l Or 6p snsojnqau ‘py 46 BREVIORA No. 454 other anoles in their diets (Rand and Andrews, 1975; O. J. Sexton, pers. comm.). Furthermore, the environment may place physiologi- cal constraints on anoles in some habitats that override the biotic factors considered here. As examples, A. onca, which occupies xeric coastal regions of Venezuela, has a very high SDR for a mainland anole (Table 12), and the low growth rates of A. tropidolepis in Costa Rica may be a function of low environmental temperatures in its cloud forest habitat (Fitch, 1972). Whether or not a distinct island-mainland dichotomy can be extended to such “atypical” species is yet to be resolved. ACKNOWLEDGMENTS This study was partially funded by NSF Grant Nos. B0O19801X and GB-37731X to E. E. Williams. Research was conducted during tenure as a Smithsonian postdoctoral fellow and a visiting research associateship at the Smithsonian Tropical Research Institute. I would like to thank G. Gorman, L. Houck, A. R. Kiester, E. G. Leigh, A. S. Rand, T. W. Schoener, E. E. Williams, and one anonymous reviewer for useful comments on the many versions of this paper. For assistance in Dominica, I am grateful to Christopher Maximae of the Forestry Office for information and logistic support and to Frobel Laville for permission to work on London- derry Estate. In Costa Rica, IICA also provided logistic support for field work. The kindness and assistance of Alfredo Paredes and Albertina Johnson at La Lola was most appreciated. LITERATURE CITED ALLAN, J. D., L. W. BARNTHOUSE, R. A. PRESTBYE, ANDD. R. STRONG. 1973. On foliage arthropod communities of Puerto Rican second growth vegetation. Ecology, 54: 628-632. ALTMANN, J. 1975. Observational study of behavior: sampling methods. Be- haviour, 29: 227-267. ANDREWS, R. M. 197la. Structural habitat and time budget of a tropical Anolis lizard. Ecology, 52: 262-270. ANDREWS, R. M. 1971b. Food resource utilization in some tropical lizards. Ph.D. Thesis, Univ. of Kansas, Lawrence. ANDREWS, R. M. 1976. 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WIEGERT, R. G. AND D. F. Owen. 1971. Trophic structure, available resources, and population density in terrestrial vs. aquatic ecosystems. J. Theoret. Biol., 30: 69-81. WILLIAMS, E. E. 1972. The origin of faunas. Evolution of lizard congeners in a complex island fauna: A trial analysis. Evol. Biol., 6: 47-90. aa & ea 7 ae at 2 jane 4 ‘ + le if kesh . Ss oo 6s eu ty: * 2 Rem Pe eds = ‘ Ay Pm) ced MUS. COMP. ZOOL LIBRARY B R k; \ le Rk A EL A UNIVERSITY Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, MASS. JULY 31, 1979 NUMBER 455 A NEW PALEOCENE PALAEANODONT AND THE ORIGIN OF THE METACHEIROMYIDAE (MAMMALIA) KENNETH D. ROSE! ABSTRACT. A new genus and species of palaeanodont, Propalaeanodon schaffi, from the late Paleocene (late Tiffanian) Polecat Bench Formation of the Bighorn Basin, Wyoming, is the oldest and most primitive known metacheiromyid. Pro- palaeanodon foreshadows Palaeanodon in the reduction of its postcanine teeth. It resembles epoicotheriids in the presence of postcanines along the entire length of the horizontal ramus, and resembles the slightly older Amelotabes, in particular, in the retention of seven postcanines. Propalaeanodon thus provides additional evidence for uniting the Metacheiromyidae and the Epoicotheriidae in the suborder Palaean- odonta, and it strongly suggests that the Metacheiromyidae evolved from the Epoicotheriidae, probably during the Tiffanian. Two humeri from the same stratigraphic level as the holotype are tentatively referred to Propalaeanodon. They possess specializations typical of Eocene palaean- odonts, including a prominent deltopectoral crest and a large supinator crest. These features indicate that Propalaeanodon was, already in the late Tiffanian, a highly specialized fossorial animal. The systematic position of the Palaeanodonta is uncertain. They have been considered to be related to the orders Xenarthra or Pholidota or both by all recent students and, indeed, they are the only known fossil group that evolved specializa- tions similar to those in these two orders. Although it is possible that palaeanodonts were an early radiation of probably myrmecophagous, fossorial mammals merely convergent to these orders, the evidence now available enhances the probability that the three groups share a special relationship. INTRODUCTION The edentate-like members of the rare mammalian suborder Palaeanodonta have long been considered early relatives of the Xenarthra, the Pholidota, or both. As now known, they are restricted to the earlier Tertiary of western North America. Palaean- odonts are classified in two families, the Epoicotheriidae and the 'Museum of Paleontology, University of Michigan, Ann Arbor, Michigan 48109. 2 BREVIORA No. 455 Metacheiromyidae. To date, nine genera and fifteen species have been recognized, several of which were described quite recently (West, 1973; Rose et al., 1977; Rose, 1978). They range in age from Tiffanian (late Paleocene) to Orellan (middle Oligocene). Recently the dentary of a primitive new palaeanodont was discovered by Charles Schaff, Museum of Comparative Zoology, in late Tiffanian rocks in the northern Bighorn Basin, Wyoming. The oldest and most primitive metacheiromyid, it sheds new light on the origin of this peculiar family and further documents the unity of the Palaeanodonta. Two palaeanodont humeri, collected much earlier from the same stratigraphic interval, probably represent the same new species. Abbreviations of institutional names cited herein are as follows: AC Pratt Museum, Amherst College, Amherst, Massachusetts. AMNH_ American Museum of Natural History, New York. MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. PU Princeton University Museum, Princeton, New Jersey. UM_ University of Michigan Museum of Paleontology, Ann Arbor, Michigan. SYSTEMATIC PALEONTOLOGY Suborder PALAEANODONTA Matthew, 1918 Family METACHEIROMYIDAE Wortman, 1903 PROPALAEANODON, gen. nov. Type species. Propalaeanodon schaffi, sp. nov. Included species. Type only. Distribution. Late Paleocene (late Tiffanian) o1 the Bighorn Basin, Wyoming. Diagnosis. Only metacheiromyid with seven postcanine teeth that extend to the back of the horizontal ramus. All postcanines single- rooted except P,, which is two-rooted. Postcanines small, peglike, separated by short diastemata, and with rounded crowns, in these features resembling Palaeanodon; but crowns higher and more pointed than in Palaeanodon. Dentary slender, slightly smaller than in Palaeanodon ignavus; deepest beneath first postcanine (P)), shallowest beneath last postcanine (M3), as in Pa/laeanodon. Medial buttress prominent but less so than in Palaeanodon. 1979 PALEOCENE PALAEANODONT 3} Etymology. In allusion to its resemblance to, lower stratigraphic level than, and probable relationship to Palaeanodon. PROPALAEANODON SCHAFFI, sp. nov. Fig. 1A and B, 2A, and 3B Holotype: MCZ 20122, left dentary with P23, alveoli for P), Ps, M, 3, and posterior margin of canine alveolus. Hypodigm. Holotype and tentatively PU nos. 13928, 13929. Horizon and Locality. Latest Paleocene (late Tiffanian), “Silver Coulee beds” of Polecat Bench Formation (of Jepsen, 1940), Jepsen Valley Quarry, NW, Sec. 27, T.57N., R.100W., Park County, Wyoming. ject Figure 1. Lateral (A) and dorsal (B) views of holotype of Propalaeanodon schaffi, MCZ 20122, left dentary with second and third postcanines (P2_;). Dorsal view of left dentary of Palaeanodon sp. (C), showing extended edentulous part of ramus behind fifth postcanine; reconstructed from UM 63611 and 66243. 4 BREVIORA No. 455 c Figure 2. Medial views of left dentaries of palaeanodonts. (A) Propalaeanodon schaffi, holotype, MCZ 20122. (B) Palaeanodon sp., reconstruction based on UM 63611 and 66243. (C) Amelotabes simpsoni, holotype, PU 14855 (right dentary, reversed). Diagnosis. Only known species of the genus; measurements in Table: I: Etymology. For Charles R. Schaff, expert collector and prepara- tor, who discovered the holotype. Description. Only two teeth are preserved in the holotype, although the dentary is nearly complete and provides much in- formation about the mandibular morphology of Propalaeanodon schaffi. The horizontal ramus is intact as far forward as the posterior margin of the canine alveolus. This alveolar margin reveals that the canine was large, as in other palaeanodonts, and its 1979 PALEOCENE PALAEANODONT 5) root robust, a feature also indicated by thickening of the dentary anteriorly. The dentary contained seven postcanine teeth, presumably four premolars and three molars, which were greatly modified from the primitive eutherian condition. Seven is a greater number of teeth than in any other metacheiromyid, indeed more than in any other palaeanodont except Amelotabes, which also has seven (Rose, 1978). Alocodontulum, in which the lower dentition is unknown, has seven upper postcanines (Rose et al., 1977, 1978), and thus probably had seven lowers as well. The postcanines of Propalaean- odon occur along the entire length of the horizontal ramus, as in Figure 3. Left humeri of palaeanodonts. Scale is 10 mm. (A) Early Eocene metacheiromyid Palaeanodon ignavus, based on AMNH 15088 and 16832; after Matthew (1918), but with corrected orientation of proximal end. (B) ?Propalaeano- don schaffi, PU 13928. (C) Early Eocene epoicotheriid Pentapassalus woodi, part of holotype, AC 2766. Note that this species is referable to Pentapassalus (Rose, 1978), not to Palaeanodon (Guthrie, 1967; Emry, 1970). The supinator crests in (A) and (C) are incomplete and have not been reconstructed because of the variable develop- ment of this crest in aifferent taxa of palaeanodonts; they were in any case larger than depicted. 6 BREVIORA No. 455 TABLE | Dimensions of the holotype of Propalaeanodon schaffi, MCZ 20122. (Measurements are in mm to nearest .05 mm) Length Breadth P, alveolus 1E35 0.85 P2 alveolus damaged P) crown 1225 1.00 P; alveolus 2.40 1.15 P; crown 1.80 1.10 P; alveolus 2.80 1.10 M, alveolus 2.00 eS M> alveolus 2.00 1.15 M; alveolus 1.60 1.00 P,-M;3 17.10 (approx.) Depth of mandible, lingually at P;: 6.10 (approx.) Depth of mandible, lingually at M;: 4.80 epoicotheriids but not in later metacheiromyids, in which the back of the horizontal ramus is edentulous. All teeth in Propalaeanodon are single-rooted except Ps, which has two roots (probably a primitive feature). Judging from the alveoli, M; and Mb», and possibly P3;, had single roots that were faintly bilobate. The roots of all postcanines were tapered and closed, as in Palaeanodon. The epoicotheriids Amelotabes (Fig. 2C) and Tubulodon are similar in having a two-rooted P,, but in these taxa several other postcanines are also two-rooted. In Pentapassalus M, has a bilobate root but P4 is single-rooted (Gazin, 1952). The postcanines in Palaeanodon are all single-rooted (Fig, IC, 2B). The crowns of P2 and P; are preserved. P; has become dislodged from its alveolus as a result of damage to the front of the jaw and appears higher than P;. Both teeth are peglike with rounded crowns apparently devoid of enamel. In these characters they closely resemble the postcanines of Palaeanodon (Fig. 1C, 2B and Rose, 1978), although they are slightly larger, particularly longer, higher crowned, and less rounded. As in Palaeanodon they show no cusp pattern on the crowns, nor is there any evidence of direct occlusion with the upper teeth. Occlusal wear is discernible in molars and even in premolars of some epoicotheriids. The alveolus of P, indicates 1979 PALEOCENE PALAEANODONT 7/ that it was the largest cheek tooth, a feature in common with Amelotabes but in contrast to Palaeanodon. There is a slight size reduction from P, to M3. All postcanines are separated by short diastemata. At the alveolar margin, the alveoli of P24 slightly exceed the size of the roots they contain; this probably obtained for the molars as well. These traits are characteristic also of Palaeanodon, some advanced epoicoth- erlids, and many toothed xenarthrans. The ascending ramus and condyle of Propalaeanodon resemble those of Pentapassalus pearcei and Metacheiromys tatusia (Gazin, 1952; Simpson, 1931). The coronoid process is slightly smaller than in contemporary proteutherians (sensu Butler, 1972) and rises at an angle of about 110° to the ventral border of the mandible. The condyle, which is narrower transversely than in most toothed xenarthrans and contemporary proteutherians, is situated relatively low but above the level of the tooth row. Its convex articular surface is directed dorsally and backward, as in P. pearcei. The mandibular foramen is also relatively low, being situated on the posterior pro- jection of the tooth row. The internal mandibular groove extends from just anterior to this foramen to a point beneath M;. As in all metacheiromyids, there is a prominent medial buttress, forming anteriorly a low, narrow ridge lingual to the molars. The dorsal surface of the buttress is roughened, particularly behind M3. On the external surface of the mandible the masseteric fossa, though rather poorly defined, is deeper than in most other palaeanodonts. The fossa is bounded in front by a crest formed by the thickened anterior border of the coronoid. A more rounded ridge running forward from the condyle forms its lower limit. The horizontal ramus 1s rather slender and shallow, deeper in front than in back, as in Palaeanodon. A small mental foramen is present beneath P3. Two palaeanodont humeri from Princeton Quarry are probably referable to Propalaeanodon schaffi (Princeton Quarry is about 4 mile northwest of Jepsen Valley Quarry and at about the same stratigraphic level). They are the only other palaeanodont specimens from the same stratigraphic interval as the holotype. PU 13928, a complete left humerus (Fig. 3B), and PU 13929, a distal end of a left humerus, are of appropriate size for Propalaeanodon. They are smaller than the humeri of Palaeanodon ignavus (Fig. 3A) and Pentapassalus woodi (Fig. 3C and Rose, 1978) and slightly larger 8 BREVIORA No. 455 TABLE 2 Dimensions (mm.) of humeri in certain palaeanodonts. (Known humeri of Palaeanodon are too incomplete to allow accurate measurements.) ? Propalaeanodon schaffi, PU 13928 dasypus, AM 11718 pearcei (from Gazin, 1952) Pentapassalus AC 2766 Metacheiromys Pentapassalus woodi, length 39.3 74.0 37.0 47.1 length of deltopectoral crest 26.3 54.7 28.0 35.5 (from proximal end of greater tuberosity) length of supinator crest 1957 38.8 22.0 20.8a (from distal end of humerus) greatest transverse breadth 19.8 34.0 DEF 22.6 of distal end of humerus a = approximate than the humeri of Pentapassalus pearcei (Gazin, 1952). However, all these specimens are essentially similar in structure (see Table 2 for comparative dimensions). The deltopectoral crest is broad and well developed in PU 13928. It may be relatively slightly longer than in Palaeanodon, but this is not certain because the known humeri of the latter are fragmentary and poorly preserved (AMNH nos. 15088, 16831, 16832). The crest is relatively shorter than in Pentapassalus and Metacheiromys (see Simpson, 1931: 340-342). Its distal end is developed into an anteromedially directed prominence for the insertion of the pecto- ralis major. Proximolaterally the crest gives rise to a low process for the insertion of the deltoid. A less well-defined crest extends distally from the lesser tuberosity almost to the entepicondylar foramen and bears a small teres tubercle midway along the shaft, on which the teres major inserted, as in Metacheiromys dasypus (Simpson, 1931). The supinator crest is somewhat smaller than in Pentapassalus pearcei and Metacheiromys dasypus and lacks the hooklike, dor- sally projecting flange characteristic of the latter two species. 1979 PALEOCENE PALAEANODONT 9 Nonetheless, this crest is very large and served as the site of origin for the powerful supinator muscle and carpal extensors, and probably the brachioradialis. The median epicondyle (the point of origin of the pronator teres and the carpal flexors) is also large, although smaller than in Palaeanodon and Pentapassalus. These features of the humerus indicate an animal highly adapted for fossorial habits (Reed, 1954; Hildebrand, 1974). Although the development of the deltopectoral crest in palaeanodonts is (as Emry, 1970, asserts) more like that of manids than of armadillos, the development of the supinator crest is more extreme, even in PU 13928, than in either manids or xenarthrans. DISCUSSION Propalaeanodon is the second known Paleocene palaeanodont and the only known Paleocene metacheiromyid. A number of resemblances to epoicotheriids were described above — e.g., seven postcanines, teeth present at the back of the horizontal ramus, two- rooted P; — but these are primitive features of palaeanodonts. Propalaeanodon resembles Palaeanodon (Fig. 1 and 2) in the manner of reduction of its postcanines to peglike teeth (all but one single-rooted) with rounded crowns that apparently lack enamel. Like Palaeanodon, its postcanine teeth are housed in tapered alveoli that are larger than the roots and separated by diastemata. Al- though some of these features also evolved in advanced epoicothe- riids (e.g., Bridgerian Tetrapassalus mckennai, Chadronian-?Orel- lan Xenocranium pileorivale; Simpson, 1959; Colbert, 1942), the total suite of features evolved much earlier in the metacheiromyids. Thus when metacheiromyids had already achieved these traits, contemporary epoicotheriids, as now understood, retained rela- tively unreduced teeth with enamel and with very short or no diastemata between them. Tetrapassalus and Xenocranium are more specialized than Propalaeanodon in having fewer teeth and a shortened dentary, but they are more primitive (and typically epoicotheriid) in showing occlusal wear on the teeth. As metacheiromyids reduced the number of postcanine teeth, they retained a long dentary, hence leaving the back of the horizontal ramus edentulous. The trend in the Epoicotheriidae was toward shortening the dentary as teeth were lost. The antiquity and primitive grade of Propalaeanodon results in resemblances to both 10 BREVIORA No. 455 metacheiromyids and certain epoicotheriids and highlights the extent of parallelism that has occurred within the Palaeanodonta. Propalaeanodon is one of those rare fossils that is morphologi- cally intermediate between two well defined families, and it strength- ens the alliance of the Epoicotheriidae and the Metacheiromyidae in the suborder Palaeanodonta. Because of its very early acquisition of advanced traits found also in Palaeanodon and its stratigraphic and geographic proximity to Palaeanodon, | believe Propalaeanodon is best regarded as a primitive metacheiromyid, probably lying in or very near the ancestry of Palaeanodon. Its very late Paleocene age makes derivation of the Metacheiromyidae from an epoicotheriid similar to early late Tiffanian Amelotabes highly probable. More- over, the primitive aspect of Propalaeanodon suggests that meta- cheiromyids did not become distinct from epoicotheriids before the Tiffanian. The humeri tentatively referred to Propalaeanodon possess spe- cializations characteristic of fossorial mammals (see Simpson, 1931 and Rose, 1978 for additional functional interpretations). They are of interest in demonstrating that the humerus was already highly specialized in this late Paleocene palaeanodont. The deltopectoral crest is very similar to that in Palaeanodon. The supinator crest may have been slightly larger than in Palaeanodon, but no specimen of Palaeanodon preserves a complete supinator crest. AFFINITIES OF THE PALAEANODONTA Resemblances of palaeanodonts to Xenarthra and Pholidota have been enumerated by several authors (Matthew, 1918; Simpson, 1931; Emry, 1970). Two questions arise from these comparisons: To what extent are these features demonstrably synapomorphic with one or the other of these orders? To what extent do they reflect parallelism or convergence? Detailed comparisons led Matthew (1918) to conclude that the Palaeanodonta are definitely related to the Xenarthra and probably to the Pholidota as well. He found no morphologic obstacles to direct descent of both orders from Palaeanodon, but for geographic and faunal reasons he deemed this origin of Xenarthra unlikely. He speculated that “early Tertiary faunas of South America . . . must be derived apparently from some late Cretacic fauna, unknown to us 1979 PALEOCENE PALAEANODONT 11 but presumably inhabiting some part of North America” (Matthew, 1918: 653). Simpson (1931) detailed many characters to support alliance of the Metacheiromyidae and the Xenarthra, but he concluded that they are not so closely related to the Pholidota. Nearly four decades later, Emry (1970) critically reviewed Simpson’s criteria, presenting arguments that he believed vitiated them. In Emry’s view, the Metacheiromyidae were directly ancestral to the Manidae, and he included both in the order Pholidota. He rejected any special relationship between palaeanodonts and Xenarthra. Emry therefore discontinued formal recognition of the suborder Palaeanodonta, questioning whether epoicotheriid-metacheiromyid ties were really closer than metacheiromyid-manid relationship. A primary concern of these and other authors has been that known palaeanodonts were too specialized too early in the record to be direct ancestors of xenarthrans, since contemporary and dif- ferently specialized xenarthrans were known from South America (e.g., the early Eocene dasypodid Utaetus). In particular, tooth reduction was regarded as “prematurely specialized.” The discovery of a primitive epoicotheriid (Rose, 1978) and now a primitive metacheiromyid, both from the late Paleocene and both with much less reduced dentitions, may obviate this objection. Meager evidence attests to the presence of dasypodids in the Riochican of Patagonia (Simpson, 1948), and these are possibly contemporaneous with the Paleocene palaeanodonts. However, while the age of the Riochican is usually given as late Paleocene, its precise correlation with North American land mammal ages is uncertain (Simpson, 1978). At least part of the Riochican is considered by some authors to be younger than late Paleocene, i.e. equivalent to the Clarkforkian and early Wasatchian (e.g., Marshall et al., 1977). It is known that the teeth of early xenarthrans had enamel and developed gabled wear surfaces (Simpson, 1932). Therefore Propalaeanodon itself, whose post- canine teeth apparently had already lost enamel and did not develop wear facets, may have been too specialized to be directly ancestral to the Xenarthra (but not Pholidota). These objections do not apply to Amelotabes, and nothing now known about it precludes it from possible direct ancestry to Xenarthra or Pholidota. Some post- cranial features, sich as the extreme development of the supinator crest in palaeanodont humeri, are more specialized than in any known xenarthrans or pholidotans. However, some members of both groups possess an expanded, though smaller, supinator crest. 12 BREVIORA No. 455 If known palaeanodonts are ancestral to either Xenarthra or Pholidota, secondary reduction of the supinator crest must have occurred. This would not be required, however, if either or both orders were derived from more primitive palaeanodonts in which the supinator crest was less developed. Paleogeographic evidence must also be considered. There was no land connection between North and South America in the late Cretaceous or in the Paleocene (Freeland and Dietz, 1971; Malfait and Dinkelman, 1972). Therefore if palaeanodonts gave rise to xenarthrans, they could only have reached South America by rafting (Patterson and Pascual, 1972). In view of the specializations of metacheiromyids and the probability that Metacheiromyidae did not evolve before the Tiffanian, their resemblances to xenarthrans are more likely the result of convergence than of true affinity. Among known fossil forms, epoicotheriids are the most likely candidates for xenarthran ancestors, although this is probably dependent on the existence of earlier representatives than are now known. Derivation of manids from early palaeanodonts is possible and may be supported by the presence of an early manid, Patri- omanis, in the Chadronian of North America. However, contem- porary (or possibly older) manids have long been known from the Quercy Phosphorites of Europe (e.g. von Koenigswald, 1969) and a middle Eocene manid was recently reported from Messel, Germany (Storch, 1978). A direct land connection between North America and Europe existed until the end of Sparnacian (early Eocene) time (McKenna, 1972), hence it was presumably possible for manids to disperse in either direction before the middle Eocene. If manids evolved from palaeanodonts, it is most parsimonious to postulate North America as the place of origin, with subsequent dispersal to Europe. There can be little question that palaeanodonts were the closest ecological analogues of xenarthrans and pholidotans in the early Tertiary. It may be that palaeanodonts represent an early independ- ent radiation convergent to these orders. Evidence presented here and in other recent studies, however, strengthens the possibility of relationship among these groups. Once again it appears reasonable, as Matthew (1918: 655) wrote sixty years ago, “that there is, after all, a real affinity between the Pholidota and Xenarthra.” 1979 PALEOCENE PALAEANODONT 13 ACKNOWLEDGMENTS I am greatly indebted to Dr. Farish A. Jenkins, Jr. and Mr. Charles R. Schaff (MCZ) for permission to describe the holotype, and to Drs. Donald Baird (PU) and Frederick Szalay (Hunter College) for permission to study the humeri from Princeton Quarry. To Mr. William Amaral (MCZ), I extend my gratitude for prepar- ing the holotype. I thank also Drs. Walter Coombs (AC), Robert J. Emry (National Museum), Philip D. Gingerich (UM), and Malcolm C. McKenna (AMNH) for access to specimens used in this research, and Dr. George Gaylord Simpson for calling my attention to the recent article by Dr. Gerhard Storch. The figures were skillfully drawn by Ms. Karen Payne. Finally, lam grateful to Drs. Emry and Gingerich, Mr. David Krause and Prof. Bryan Patterson for their careful critical reading of the manuscript and suggestions for its improvement. REFERENCES BuTLeR, P. M. 1972. The problem of insectivore classification. In K. A. Joysey and T. S. Kemp (eds.), Studies in Vertebrate Evolution. Edinburgh: Oliver and Boyd, p. 253-265. COLBERT, E. H. 1942. An edentate from the Oligocene of Wyoming. Notulae Naturae, No. 109: I-16. Emery, R. J. 1970. A North American Oligocene pangolin and other additions to the Pholidota. Bull. Amer. Mus. Nat. Hist. /42: 455-510. FREELAND, G. L., AND R.S. Dietz. 1971. Plate tectonic evolution of Caribbean —Gulf of Mexico region. Nature 232: 20-23. GAZIN, C. L. 1952. The Lower Eocene Knight Formation of western Wyoming and its mammalian faunas. Smithson. Misc. Coll. //7: 1-82. GuTurig£, D. A. 1967. The mammalian fauna of the Lysite Member, Wind River Formation, (Early Eocene) of Wyoming. Mem. S. Calif. Acad. Sci. 5: 1-53. HILDEBRAND, M. 1974. Analysis of vertebrate structure. New York: John Wiley and Sons, 710 pp. JEPSEN, G. L. 1940. Paleocene faunas of the Polecat Bench Formation, Park County, Wyoming. Part I. Proc. Am. Phil. Soc. 83: 217-341. MALFAIT, B. T., AND M. G. DINKELMAN. 1972. Circum—Caribbean tectonic and igneous activity and the evolution of the Caribbean plate. Geol. Soc. Amer. Bull. 83: 251-272. MARSHALL, L. G., R. PASCUAL, G. H. Curtis, AND R. E. DRAKE. 1977. South American geochronology: radiometric time scale for middle to late Tertiary mammal-bearing horizons in Patagonia. Science 1/95: 1325-1328. 14 BREVIORA No. 455 MATTHEW, W. D. 1918. Part V. Insectivora (cont.), Glires, Edentata. Jn W. D. Matthew and W. Granger, A revision of the Lower Eocene Wasatch and Wind River faunas. Bull. Amer. Mus. Nat. Hist. 38: 565-657. McKeENnaA, M. C. 1972. Was Europe connected directly to North America prior to the middle Eocene? Evol. Biol. 6: 179-188. PATTERSON, B., AND R. PASCUAL. 1972. The fossil mammal fauna of South America. /n A. Keast, F. C. Erk, and B. Glass (eds.), Evolution, Mammals, and Southern Continents. Albany: State University of New York Press, p. 247-309. Reep, C. A. 1954. Some fossorial mammals from the Tertiary of western North America. J. Paleontol. 28: 102-111. Rose, K. D. 1978. A new Paleocene epoicotheriid (Mammalia), with comments on the Palaeanodonta. J. Paleontol. 52: 658-674. Rose, K. D., T. M. Bown, AND E. L. SIMons. 1977. An unusual new mammal from the early Eocene of Wyoming. Postilla, No. 172: 1-10. 1978. Alocodontulum, a new name for Alocodon Rose, Bown and Simons, 1977, non Thulborn 1973. J. Paleontol. 52: 1162. Simpson, G. G. 1931. Meracheiromys and the Edentata. Bull. Amer. Mus. Nat. Hist. 59: 295-381. 1932. Enamel on the teeth of an Eocene edentate. Amer. Mus. Novitates, No. 567: 1-4. 1948. The beginning of the age of mammals in South America. Bull. Amer. Mus. Nat. Hist. 9/: 1-232. 1959. Anew middle Eocene edentate from Wyoming. Amer. Mus. Novi- tates, No. 1950: 1-8. 1978. Early mammals in South America: Fact, controversy, and mystery. Proc. Am. Phil. Soc. /22: 318-328. StorcH, G. 1978. Ein Schuppentier aus der Grube Messel—zur Palaobiologie eines mitteleozanen Maniden. Natur und Mus. /08: 301-307. VON KOENIGSWALD, W. 1969. Die Maniden(Pholidota, Mamm.) des europaischen Tertiars. Mitt. Bayer. Staatssamml. Paldont. hist. Geol. 9: 61-71. West, R. M. 1973. An early middle Eocene epoicotheriid (Mammalia) from southwestern Wyoming. J. Paleontol. 47: 929-931. co BRE VI OR Museum of Comparative Loology US ISSN 0006-9698 CAMBRIDGE, MASS. JuLY 31s 1979 NUMBER 456 a DESCRIPTION OF A NEW HAWAIIAN GOBIID FISH OF THE GENUS TRIMMA PHILLIP S. LOBEL! ABSTRACT. A new gobiid fish species, Trimma taylori from Hawaii is described. A key to the nine nominal species of Trimma and a discussion of the Hawaiian gobies are included. The genus Trimma is reviewed and a preliminary assessment it presented of the possible relationships of Trimma. The nominal species of Trimma as now recognized may eventually be seen to comprise more than one group. INTRODUCTION Gobiid fishes are among the least known of the tropical marine fauna due to their generally miniscule size and secretive habits. Despite the difficulty of collecting and identifying such small reef fishes, a great number of species (over 1500) and genera (ca. 550) have been described (E. Lachner, pers. comm.). This high species diversity of gobiids rivals and exceeds that of most other fish families. However, the full extent of goby species diversity is far from known. For example, Trimma presently contains eight nominal species but an additional 25 to 30 remain to be described (D. Hoese, pers. comm.). The primary task of gobiid systematists at this time remains the study of species identification and distribution. The new goby described herein is the first Trimma to be found in Hawaiian waters. This species brings the total number of Hawaiian fishes to 682 (see Randall 1976). 'Museum ot Comparative Zoology, Harvard University, Cambridge, Massachu- setts 02138. 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CHE yisug] uy yepneD (€pE-L'7E)S CE CiTisesOlale. | 6i8¢ (UEC=S'6O)T TE (E°SE-89TO OE 0°67 uly o1Ajad JO UISIO 01 NOUS (9°LS-8'°SS)L'9¢ (T9S-TISNE9S 79S (p€S-0°08)6' 1S (S19-O'0S)S'ES ¥'7S UIJ [PSIOP PUOIIS JO UITIIO 0} INOUS (9'SE-€ EL)S VE (SvE-I'8061E OE (7 8€-€'€£)6°SE (68E-S6DESE OF UL} [PSIOP ISAT} JO UIBIIO 01 NOUS (8°97-7 610 €Z (SEZ-E8DT7 =H EZ (€°97-$°61)0°7Z (SC=8EWS IG IOC yysus] sjounpad jepney (9O0I-00DE 01 (7T71-8'6)9 01 GGl (L;OI-¢'L)S°8 (L'0I-¥'L)P'6 SOI ajounpad Jepnes jo yidap isea] (87-7 7)9'7 (OP 6 DLT IC (9 p-L'78'E (9S 6 O)6'E Ov YIPIM [eUqsosajur Auog (6 01-¢ 67 01 (T1I-16)7 01 Gl (€1l-p 6)r' Ol (8'11-2'6)8'01 L6 Ja}9W Ip Aq (0°S-0'°S)0'S (¢'p-€'©)0' ly (OS PHT (09-0' Orr 0'P yysuaj jnoug (€O£-€'67)8' 67 (BGGEGECIEUGe UCREC (0° 1€-6'$7)9' 87 (OTE-PZ9LZ O'S7 yisua] pray (SEI-91TI)9TI (SLI-V EDI II SLI (7SI-8' OP II (eS1-S 6)S TI I Sq[!3 puryaq yIpim Apog t USiCCSL BO Iie 89 iS (9'€7-7'07)6 IZ (C.SCsORD9 UC | Cie (C1E-1i6D)S ce CC Apog Jo yrdap Isayea1H UU(g'OZ-S EEOC WWUCL'OC-O'ECh PC LOT WUW(GPI-VEND Sel Ww(sCI-S IDL El SSI uw YyiBu2] prepuris é 2 2 é ° e Z=N adAJOJOH OUT G=N adAojoH VaNISAVO VAWIEL 9=N edAjO[OH [oul Z[=N adAiojoy IYOTAVL VAWISL yidua] prepueys jo uadiad ul [(adue1) ues] Dunia) “7 pur 10/.tD) “7 JO UOSURdWOD dLUIAWOYdIO |W | ATV 1979 GOBIID FISH OF THE GENUS TRIMMA 3 MATERIALS AND METHODS All specimens were measured with dial calipers. Proportional measurements were made by micrometer in a dissecting microscope and with calipers. Proportions are expressed as percent of standard length (SL) in millimeters (Table 1). Fish were collected with the anesthetic quinaldine mixed with 95% isopropyl alcohol. Specimens of T. taylori were captured in a fine meshed net and immediately placed into vials to prevent damage. Color photographs were taken as records of live coloration. Fifty specimens have been deposited as representative types of the species. Abbreviations of institutional names cited herein are as follows: AMS, Australian Museum, Sydney; BPBM, Bernice P. Bishop Museum, Honolulu; BMNH, British Museum of Natural History, London; CAS, California Academy of Sciences, San Francisco: MCZ, Museum of Comparative Zoology, Harvard University: USNM, National Museum of Natural History, Washington, D.C. Comparative material examined: Trimma caesiura, holotype, USNM 51772; USNM 156727 to 156738 (25+ specimens); USNM 218362. T. tevegae, paratypes, USNM 203437 (9). T. eviotops, holotype, USNM 116169; paratypes, USNM 116170 (3); paratype, MCZ 37267 (1). X-ray photographs of these species are on file at the USNM and BPBM. I have also examined species of related genera deposited at the USNM, MCZ, AMS, and BPBM. Trimma Taylori sp. nov. Pig. 1; Tablet: HOLOTYPE: BPBM 19919, 15.5 mm SL, taken ina cave at 44 meters depth off the leeward coast of Oahu, Hawaii, near Makaha Beach, uly 121975, P.S: Lobel: PARATYPES: BPBM 19920, 15 mm SL, locality same as holotype. July 20, 1975, P. S. Lobel (specimen photographed in color and figure 1). MCZ 51685, 14 mm SL, collected with holotype. BPBM 19921, 2 specimens each 14 mm SL; collected with BPBM 19920 (cleared and stained). MCZ 51686, 2 specimens, 13-15 mm SL; collected with BPBM 19920. MCZ 51687, five specimens, 12—13.5 mm SL; collected with holotype. USNM 215323, 9 specimens, 6-13 mm SL; same location as holotype, Aug. 13, 1975, P.S. Lobel. AMS 118686-001, eight specimens, 6-13 mm SL; same location as 4 BREVIORA No. 456 Figure 1. Trimma taylori, male 15 mm. s.1. Paratype BPBM 19920. holotype, Aug. 13, 1975, P. S. Lobel. CAS 35450, six specimens, 11-13 mm SL; collected with BPBM 19920. BPBM 19922, eight specimens, 6-14.5 mm SL; same location as holotype, Aug. 13, 1975, P. S. Lobel. BMNH 1976.4.6.1—7, seven specimens, 6-13 mm SL; same location as holotype, Aug. 13, 1975, P. S. Lobel. Diagnosis. This species can be easily distinguished from all other Trimma spp. by a combination of characters: Dorsal VI-I, 10, anal I, 10, and pectoral 14; second spine of the first dorsal fin filamentous, longest, extending past the midpoint of the second dorsal fin when depressed. Description. Morphometric characters of specimens are presented in Table |. Tooth shape and pattern are illustrated in Figure 2. Head length 25% of standard length. Eye diameter about 40% head length. First dorsal fin originates one third the standard length distance from snout. Mouth slanted about 45° upward, corner of mouth in line with midpoint between forward margin of eye and pupil. Eye to snout distance approximately one-half eye diameter. Body scaled except for head and operculum, but including nape. Scales in longitudinal series from opercule to hypural base, 22-23. Seven scales from the origin of the anal to the first dorsal. Middorsal area from first dorsal spine to head with three scales. Gill raker count of three specimens: 3-1-12, 3-1-13, and3-1-14. Five lobes on 1979 GOBIID FISH OF THE GENUS TRIMMA 5) pseudobranch. Fin ray formulae: Dorsal VI-I, 10, anal I, 10, pelvic 15, and pectoral 14. Vertebral count 25, not including urostyle. Dorsal fins separate. Second spine of first dorsal fin elongated, extending over second dorsal fin. Posterior most ray of second dorsal fin and of anal fin split to base but counted as one. Pelvic fins connected at base but not united by frenum. Additional characters are the same as described for the genus (see Cohen and Davis 1969). Color in life. Overall color pale straw yellow with darker yellow pigment spots bordering base of each scale. Immediately posterior to the eye, orange spot almost equal in area to eye. Dorsal portion of this orange spot speckled with small melanophores. Head dorsally speckled with numerous orange and black chromatophores. Dorsal and anal fins with broad deep orange band at base and with thin band of black pigmentation above it; fins overall light orange, with a diffuse dark edge. Caudal fin similar in color to dorsal and anal fins but without deep orange band. Pelvic and pectoral fins yellow- orange. Figure 2. Trimma taylori, detail of mouth and teeth. S.E.M. photo by E. Seling. 6 BREVIORA No. 456 FEMALE Figure 3. Trimma tavlori, sexual dimorphism of genital papillae. Drawn from MCZ 51686. Paratypes by Karsten Hartel. Color in alcohol. All colors vanish, body pale. Pigment spots on dorsum of head visible after one year but fade thereafter. Fins transparent. Sexual dimorphism. Morphometric differences are not immedi- ately apparent between the sexes; however, males and females may be distinguished on the basis of genital papillae (Fig. 3). The papilla of the male is slender, curved and tapers to a point posteriorly. The female’s papilla is greater in diameter than that of the male and uniform in thickness, terminating ina round opening with a bilobed flap bilaterally; it is also directed posteriorly but it is not as long as that of the male. Immature gobies exhibit no sexual difference in the shape of the papilla (see also Egami 1960). There are significant differences in both shape and size of male and female genitalia among Trimma eviotops, T. taylori and T. caesiura. The genitalia of 7. tay/ori and T. caesiura are very similar in shape and structure but differ in relative length of the papilla of the male. 1979 GOBIID FISH OF THE GENUS TRIMMA 7 The relative lengths of male papillae (in percent of standard length) are: T. taylori6%, T. caesiura 3.4%, and T. eviotops 2.2%. The papilla of T. eviotops is not curved as 1s that of T. ray/ori (Fig. 3), but extends straight posteriorly and is comparatively thinner and not tapering to a point, ending bluntly. A pigment line runs ventrally along the length of the male papilla of T. eviotops. Arai (1964) illustrated the genital papillae of 25 goby species. Four are particularly relevant here: Eviota abax (Jordan and Snyder), Quisquilius eugenius (Jordan and Evermann), Zonogobius semi- doliatus (Cuvier and Valenciennes) and Fusigobius neophytus (Giinther) (Fig. 4 modified from Arai, 1964). The genitalia in both sexes of 7. tav/ori and T. caesiura appear most like that of Fusigobius neophytes. The genital papilla of 7. eviotops is more similar to those of Zonogobius semidoliatus. The genitalia of Quisquilius and Zonogobius are superficially more similar to each other than either is to those of Fusigobius. Such comparisons may become important when the phylogeny of these taxa is revised. SE EVIOTA ABAX ee Fa) ae a QUISQUILIUS EUGENIUS ZONOGOBIUS SEMIDOLIATUS as eG HISTSOBIIIS NEORHYTES Figure 4. Genital papillae of some related Gobiids. Modified from Arai (1964). 8 BREVIORA No. 456 Figure 5. Trimma taylori, detail of scales on flank. S.E.M. photo by E. Seling. Scale Morphology. The number of ctneii in a single row along the posterior margin of the scale may differentiate species of Trimma. The pattern and structure of scales of T. tayv/ori (Fig. 5), and of T. caesiura (Fig. 6), 1s characteristic for each species. Trimma taylori possess fewer and larger scale ctenii (mean 10.9, range 8-14 per scale, N = 12) than does T. caesiura (mean 31.6, range 28-36, per scale, N = 5). Comparison to other species was not possible due to the unavailability of specimens. Consequently, the taxonomic value of such scale morphology is uncertain. Zoogeography. Trimma taylori is known only from the island of Oahu, Hawaiian archipelago. All other 7rimma species occur in the Indopacific oceanic region or in the Indian Ocean. None are recognized from the Atlantic Ocean or within the temperate zones. All Trimma spp. with known habits are reef dwellers. Entymology. Trimma taylori is named in honor of Dr. Leighton R. Taylor, Jr. on the occasion of his appointment as director of the Waikiki Aquarium. A tradition of the Waikiki Aquarium has been 1979 GOBIID FISH OF THE GENUS TRIMMA 9 that each of its directors have a uniquely Hawaiian fish as a namesake. DISCUSSION The nominal species of Trimma The original description of the genus Trimma was not adequate and there has consequently been confusion in the generic assignment of the species. The genus 7rimma was defined on the basis of T. caesiura by Jordanand Seale (1906). It was diagnosed as “near Eviota but the region before the dorsal is fully scaled. In texture the species is firmer and less translucent” (Jordan and Seale, 1906, p. 361). This description does not adequately define the genus with regard to some of the species that have since been included in it. Additional morphometric data of 7. caesiura are listed in Table | with comparisons to 7. taylori. The species included in the genus Trimma which appear closely related to 7. caesiura based on the original generic characters and other characters described below are Trimma naudei Smith (1956) and 7. tevegae Cohen and Davis (1969). Other species placed in 7 Bet k : C Figure 6. Trimma caesiura, details of scales on flank. S.E.M. photo by E. Seling. 10 BREVIORA No. 456 Trimma present an enigma. 7Trimma eviotops Schultz (1943) is presently considered a valid species but its placement in Trimma will soon be reconsidered (S. Karnella, pers. comm.). It clearly differs from 7. caesiura in the lack of scales along the nape. 7rimma eviotops does not fit comfortably in other closely related genera. For example, it differs from species of Eviota by lacking head pores, although this may not be taxonomically significant. Smith(1956) remarked that 7. eviotops may well become the type of a new genus. Consequently, it is included here only provisionally. However, there is no difficulty in assigning Trimma taylori to the genus Trimma. I follow Hoese’s (pers. comm.) unpublished classification for species of Trimma. The species (as originally designated) are Eviota macrophthalma Tomiyama (1936), Eviota grammistes Tomiyama (1936), Zonogobius flammeus Smith (1959) and Zonogobius coral- linus Smith (1959). These all differ from 7rimma caesiura in lacking predorsal scales along the nape but resemble 7rimma eviotops in that character. Until related species and genera are better known the decision to include these species in 7rimma involves the fewest complications. However, if 7. eviotops is removed from Trimma, then the generic placement of these other species should also be carefully reconsidered since some are more similar to 7. eviotops than to 7. caesiura, the species on which the genus is based. Meristic and color traits of the nominal species of 7rimma are contrasted in Table 2. There exists numerous problems with the classification of the genera and species of gobies. However, most of the conflicts cannot be solved until both more species are described and the characters defining the genera are delineated. The genus 7rimma is no exception but since assignment of 7rimma taylori into this genus offers no conflicts, discussion of other issues 1s postponed until more data are available. Ecology. Trimma taylori appears to be an exclusively deep reef species that inhabits the recesses of caves and crevices. Its habits are somewhat like those described for 7. tevegae (Cohen and Davis, 1969). During the daytime, swarms of 50 to 150 individuals hover near the ceiling, in about the midsection of a cave or pocket. The shallowest location at which adults have been observed is at a depth of 30 meters. A few juveniles have been seen somewhat shallower. 1979 GOBIID FISH OF THE GENUS TRIMMA 11 The cave habitat in which all specimens were collected was typically a small crevice (one to two meters deep) alonga vertical wall of the deep reef, Oahu, Hawaii. Perhaps it is significant that these shallow pockets did not house holocentrids or priacanthids. Caves containing the latter were inspected for 7rimma taylori but none were found. Sympatric cave inhabitants which were conspicuous included a prawn, Stenopus (an undescribed species) and the fishes Prerois sphex (Scorpaenidae), Apogon maculiferus (Apogonidae), Gymno- thorax spp. (Muraenidae), an undescribed species of Quisquilius (Gobiidae), Scorpaenid sp., and Pseudanthias sp. (Serranidae- Anthiinae). Of these fishes, only Prerois sphex, Gymnothorax spp. and Apogon maculiferus seem likely potential predators of T. taylori. Trimma taylori feeds on harpacticoid copepods. Eight specimens were examined for gut contents and three of these were empty. KEY TO THE NOMINAL SPECIES OF TRIMMA [Peamebredorsal-scalestonmnaper tices stais tells isola viccns soreeus anaes olsae etevele aiatecere 2 [, IN@ GeAlles Cine Boeke asoonsacecoUDeS acoDaN So Osos oma baDactotoou™ 5 rer beg) OLS SIM WAT SPRAYS ws, creeteva cic ke rete ral avel eaterebocucpecsyoruslonene ale locreteratensheveeste ae 3} bseDorsalafinewithemrore) than Sora ySicnusicesioe erie cleric cre clevsie-etereners achotehe aero 4 Bare Gillirakeron first anchystitllG; syacqeevactaescraraelol artery era T. caesiura beeGuillirakeronifirst arch! 3113) on 14) Sees eee siete ees ete T. naudei 4\ ay IDosaliaiy WIE UO. letxcorell ue Gesu osonpasuncosncovscoodsoDonS T. tayvlori by, IDorviliiin WiC) taco rililS) GoasooeuoumondsoosacdosudduDens T. tevegae rasebectoraluini withtewer tham) 19) rays) cv. c-teeis cro) erosive oravcicvets rere 6 be Rectorallifim swath U9 raysy ec crerevercneiore cere oer oye avevonsitovel(eneve eye) sree T. coralinus Omdeerectoraletini with ara ys cs sete ccistrsiee lela cielciietcinie letters ci ieusinenstcten 7 baeectoralttim: withwfewermthanull/mayse ee crcisciee i ctaers er tocracircrerserrcrac. 8 (pea body with eight dark bars) or saddiles' 2% 42.2... 2.4. 2-4-0055) T. eviotops beeBodviwithoutibarsromsaddlestepacemrieie acictaciicricrroetr tc are T. flammeus 8. a. Pectoral with 15 rays, 11 scales transverse series ............ T. grammistes b. Pectoral with 16 rays, 9 scales transverse series ......... T. macrophthalma SUPPLEMENTARY KEY TO HAWAIIAN GOBIID FISHES [Pease pelvicatinsyseparater(n siren) mey-toreiiste ere iicrocterciere casteisielevsiiona axe ere yonel ones 2 b. pelvic fins united to forma cup ........ See Gosline and Brock 1960, p. 266 2. a. each scale with a dark crescent shaped mark, no spines on the preopercle b. no markings on scales, head compressed, 3 to 5 spines on the preopercle Ue oe or Suse ate, cub lchalte: sroaaranesausovene © evaiec elavain cus sles Asterropteryx semipunctatus Seaawnolpredorsallscalesyoninapelccrs-ieisciiinctce aces ere Eviota epiphanes bss predorsallscalesontnape presents. a. aren oe sa slo e ois tere ote Trimma taylori 12 BREVIORA No. 456 TABLE 2 Comparison of the Nominal species of Trimma Pec- Pel- Longi- Nominal Dorsal Anal toral vic tudinal Species Author fin fin fin fin scales T. caesiura' Jordan and VI-I.8 1,8 15 1,5 23-25 Seale 1906 T. coralinus (Smith 1959) VI_-I,10 1,9 19 Zs 30 T. eviotops Schultz VI-1,9-10 1,8-9 17 1,5 26-27 1943 T. flammeus (Smith 1959) VI-I,9-10 1,8—10 17 — 26-36 T. grammistes? (Tomiyama VI-1,10 1,9 15 IES 30 1936) T. macroph- (Tomiyama VI,1,9 1,9 16 les 23 thalma? 1936) T. naudei Smith 1956 VI-1.8 1,8 16 — 24-25 T. taylori n.sp. VI-I,10 1,10 14 1,5 22-23 T. tevegae Cohen and VI-I,9 1,9 13 It) 28 Davis 1969 'l examined 21 specimens for additional counts. ?Pectoral fin count from drawing with description. 1979 Scale Trans- verse 6-7 10-11 8-9 GOBIID FISH OF THE GENUS TRIMMA 13 TABLE 2 Comparison of the Nominal species of 7rimma Gillrakers on first Verte- arch brae Locality 3-1-16 25 Samoa 3+1+10 - Seychelles 3+1+11-12 25 Phoenix & Samoan Islands 3+1+11-12 = Australia & S. Africa- — — Japan — — Japan 3+1+13-14 — Seychelles 3+1+12-14 25 Hawaii — 25 New Britain Live Coloration Plain bright red with grey spots on back of tail and grey spots on many scales of back and head. Fins all bright red. Body red with two silvery areas on back below first, and three below second dorsal fin. Orange spots on head and peduncle. Rays red with pink membrane. Pale with eight vertical bars, saddles on head and body. Light orange with darker spots; edge of soft dorsal and caudal, dusky. Longitudinal dark bands. (not described) Body and fins brilliant orange- scarlet, irregularly mottled with iridescent light areas; dorsal fin with two light pink bars between the orange. Allstraw yellow with orange-red and dark pigment spots on head. (Live body color not described) Dark reddish spot at end of caudal peduncle. 14 BREVIORA No. 456 NOTE: Asterropteryx semipunctatus and Eviota epiphanes were placed in the Eleotridae by Gosline and Brock (1960). However, the only eleotrid in Hawaii is Eleotris sandvicensis, which is restricted to freshwater. It can be easily distinguished from the gobies. E. sandvicensis possesses about 75 scales in longitudinal series whereas gobies generally have less than 30 longitudinal scales. In addition, eleotrids are primarily freshwater and have six branchiostegal rays; gobies are mostly marine and have five branchiostegal rays. There are 2] other gobies presently known in the Hawaiian Islands. ACKNOWLEDGEMENTS I thank Douglas Hoese (AMS) who first examined my specimens and confirmed them as a new species. D. Hoese also generously supplied valuable information which made my task considerably more manageable. Ernest Lachner (USNM) and Susan Karnella (USNM) were especially helpful by explaining to me many of the problems associated with goby systematics and loaning specimens. I thank Karel Liem (MCZ), John Randall (BPBM) and especially William Fink (MCZ) for their comments. Ed Baughman, John Earle and Janie Culp helped to collect the specimens. Ed Seling (MCZ) prepared the scanning electron microphotographs with support from NSF grant BMS 7412494. Karsten Hartel (MCZ) prepared Figure 3. Kathie Cunningham and Terry Dash kindly typed the manuscript. Support for field work was provided by the Department of Zoology, University of Hawaii while I was an undergraduate there. Additional support and travel were provided by the Biology Department, Harvard University. LITERATURE CIPED ARAI, R. 1964. Sex characters of Japanese gobioid fishes (1). Bull. Nat. Sci. Mus. Tokyo, 9: 295-306. COHEN, D. M. AND W. P. Davis. 1969. Vertical orientation in a new gobioid fish from New Britain. Pac. Sci., 23: 317-324. EGAM1, N. 1960. Comparative morphology of the sex characters in several species of Japanese gobies, with reference to the effects of sex steroids on the characters. J. Fac. Sci. Univ. Tokyo, 9:67-100. GOSLINE, W. A. AND V.E. Brock. 1960. Handbook of Hawaiian fishes. Honolulu: University of Hawaii Press, IX + 372 pp. JoRDAN, D. S. AND A. SEALE. 1906. The fishes of Samoa. Bull. Bur. Fish., 25: 173-455. RANDALL, J. E. 1976. The endemic shore fishes of the Hawaiian Islands, Lord Howe Island and Easter Island. Collogue Commerson 1973. O.R.S.T.O.M. Travaux et Documents., No. 47: 49-73. 1979 GOBIID FISH OF THE GENUS TRIMMA 15 SCHULTZ, L. P. 1943. Fishes of the Phoenix and Samoan islands collected in 1939 during the expedition of the U.S.S. “Bushnell”. Bull. U.S. Nat. Mus., 180: 1-316. SmiTH, J. L. B. 1956. The fishes of Aldabra. Part 6. Annals and Magazine of Natural History, Ser. 12., Vol. 9. pp. 817-829. SmiTH, J. L.B. 1959. Gobioid fishes of the families Gobiidae, Periophthalmidae, Trypauchenidae, Taenioididae, and Kraemeriidae of the Western Indian Ocean. Rhodes Univ. Ichthy. Bull., 13: 185-225. TOMIYAMA, I. 1936. Gobiidae of Japan. Jap. J. of Zool., 7: 37-112. MAR 1 8 19x5 HARVARD BhRE VI O“RTA Museum of Comparative ZLoology US ISSN 0006-9698 CAMBRIDGE, MASS. JANUARY 31, 1980 NUMBER 457 TWO NEW SPECIES OF ELEUTHERODACTYLUS (AMPHIBIA: LEPTODACTYLIDAE) FROM THE LOWLANDS AND LOWER CLOUD FORESTS OF WESTERN ECUADOR JOHN D. LYNCH! AND KENNETH MIYATA2 ABSTRACE: Eleutherodactylus muricatus sp. nov., an ally of the large, flare- snouted frogs of the rubicundus assembly, is named from lowland and lower cloud forest localities in Provincia Pichincha, Ecuador. The new species is smaller than its sympatric allies E. crenunguis and E. latidiscus. Eleutherodactylus tenebrionis sp. nov., is also found in primary lowland and lower cloud forests in west-central Ecua- dor. It is allied to a species found in the high cloud forests in western Ecuador but differs in color pattern and lacks a calcar and ulnar tubercles. INTRODUCTION Approximately I5 species of Eleutherodactylus inhabit the Pacific lowlands of Ecuador (Lynch, in press) and perhaps another 40 species are found in the cloud forests of the Pacific versant in the Ecuadorian Andes. Most of these frogs are small organisms (less than 35 mm SVL) but two members of the rubicundus assembly [ Eleutherodactylus crenunguis Lynch and E. latidiscus (Boulenger)] are much larger species having long, slender limbs and digits and large emarginate (or notched) digital pads. Our field work in western Ecuador over the past several years has revealed that crenunguis is a frog of the lower cloud forests (800-1500 m) and is not an altitudinal replacement for Jatidiscus (20-1500 m). In the course of this field work two additional species having notched or indented digital pads were found in sympatry ‘John D. Lynch, School of Life Sciences, The University of Nebraska, Lincoln, Nebraska 68588 *Kenneth Miyata, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 9) BREVIORA No. 457 with both of the larger species. In spite of sharing the notched pads, only one of these species seems allied to the rubicundus assembly as defined by Lynch (1979). The other species appears to be most closely allied to a species found in the high cloud forests of western Ecuador (Lynch and Trueb, in press). The format of the descriptions follows Lynch’s long-used style. Abbreviations employed below include the following: SVL — snout-vent length; HW — head width; IOD — interorbital distance; E-N — eye to nostril distance. Eleutherodactylus muricatus sp. nov. Holotype: MCZ 94469, an adult male collected at the Rio Fai- sanes where it is crossed by Ecuador Highway 28 (the road from La Palma to Quito via Chiriboga), 14.4 km from the junction with Highway 30 (the Aloag to Santo Domingo de los Colorados road) at La Palma, Provincia Pichincha, Ecuador, 1380 m, on 12 Novem- ber 1977 by Ken Miyata. Paratypes. MCZ 92091, 92095, 92100-01, 94848, 97528-31, USNM 211172-74, topotypes; MCZ 97592, Centinela, 14.1 km SE Patricia Pilar by road, Provincia Pichincha, 570 m; MCZ 90337, 94456, 94460, Centro Cientifico Rio Palenque, 47 km S Santo Domingo de los Colorados, Provincia Pichincha, 220 m. Diagnosis. 1) skin of dorsum smooth with numerous conical tubercles, that of venter feebly areolate; no dorsolateral folds; no anal sheath; 2) tympanum moderately distinct, its length 4 eye length; 3) snout subacuminate in dorsal view, rounded in lateral profile; canthus rostralis moderately distinct; 4) upper eyelid much wider than IOD, bearing many conical warts: no cranial crests; 5) vomerine odontophores large, triangular in outline, narrowly separ- ated; 6) males with vocal slits, subgular vocal sac; no nuptial pads; 7) first finger shorter than second; all digits bearing broad discs, pads on fingers II-IV, those of II] and IV notched; 8) fingers bear- ing lateral keels; 9) 1-2 small ulnar tubercles; 10) one large conical tubercle on heel; low tubercles along outer edge of tarsus; short inner tarsal fold; 11) two metatarsal tubercles, inner elongate, 10 times size of round, subconical outer; supernumerary plantar tuber- cles at base of toes; 12) toes bearing lateral fringes, not webbed; toe TWO NEW SPECIES OF ELEUTHERODACTYLUS 1979 Wt 9'0¢ “P7Z6L1 (dq) “IAS WU gee “9Sbhh6 ZOW Ay ‘ 7 ‘S1UOLAG Ue AS wu A (OD) ‘ Sp SZIQIEL AM ‘SMesipur] “A (d) -TAS AS RECO SIL TORT Med SiUOiG obey ‘snipoaNiU SnpALopposay nay (W) “| ans 4 BREVIORA No. 457 pads notched, smaller than those of outer fingers; 13) dorsum brown with black spots; ventral surfaces brown with cream flecks; slightly darker brown chevrons on throat; posterior surfaces of thighs brown; 14) adults from type-locality moderate-sized, males 31.8- 40.7 (x= 36.0, n = 4) mm SVL, one female 46.3 mm SVL; two gravid females from the lowlands (Centinela and Rio Palenque) are only 33.8-36.0 mm SVL. Eleutherodactylus muricatus is most similar to E. crenunguis but differs in coloration (no orange patch on the breast) and in having fewer, larger tubercles on the dorsum (Fig. 1). It is also smaller than E. crenunguis (Lynch 1976). Description. Head as wide as or wider than body, wider than long: HW 37.1-39.9 (x = 38.4, n= 7) per cent SVL; snout subacum- inate in dorsal view, rounded in lateral profile; nostrils weakly pro- tuberant, directed dorsolaterally; canthus rostralis relatively sharp (swollen), straight or weakly convex; loreal region concave, sloping gradually to lips; lips flared; E-N 80.7-100.0 per cent (x= 89.4, n= 7) eye length; upper eyelid 100.0-132.3 per cent (x = 119.7, n= 7) IOD, bearing many pungent tubercles; no cranial crests; supratym- panic fold moderately distinct, obscuring upper edge of tympanum; tympanum not prominent, round, separated from eye by distance equal twice tympanum length; tympanum length 22.0-27.9 (x = 25.1, n= 6) per cent eye length except in MCZ 94456 (small female from Rio Palenque has ratio of 37.5 per cent); postrictal tubercles present, not prominent; choanae round, not concealed by palatal shelf of maxillary arch; vomerine odontophores median and poste- rior to choanae, large, triangular in outline, separated on midline by distance less than 1/3 an odontophore width, each larger than a choana, bearing 5-12 teeth in a transverse row; tongue longer than wide, its posterior edge notched; posterior 1/3 not adherent to floor of mouth; males with vocal slits posterolateral to tongue, median subgular vocal sac. Skin of dorsum smooth but bearing many warts, most numerous on lower back (where skin is tuberculate); many elongate, subconi- cal warts on flanks; no dorsolateral folds; no anal sheath; skin posterior and posterolateral to anus areolate but also bearing coni- cal warts; skin of throat areolate, that on other ventral surfaces smooth with very feeble areolations; discoidal folds well anteriad to groin; one or two small ulnar tubercles; palmar tubercle bifid, larger 1979 TWO NEW SPECIES OF ELEUTHERODACTYLUS 5 than oval thenar tubercle; several prominent supernumerary palmar tubercles; subarticular tubercles round, pungent; fingers bear lateral keels; all digits with pads, that of thumb scarcely wider than digit below pad, on II twice as wide, on III-IV three times as wide (wider than tympanum); pads of fingers III-I1V notched apically; discs on all pads broader than long; fingers long, first slightly shorter than second; thumb of male lacking nuptial pad or swelling. Numerous tubercles on heel, one large conical tubercle (not a calcar); low tubercles along outer edge of tarsus; short fold at base of inner metatarsal tubercle which is three times as long as wide, 10 times size of round, subconical outer metatarsal tubercle; supernu- merary plantar tubercles at bases of each toe; subarticular tubercles longer than wide, pungent; toes bearing distinct lateral fringes, broad discs, expanded pads; pads notched apically, smaller than those of outer fingers; heels of flexed hind limbs broadly overlap; shank 54.7-62.2 per cent (x = 58.4, n = 7) SVL. Brown above with black spots enclosing tubercles; bars evident on thighs but remainder of pattern (limbs, labial bars, canthal- supratympanic stripe, dorsal chevrons, etc.) only suggested by black spots; venter brown with cream flecks and small spots; inverted brown chevrons on chin and throat; undersides of limbs and groin brown with cream flecks; anterior and posterior surfaces of thighs brown with cream flecks. In small individuals, ground color gray and with evident shank bars (narrow and oblique), labial bars, and canthal-supratympanic stripe. In life, E. muricatus from the type locality are pale to dark brown with black rings around large tubercles; flanks paler brown with yellowish wash; venter dark purplish-brown with yellow-brown mottling; iris gold flecked with black. Lowland specimens lack the black rings around the tubercles; the venter is a muddy yellow mottled with dark brown; the lower flanks, groin, and underside of the limbs have a purplish-brown wash; the iris is copper. The above color descriptions apply to daytime patterns; at night all individuals are much paler in dorsal coloration, ranging from pale buff to olive brown, but the various markings remain intact. Measurements of holotype (in mm). SVL 32.3; shank 20.1; HW 12.5; head length 11.6; upper eyelid 2.9; IOD 2.9; tympanum length 1.0; eye length 4.3; E-N 3.9. 6 BREVIORA No. 457 Etymology. The specific epiphet is derived from the Latin, meaning spiny, in reference to the pungent tubercles on the dorsum and upper eyelid. Variation. The two adult females from low elevations (MCZ 94456 from Rio Palenque, 220 m, and MCZ 97592 from Centinela, 570 m) are much smaller than the only adult female from the type- locality. They are also peculiar (Fig. 1) in having prominent verte- bral stripes which are yellow-cream to buffy orange in life. All of the material from the lower elevations has venters more pale than the topotypic material; in life the ventral surfaces of the topotypic popu- lation are predominantly brown with some yellowish mottling while those of the low elevation populations are predominantly yellowish with brown mottling. The lowland populations also have more prominent tubercles on the eyelids and dorsum. All of these differ- ences are rather minor and probably represent slight geographical or altitudinal variation. We are taking a conservative approach in assigning all of these specimens to the same taxon; further work may well prove we have confused two species. Remarks. Adults of E. crenunguis and E. latidiscus are seldom encountered. No adult females of E. crenunguis have been found, although juvenile females to 41.3 mm have been examined, and adult males range in size from 35.0 to 49.2 mm SVL. Very few adults of E. latidiscus are available even though juveniles are often quite common. Three males with vocal slits are 43.9-50.0 mm SVL and four females having convoluted oviducts are 49.7-64.5 mm SVL. Both of these frogs are considerably larger than the sympatric E. muricatus. Natural History. Eleutherodactylus muricatus from the type- locality have been collected on streamside vegetation within 1.5 m of the water surface. Several specimens were taken on logs just above water level, but the majority were perched on large leaves adjacent to the stream. For a description of the type-locality see Miyata (in press). Individuals are found in low density on most nights at this locality. The lowland specimens have all been taken from primary forest or, in the case of the Centinela specimen, from very recently eut primary forest. The Rio Palenque specimens were taken from low vegetation in forest with a dense canopy on rainless nights. 1979 TWO NEW SPECIES OF ELEUTHERODACTYLUS if Eleutherodactylus muricatus appears to have a spotty distribu- tion, somewhat analogous to that seen in other anurans in this region (Lynch 1977). E. muricatus has not been collected at the Rio Orito, a locality very close to the type locality, despite several visits by field parties from the University of Kansas. More field work is needed to confirm the spottiness of the distribution. “ Eleutherodactylus tenebrionis Holotype: MCZ 90326, an adult male collected at the Hotel Tinalandia, 16 km E Santo Domingo de los Colorados by road, Provincia Pichincha, Ecuador, 800 m, on 6 August 1975 by Ken Miyata. Figure 2. Outlines of heads of western Ecuador Eleutherodactylus. (A) E. crenunguis, MCZ 92099; (B) E. muricatus, MCZ 92095; (C) E. latidiscus, cotype, BM 98.4.28.109; (D) FE. tenebrionis, MCZ 92081. Scale for A-C is between figs. A and C; that for D is to its right. Scales equal 5 mm. 8 BREVIORA No. 457 Paratypes: MCZ 88890, 90325, 90327-29, 92079-81, 94712, top- otypes; KU 179224-27, Santo Domingo de los Colorados, Provincia Pichincha, 580 m; KU 179228-30, 2 km E, 1 km S Santo Domingo de los Colorados, Provincia Pichincha, 600 m; MCZ 97596-97, USNM 211176, Centinela, 14.1 km SE Patricia Pilar by road, Pro- vincia Pichincha, 570-600 m; KU 146171, 165874-77, MCZ 94864-— 65, 94867, 98164-66, USNM 211175, Centro Cientifico Rio Palenque, 47 km S Santo Domingo de los Colorados by road, Provincia Pichincha, 170-220 m. Diagnosis. 1) Skin of dorsum smooth with occasional tubercles, that of venter areolate; no dorsolateral folds; no anal sheath; 2) tympanum distinct, round, its length |/4—2/5 eye length; 3) snout round in dorsal view, truncate in lateral profile; canthus rostralis distinct; 4) interorbital space narrower than upper eyelid; low cra- nial crests in females; small tubercle on upper eyelid; 5) vomerine odontophores elevated, triangular in outline, narrowly separated; 6) males with vocal slits; males lack nuptial pads on thumb; 7) first finger shorter than second; all digits bearing broad discs on expanded pads, pads of fingers III-IV largest, emarginate, those of I-II smaller, rounded apically; 8) fingers lack lateral fringes; 9) no ulnar tubercles; 10) small tubercles on heel, none on knee or tarsus; 11) two metatarsal tubercles, inner oval, 4 times size of flat outer; low supernumerary tubercles at bases of toes II-IV; 12) toes lack lateral fringes; discs broader than long on weakly emarginate pads, pads smaller than those on fingers; 13) brown above with little indication of pattern (Fig. 1); venter cream with extensive brown reticulation; undersides of limbs brown with cream flecks; anterior and posterior surfaces of thighs brown with small cream flecks; in life, iris blue; 14) adults moderate-sized, males 20.8-26.8 (x = 23.8 + 0.9,n= 16) mm SVL, females 30.6-36.9 (x = 33.8 = 1.6, n= 7) mm SVL. Eleutherodactylus tenebrionis has no close relatives known to us except for an undescribed species from the upper cloud forests (2000-2700 m) in Provincias Imbabura and Pichincha, Ecuador (Lynch and Trueb, in press) which differs from E. tenebrionis in having prominent conical tubercles on the forearm, tarsus, and upper eyelid and a calcar on the heel. Description. Head as wide as or wider than body, wider than 1979 TWO NEW SPECIES OF ELEUTHERODACTYLUS 9 long; HW 37.0-41.4 per cent (x= 39.3, n= 23) SVL; snout round in dorsal view, truncate in lateral profile; E-N in males 72.0-84.8 per cent (x = 77.9, n= 16) eye length, in females 75.9-87.0 (x = 82.9, n = 7); nostrils protuberant, directed laterally; canthus rostralis dis- tinct, weakly concave; loreal region concave, sloping abruptly in males and gradually in females to lips; lips not flared (Fig. 2); upper eyelid 110.3-165.7 per cent (x = 125.8, n = 23) IOD, bearing 1-2 small conical tubercles; adult females have low cranial crests (not evident in smaller frogs); lower 3/4 of tympanum distinct, upper edge hidden by diffuse supratympanic fold; tympanum separated from eye by almost its length; tympanum length in males 25.0—35.0 per cent (x= 29.4, n= 16) eye length, in females 29.6—38.5 (x= 32.9, n = 7); small postrictal tubercles present; skin on rest of head smooth; choanae longer than wide, not concealed by palatal shelf of maxillary arch; vomerine odontophores median and posterior to choanae, pungent, triangular in outline, bearing 5S—7 teeth in a trans- verse row, separated on midline by a space equal to 1/3-1/2 of odontophore width; odontophores nearly as large as a choana; tongue longer than wide, its posterior border notched, posterior 1/5 not adherent to floor of mouth; male with vocal slits. Skin of dorsum essentially smooth but some small, low, flat warts on lower back (in a few examples these warts are pungent) and a few tiny warts scattered on back; no dorsolateral folds; flanks become areolate, areolation continuing onto venter; throat smooth; discoi- dal folds present; no anal sheath; no ulnar tubercles; palmar tuber- cles bifid (outer lobe the smaller), as large as or smaller than oval thenar tubercle; if present, supernumerary palmar tubercles flat, indistinct; subarticular tubercles relatively low, round; fingers lack- ing lateral fringe or keel; all fingers bearing broader than long discs; pad smallest on I, intermediate on II, largest on III and IV; pad of III as large as tympanum; pads on III and IV feebly emarginate; II longer than I; males lack nuptial pads. No tubercles on knee or tarsus; |—2 small tubercles on heel; inner metatarsal tubercle twice as long as wide, outer 1/4 (or less) size of inner, flat; supernumerary plantar tubercles low, at bases of toes II-IV; subarticular tubercles longer than wide, not conical; toes lack lateral fringes, bearing broad discs on expanded, feebly emarginate pads (toe pads smaller than those on outer fingers); heels of flexed hind legs overlap; shank of males 52.8-62.5 (x = 57.9, n= 16) per Eent SV i of females 52.2-59.7 (x = 56.1, n = 7). 10 BREVIORA No. 457 ” Brown above with diffuse dark brown or black scapular “W”, supratympanic stripe, interorbital bar, and indefinite sacral chev- ron; canthal stripe, labial bars, anal triangle brown, not distinct; limb bars brown, nearly transverse on shanks, about as wide as interspaces; flanks pale brown (cream invasion of brown); venter cream with dense fine brown reticulation; some cream flecks on limbs, fewest on underside of shank; throat and breast heavily stippled with brown (relative to venter); anterior and posterior sur- faces of thighs (and groin) brown with small cream flecks. In life, E. tenebrionis is burnt umber with black and dull golden flecks above; the venter is gray with gray-brown mottling. The iris is gray-blue. At night they tend to be a much paler yellowish-tan dorsally with prominent black spots in the shoulder region. Measurements of Holotype(inmm). SVL 26.4; shank 15.0; HW 10.8; head length 10.4; upper eyelid 3.0; IOD 2.6; tympanum length 1.4; eye length 4.4; E-N 3.3. Etymology. The specific epiphet is derived from the Latin, meaning a lover of darkness, in reference to the restriction of E. tenebrionis to primary forest. Natural History. Eleutherodactylus tenebrionis 1s restricted to primary forest, where it is most frequently found at night on low vegetation along stream courses. The lowland rain forests in western Ecuador which this species inhabits are characterized by full cano- pies and relatively sparse understories. Epiphytes are especially prominent in these wet forests, and some of the lowland forests appear superficially like cloud forest. Much of this region is covered with clouds during the dry season which encourages this lush growth even though annual rainfall is only moderately heavy (Dod- son and Gentry 1978). The preference of E. tenebrionis for streamside vegetation appears to be real and not a sampling bias. One of us (KM) has spent considerable time working study plots in primary forest at Rio Palenque located on top of a plateau and lacking any streams and has never encountered FE. tenebrionis there. Along the small creeks elsewhere in the Rio Palenque forest they are encountered regularly, if not commonly. The specimens from Centinela likewise all came from along a small forest stream rather than from the primary ridgetop forest or the recently cleared hillside forest. At Tinalandia 1979 TWO NEW SPECIES OF ELEUTHERODACTYLUS 11 occasional individuals were found perched on low vegetation within 2 m of the ground along a hillside about 30 m above stream level, but this area at 800 m elevation is more mesic than the lower localities. DISCUSSION The flared lips and long snouts of E. crenunguis, E. latidiscus, E. muricatus, and E. rubicundus are in marked contrast to the condi- tion seen in E. tenebrionis (Fig. 2). All of these species share the narrow IOD, a relatively uncommon trait among Eleutherodactylus (Fig. 2). Low cranial crests are present in most of these species (except E. muricatus) but are generally evident only in adult females. These traits are also exhibited by E. cruentus (Peters), a species sometimes confused with E. /atidiscus which may be allied to E. crenunguis and E. latidiscus (Lynch 1976). As noted by Lynch (in press) only E. crenunguis, E. latidiscus, and E. rubicundus of the rubicundus assembly have emarginate digital pads. We can now add E. muricatus to this list but we are not convinced that these four species are more closely related to one another than any is to the other two known species of the assembly. The weakly emarginate digital pads of EF. tenebrionis are not convincing evidence of its relationship with the rubicundus assembly; its snout physiognomy suggests that it is allied elsewhere. ACKNOWLEDGEMENTS Lynch’s fieldwork was supported by the University of Nebraska Research Council and Miyata’s by the Barbour Fund of the Museum of Comparative Zoology, the Anderson and Richmond Funds of the Department of Biology, Harvard University, the Museu de Zoologia of the Universidad de Sao Paulo, and Earth- watch and the Center for Field Research of Belmont, Massachu- setts. Facilities were provided and specimens were loaned by William E. Duellman, Museum of Natural History, University of Kansas (KU), Ernest E. Williams, Museum of Comparative Zool- ogy, and Roy W. McDiarmid, National Museum of Natural History (USNM). We are grateful to Fernando Ortiz and Eugenia del Pino of the Universidad Catolica del Ecuador in Quito for logistic sup- port in the field. For permission to collect specimens we would like to thank Calaway Dodson and Alcides Marmol (Centro Cientifico Rio Palenque) and Alfredo and Tina Garzon (Tinalandia). 12 BREVIORA No. 457 oii : LITERATURE CITED } Dopson, C. H. AND A. H. GENTRY. 1978. Flora of the Rio Palenque Science Center. Selbyana, 4(1-6): xxx + 628 p. LyNcH, J. D. 1976. New species of frogs (Leptodactylidae: Eleutherodactylus) from the Pacific versant of Ecuador. Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 55:1—33. 1977. A new frog (Leptodactylidae: Eleutherodactylus) from the Pacific lowlands of Ecuador. Copeia, 1977:282-284. In press. A new species of Eleutherodactylus from northern Ecuador (Amphibia: Leptodactylidae). Proc. Biol. Soc. Washington. In press. Taxonomic and distributional notes on poorly-known frogs (Eleutherodactylus: Leptodactylidae) from the chocoan lowlands of South America. Herpetologica. Lyncu, J. D. AND L. TRuUEB. In press. A new species of Eleutherodactylus from the cloud forests of western Ecuador (Leptodactylidae) Copeia. MIYATA, K. In press. A new species of Ate/opus (Anura: Bufonidae) from the cloud forests of northwestern Ecuador. Breviora. LIDAR Y MAR 1 8 4985 BRE VIO Re& Museum of Compa “ative Zoology US ISSN 0006-9698 CAMBRIDGE, MASS. JANUARY 31, 1980 NUMBER 458 “A NEW SPECIES OF ATELOPUS (ANURA:BUFONIDAE) FROM THE CLOUD FORESTS OF NORTHWESTERN ECUADOR KENNETH MIYATA! ABSTRACT: Atelopus covnei is described from the Rio Faisanes in Pichincha Province, Ecuador. It appears to have an extremely restricted distribution and may be in danger of extinction as its habitat 1s altered by man. The new species is most closely to Atelopus mindoensis INTRODUCTION The western slope of the Andes in northern Ecuador has a rich anuran fauna, much of which has been described only within the past decade. Until recently access to this area has been difficult due to its precipitous terrain and dense blanket of montane forest. Many Species appear to have restricted distributions and it is not surpris- ing that they were missed by early collectors. Within this forested zone, Which extends up to at least 3000 m elevation, there are several distinct asemblages of frogs. Maximum species diversity apparently occurs between 1000 and 1600 m. Many of these species are very restricted in distribution; few are known from outside Ecuador. My field work in this region revealed the presence of an apparently new Species of frog of the genus Are/opus that seems to exhibit an excep- tionally restricted distribution even in comparison with other ele- ments of this fauna. Despite extensive collecting efforts in the cloud forests of western Ecuador by myself and field parties from the Museum of Natural History of the University of Kansas, this new Species remains known from only two localities within a few kilome- ‘Kenneth Miyata, Museum of Comparative Zoology, Harvard University Cambridge, Massachusetts 02138 2 BREVIORA No. 458 ters of each other. Given the rather precarious status of these habi- tats as human development encroaches, | take the opportunity here to describe this species before the populations become extinct. Figure |. Dorsal view of holotype of Atelopus coynei (MCZ 91444), 1979 A NEW SPECIES OF ATELOPUS 3 Figure 2. Ventral view of holotype of Atelopus coynei (MCZ 91444). Atelopus coynel sp. nov. Holotype: MCZ 91444, an adult male, one of a series collected on the banks of the Rio Faisanes where it crosses Ecuador Highway 28 (the road from La Palma to Quito via Chiriboga), 14.4 km from the junction with Highway 30 (the Aloag to Santo Domingo de los 4 BREVIORA No. 458 Colorados road) at La Palma, Pichincha Province, Ecuador, 1380 m, on II July 1976 by Godfrey Guynn, Kay Harker, Steven Kaal, Ken Miyata, David Paul, and Harrison Weed. Paratyvpes. Topotypes: MCZ 91445-91449, 96775-96756, col- lected with the holotype; MCZ 91450, collected on 7 August 1976 by Jerry Coyne and Ken Miyata; MCZ 95411, collected on 8 January 1978 by Lauren Cardullo, Andrea Dion, Ken Miyata, Hugh Tor- bert, and Lisa Schwadron; MCZ 95676, collected on 30 April 1978 by Paul Greenfield and Ken Miyata; MCZ 96754, collected on 12 November 1977 by Ken Miyata; USNM 211171, collected on 17 February 1979 by Roy McDiarmid. From 4 km E Dos Rios, Pichin- cha Province, Ecuador, 1140 m: KU 164744, collected on 2 April 1975 by William Duellman. Diagnosis. A small Atelopus (males to 23 mm, females to 32 mm) distinguished from all other known species by the following combination of characters: |.) Hind limbs relatively long, the heels overlapping slightly when held parallel to femora at right angles to the body and reaching or just falling short of the orbit when adpressed. 2.) First finger almost entirely buried in a thick, rather fleshy webbing. 3.) Ventral pattern consisting of a sparse network of fine dark reticulations on a light cpaque background. Description. Head narrower than body, somewhat longer than wide. Snout projecting past tip of lower jaw, rounded from above. Projecting snout forming fairly sharp right angle above and in front of nostril in lateral profile. Nostrils opening laterally about 2/3 of way from anterior margin of orbit to tip of snout, directly above or slightly behind tip of lower jaw. From above, canthi diverging slightly from behind nostrils to a point just anterior of orbits where they diverge outward more abruptly. Canthus rostralis rounded with slight depression in loreal region. Interorbital space wider than upper eyelid. Tympanum hidden. Skin on head generally smooth with some very sparse and fine granulation. Dorsum finely shagreened, dorsolateral folds absent. Venter and sides with numerous plate-like folds, smallest and most distinct on throat and neck, becoming larger and less distinct on sides and towards cloaca. Pair of narrow low ridges on dorsum in parotoid region. Forearm slightly thicker than upper arm. Forefeet fleshy with thick webbing. First finger almost buried in fleshy webbing with only the tip free. Other fingers basally webbed with lateral fringes 1979 A NEW SPECIES OF ATELOPUS 5 extending past distal subarticular tubercles. Subarticular tubercles indistinct; palmar tubercle prominent, oval in shape. Males with cornified pad on inner margin of first finger. Heels overlap slightly when tibiofibulae held parallel to femora at right angles to body. Adpressed heels reach to, or fall just short of, posterior margin of orbit. Tarsal fold absent. Hind feet fleshy with extensive thick webbing to tips of all toes except the fourth which 1s free distally” Subarticular tubercles indistinct; outer metatarsal tu- bercle small and rounded. Measurements. Mean standard length for adult males is 22.6 mm, the single known adult female 1s 32.1 mm. The holotype is 22.4 mm. Table | summarizes the measurements and ratios used by Peters (1973) in his review of Ecuadorian Atfe/opus. Color in life. The dorsum of males varies from green with dark brown reticulations to dark brown with green spots and blotches. In all specimens the dorsal green becomes turquoise blue laterally. The single adult female (MCZ 96754) was bright green with sparse dark brown dorsal reticulations and no trace of turquoise blue on the sides. The ventral surfaces of the males were opaque white, occa- sionally with a yellowish wash, with a sparse network of dark brown or black reticulations (Fig. 2). The female was a bright opaque yellow ventrally with dark brown reticulations and a reddish-orange wash on the palms and soles. This same reddish-orange wash on the palms and soles was seen on one male as well (MCZ 94511), but it was not as prominent as on the female. The iris varies from golden yellow to orange-copper. Color in preservative. The parts of the dorsum which were green in life are pale lavender. The dark brown parts of the dorsal pattern remain brown, but are somewhat paler and have a reddish wash. The venter remains white in the males, but all traces of the yellow wash are lost. The female retains some of the yellow ventral color after 20 months in preservative. The ventral reticulations range from pale to medium brown. Etymology. The specific epiphet is a patronym for Dr. Jerry Coyne, whose timely financial assistance helped stave off the wolves on several occasions and allowed me to complete the description. NATURAL HISTORY The Rio Faisanes is a small mountain stream flowing through a narrow forested canyon where it crosses Ecuador Highway 28, 14.4 6 BREVIORA No. 458 km from the village of La Palma on the road to Quito. The elevation of the stream where it crosses the road is given as 1380 m ona topographical map of the area prepared by the Instituto Geografico Militar of Ecuador in 1969 (Alluriquin quadrat, CT-NIII-A3, 3893- 111). This falls into the “bosque muy humedo Pre Montano” eco- logical zone (Instituto Geografico Militar 1977), and the forest is characterized by a relatively low canopy height with an extremely heavy epiphyte growth. The Rio Faisanes 1s rarely more than 5 min width and few places are more than 0.5 m deep. The bottom consists of pea-sized gravel and the bed sometimes flows over or against large rocks. The water normally runs clear and in most places the canopy completely overhangs the water (see below). The other known locality where coynei has been collected (4 km E Dos Rios, 1140 m) is along the Rio Orito and 1s located about 1.3 km NSW of the type locality. According to the topographical map, the elevation is 1280 m. The two small rivers flow into the Rio El Transito, which then flows into the Rio Pilaton a few kilometers to the SW. All of the specimens taken on II July 1976 were collected along the banks of the Rio Faisanes. Individuals were captured in the late afternoon while they were active on the rocky banks of the river during a light rain. At night additional specimens were collected sleeping on streamside vegetation, usually within | m of the ground and frequently on leaves overhanging the water. A total of 41 speci- mens was collected that day in approximately 18 person-hours of collecting along a 250 m stretch of the river. Most of these speci- mens were used for bioche-nical analysis of possible skin toxins by Harrison Weed and were not available for this description. Subsequent visits to the Rio Faisanes in July and August of 1976, May and November of 1977, Janu_ry and April of 1978, and Febru- ary of 1979 resulted in the collection of very few additional speci- mens. All of these later visits were made at night and hence no more active individuals were found. All of these specimens were sleeping on the tops of leaves within | m of the ground. These specimens were not taken along the Rio Faisanes proper, but were found on the banks of a small tributary creek which crosses the road about 50 m SW of the Rio Faisanes bridge. Since the first visit to the type locality in July of 1976 the Rio Faisanes seems to have become more silted as a result of small-scale logging operations and the canopy has been opened to a considera- ble degree in places. On the first visit the river was running clear despite the fact that it was high from the rains. On later visits the 1979 A NEW SPECIES OF ATELOPUS 7 Table |. Measurements and ratios of Atelopus coynei (as in Peters 1973). Males (N=7) Female (N=1) Standard distance 22.6 mm 32.1 mm Knee-knee 20.3 29.2 Tibofibula 10.6 15.5 Head width 6.6 8.7 Head length 7.8 10.5 Knee-knee, standard distance 87.7 91.0 Tibiofibula/ standard distance 47.1 48 3 Head length) standard distance 34.4 B2E7 Head width) head length 84.5 82.9 Tibiofibula/ head length [Sve 147.6 Tibiofibula/ knee-knee S225 53.1 water was often cloudy and turbid even though the level was low and there had been no recent rains. The scarcity of frogs after the first visit may be due partly to the increase in human activity along the stream which may have rendered it unsuitable for larval devel- opment. An increase in suspended silt in the water due to this human encroachment may cause the scraping of the algae off the surface of the rocks before the tadpoles can get to it. Only a single female covnei has been found out of a total of 47 specimens (including the ones used for biochemical work). This is not necessarily indicative of a skewed sex ratio. All of the collections have been made along stream banks and McDiarmid (1971) has Suggested that male Are/opus may spend considerably more time along the creeks than females. The large number of males taken on the first visit to the type locality is indicative of a breeding concen- tration and it is likely that the females had not yet arrived. Since then, individuals have been found in low concentrations and the bulk of the population may have dispersed into the forest. The apparent absence of coynei on recent visits may also be due in part to an absence of breeding concentrations. Other anurans collected syntopically and synchronously with active Atelopus coynei include A telopus longirostris, Bufo chancha- nensis, Eleutherodactylus achatinus, E. necerus, and Colostethus sp. COMPARISONS AND DISCUSSION The relatively long hind limbs and the extensively webbed thumbs distingush A. coynei from all but a few species of Atelopus. Of these species, the hind limbs of A. elegans, A. longibranchius, A. palmatus, 8 BREVIORA No. 458 and A. rugu/osus are very long and the adpressed heel reaches beyond the anterior margin of the orbit, rather than just reaching or falling short of the posterior margin. In A. cruciger from Venezuela and A. flavescens from French Guiana the adpressed heels reach to near the posterior margin of the orbit, but they lack webbing between the second and third fingers and are larger than A. coynei (male A. cruciger to 29 mm, male A. flavescens to 32 mm). Atelopus covnei most closely resembles 4. mindoensis. The most striking difference between these two species is ventral pattern, with A. mindoensis lacking the reticulations which are characteristic of A. coyvnei. There are a number of other differences as well (Table 2) and there is little doubt that the taxa are distinct. The description of a new species of Are/opus based on external morphology from two neighboring populations may seem unwise given the known degree of variation in such characters in some members of the genus (Peters 1973, Savage 1972). The close similar- ity of A. covnel to A. mindoensis may make it seem particularly suspect. Although 4. covnei and A. mindoensis are almost certainly closely related, the differences outlined in Table 2 are consistent enough to warrant their recognition. Although color pattern can be quite variable both within and between populations of some species of Ate/opus, the ventral patt- ern of A. covnei is unique and allows the immediate recognition of Table 2. Atelopus covnei and mindoensis compared ATELOPUS COYNEI Venter white to yellow with brown reticulations. Ventral skin with numerous small scale-like folds anteriorly. Ventral skin completely opaque. Eggs not visible through skin in one gravid female known. No tubercles or enamelled pustules on back or sides. Snout more rounded from above and coming to sharp angle from side. No evidence of throat pouch in adult males. Larger size; adult males average 22.6 mm, adult female 32.1 mm. ATELOPUS MINDOENSIS Venter reddish brown with yellow spots. Ventral skin smooth or with small folds on throat. Ventral skin translucent to transparent. Eggs clearly visible through skin in gravid females. Small tubercles and enamelled pustules on back and sides Snout more pointed from above and coming to more rounded angle from side Adult males with loose skin on throat. Smaller size; adult males average 18.7 mm (N=37), adult females 24.9 mm (N=22). 1979 A NEW SPECIES OF ATELOPUS 9 this species. | have examined the holdings of Ate/opus in the MCZ collection and made a literature survey of all of the 40 presently recognized species (those listed by McDiarmid 1971 and subse- quently described species; descriptions consulted in lieu of speci- mens include Bokermann 1962, Boulenger 1902, 1903, Cochran and Goin 1970, Donoso-Barros 1969, McDiarmid 1973, Noble 1921, Peters, 1973, Ruiz-Carranza and Hernandez-Camacho 1978, Sav- age 1972) and the only species that exhibit a reticulate ventral patt- ern are A. chiriquiensis, A. longirostris, A. pachydermus, A. pulcher, A. tricolor, and A. varius. In each of these species the reticulate pattern, when present (some species are variable in this character), 1s very bold and consists of thick lines mixed with blotches and spots. In contrast, the ventral reticulations of A. coynei are thin and sparsely distributed. Although the degree of this ventral marbling in A. covnei is variable, it is immediately recognizable in all of the specimens I have examined. Although the 40 known species of Ate/opus make it a relatively large assemblage of frogs, all of which are Neotropical, the species are distributed with remarkably little geographical overlap. Exten- sive sympatry between species of Ate/opus is unusual, a situation which is in striking contrast to that seen in other Neotropical anuran genera of similar diversity. The rather specialized breeding require- ments of Ate/opus may somehow inhibit overlap. In areas where two species of Ate/opus occur in sympatry they are usually of quite dif- ferent habitus. To date, A. covnei has been collected with the larger and more gracile A. /ongirostris but has not been found with the similar A. mindoensis. Although A. mindoensis is known from eleva- tions between 20 m and 2100 m, the lower records are from the drainage of the Rio Cachabi in Esmeraldas Province. On the Rio Toachi drainage in Pichincha Province A. mindoensis appears to be restricted to elevations above 1500 mand does not come into contact with either 4. coynei or A. longirostris. Since A. mindoensis and A. longirostris do come into contact in the Rio Guayllabamaba drain- age just to the north of the Toachi, it appears that 4. covnei may be limited to the relictual populations described here. ACKNOWLEDGEMENTS The field work resulting in the discovery of Ate/opus coynei was funded by the Richmond and Anderson Funds of the Department 10 BREVIORA No. 458 of Biology, Harvard University, the Barbour Fund of the Museum of Comparative Zoology, The Museu de Zoologia, Universidade de Sao Paulo, Brasil, and by Earthwatch and the Center for Field Research of Belmont, Massachusetts. For assistance and compan- ionship in the field I would like to thank J. Bell, L. Burnham, L. Cardullo, L. Chanler, J. Coyne, A. Dion, P. Greenfield, S. Gross, G. and R. Guynn, M. Harker, S. Kaal, D. Low, R. McDiarmid, W. Morrow, D. Paul, E. Schupp, L. Schwadron, and G. Vigle. | would also like to thank W. E. Duellman (KU), R. 1. Crombie and W. R. Heyer (USNM) for the loan of material. E. E. Williams read and commented upon the manuscript. LITERATURE CITED BOKERMANN, W. C. A. 1962. Una nueva especie de Are/opus del nordeste de Brasil (Amphibia, Salientia, Brachycephalidae). Neotropica, 8(26):42-44. BOULENGER, G. A. 1902. Descriptions of new batrachians and reptiles from the Andes of Peru and Bolivia. Ann. Mag. Nat. Hist., ser. 7, 10(59):394-402. 1902. Descriptions of new batrachians in the British Museum. Ann. Mag. Nat. Hist., ser. 7, 12(71):552-557. COCHRAN, D. M. AND C. J. Goin. 1970. Frogs of Colombia. U.S. Nat. Mus. Bull., 288:1-655. DONOSO-BARROS, R. 1969. Una nueva especie de Atelopodidae de Bolivia (Anura). Physis, 28(77):327-330. INSTITUTO GEOGRAFICO MILITAR. 1977. Atlas Geografico de la Republica del Ecuador. Quito, Ecuador. 82 pp. McDIARMID, R. W. 1971. Comparative morphology and evolution of frogs of the Neotropical genera Ate/opus, Dendrophrvniscus, Melanophrvniscus, and Oreophrynella. Los Angeles County Mus. Nat. Hist., Science Bull. 12:1-66. 1973. A new species of Ate/opus (Anura, Bufonidae) from northeastern South America. Los Angeles County Mus. Nat. Hist., Contr. Sci., 240:1-12. Nosie, G. K. 1921. Five new species of Salientia from South America. Amer. Mus. Nat. Hist. Novitates, 29:1-7. Peters, J. A. 1973. The frog genus Are/opus in Ecuador (Anura: Bufonidae). Smiths. Contr. Zool., 145:1-49. RuIZ-CARRANZA, P. M. AND J. 1. HERNANDEZ-CAMACHO. 1978. Una nueva especie colombiana de Arelopus (Amphibia: Bufonidae). Caldasia, 12(57): 181-197. SAVAGE, J. M. 1972. The harlequin frogs, genus Are/opus, of Costa Rica and western Panama. Herpetologica, 28(2):77-94. MUS. COMP. ZOOL LIBRARY BREVIO BA UNIVERGITY Museum of Comparative ZLoology US ISSN 0006-9698 CAMBRIDGE, MASS. JANUARY 31, 1980 . NUMBER 459 A NEW SPECIES OF DENDROBATES (ANURA: DENDROBATIDAE) FROM THE LOWLAND RAIN FORESTS OF WESTERN ECUADOR GREGORY O. VIGLE! AND KENNETH MIYATA2 ABSTRACT: Dendrobates eryvthromos sp. nov. is described from the lowlands of western Ecuador. The new species is tentatively assigned to the genus Dendrobates on morphological grounds and appears to be most closely allied to the Colombian D. ingeri. This species may have specific microhabitat requirements which result in a spotty distribution. INTRODUCTION In January of 1978 several specimens of an apparently unde- scribed species of dendrobatid frog were collected at the Centro Cientifico Rio Palenque in the Pacific lowlands of Ecuador. Prepa- ration of an account of the herpetofauna of the region requires that a name be assigned to this population, and here we describe the species on the basis of material in the Museum of Comparative Zoology (MCZ) and the National Museum of Natural History (USNM). The generic partitioning of dendrobatid frogs is in a state of flux and the characters which have traditionally been used to define the genera are being supplanted by new ones. Following the criteria of Savage (1968) and Silverstone (1976) this new species would be asigned to Phyllobates. However, this genus has recently been defined as a monophyletic group (in the cladistic sense) on the basis of a biochemical synapomorphy by Myers et al. (1978) and consists of only four of the species recognized by Silverstone plus one newly 'Gregory O. Vigle, Biology Department, Eckerd College, St. Petersburgh, Florida 33733. *Kenneth Miyata. Museum of Comparative Zoology,’ Harvard University, Cambridge, Massachusetts 02138. 2. BREVIORA No. 459 described species. All of the remaining species of Phyllobates recog- nized by Silverstone (the femoralis, pictus, and trivittatus groups) were reassigned to Dendrobates. The new species most closely resembles those in Silverstone’s pictus group on the basis of size, coloration, and the absence of toe webbing and we tentatively assign it to the genus Dendrobates. Dendrobates erythromos sp. nov. Holotype: MCZ 96384, an adult female collected at the Centro Cientifico Rio Palenque, 47 km S of Santo Domingo de los Colora- dos, Provincia Pichincha, Ecuador, 170 m, on 17 January 1978 by Gregory O. Vigle. Paratypes. Allare topotypes: MCZ 96381-83, 96385, 94896 (lot of 3 tadpoles from back of 96385), USNM 211169-70. Diagnosis. A dendrobatid frog with the following combination of characters: |.) Teeth present. 2.) Omosternum present. 3). First Figure 1. Paratype of Dendrobates ervthromos (MCZ 96381). The slightly paler area on the dorsal surface of the forearm is the bright orange flash mark. 1979 A NEW SPECIES OF DENDROBATES 3 finger longer than second. 4.) Toe webbing absent. 5.) Muscle tissue flecked with black pigment. 6.) In life, the dorsum is dark brown and the venter is black with pale blue marbling. 7.) A bright reddish- orange flash mark on the posterodorsal surface of the upper arm in life. 8.) Size medium, to 23.5 mm snout-vent length (SVL). The above combination of morphological characters, combined with the lack of any dorsal or dorsolateral striping and the bright orange flash marks distinguish Dendrobates erythromos from all other known dendrobatid frogs. Description. Snout-vent length to 23.4 mm; four adult females 21.5-23.4 mm; two adult males 20.4-22.6 mm; one immature female 18.6 mm. Skin smooth to very finely pitted dorsally, slightly rugose laterally and on posterior surface of thighs; smooth on venter and limbs. Head as wide as body or slightly wider, widest at jaw articulations. Eyes prominent, diameter about one and one-half times the length of the snout; width of upper eyelid less than interorbital width. Snout short, truncate in dorsal and lateral aspect. Canthus rostralis rounded; loreal region flat to slightly concave. Tympanum round, about one-half diameter of orbit; posterodorsal portion concealed. Relative lengths of appressed fingers 3>4>2>1, each having a distinct disc expanded about 1.5 times the width of the distal end of the adjacent phalanx. First finger longer than second if not ap- pressed. A large outer metacarpal tubercle and a smaller, less promi- nent inner metacarpal tubercle at the base of the palm. One slightly prominent subarticular tubercle on fingers one and two and two slightly less prominent tubercles on fingers three and four; all are low with rounded surfaces. Relative lengths of appressed toes 4>5>3>2>1. Toes have distinct discs approximately 1.5 times the width of the distal end of the adjacent phalanx. A small elongated outer metatarsal tubercle and a smaller, round inner metatarsal tubercle at the base of the foot. Indistinct subarticular tubercles on toes; one On toes one and two, two on toes three and five, and three on toe four. A slightly prominent curved tarsal fold on the inner side with a weakly developed tubercle at the proximal end. Palms and soles flattened on bottom and slightly fleshy. Digits lacking lateral fringes (or with very narrow ones on toes), webbing, and supernu- merary tubercles. 4 BREVIORA No. 459 Palatine bones absent. Maxillary and premaxillary teeth present. Omosternum present. Muscle tissue flecked with black pigment. Paired vocal slits in adult males; single subgular vocal sac. Measurements of holotype. The undissected holotype is the largest individual in the type series. The following measurements are to the nearest 0.1 mm as measured on dial calipers. Snout to vent 23.4: tibia from heel to fold of skin on knee 9.6; greatest width of body 8.4: greatest width of head 8.5; head length from tip of snout to angle of jaws 7.6; length of snout from anterior edge of orbit to tip of snout 2.4; diameter of orbit 3.6; diameter of tympanum 1.9; length from proximal edge of large palmar tubercle to tip of third finger 5.7; width of disc of third finger 0.6; width of distal end of adjacent phalanx 0.4. Color in Life. Dorsum dark brown, appearing almost black in some specimens. The sides are black or very dark gray. Bright reddish-orange flash marks on the posterodorsal surfaces of the upper arms, extending from the axilla to the elbow and sometimes slightly beyond to the forearm. Anterior surface of arms and hands dark brown. The dorsal surfaces of the hind limbs are banded dark brown and black. The throat and ventral surface of the body are black with pale blue marbling which extends slightly up the sides. The iris is brown. Color in Preservative. The dorsal and lateral surfaces are black to very dark brown. The banding on the dorsal surface of the hind limbs is obscure. The flash marks on the upper arm are white. The ventral surface is black with dull gray marbling. Tadpoles. The following description is based on the paratypic lot of tadpoles (MCZ 94896) taken from the back of an adult male (MCZ 96385) on 18 January 1978. Only two of the three tadpoles are in good condition and these are the ones used for the descrip- tion. The tadpoles are in stage 25 (Gosner 1963) and measure 9.04-9.60 mm total length (measurements taken with ocular micrometer). The head and body measure 3.20-3.68 mm and constitute 35-38% of the total length. The body is somewhat flattened with the width about 1.25 times the depth of the body. The eyes and nostrils are directed dorsolaterally. The spiracle is sinistral and the anus is dextral. 1979 A NEW SPECIES OF DENDROBATES 5 The tail has a relatively low fin and is as deep as the body or slightly deeper posteriorly; the dorsal fin extends forward slightly past the level of the anus. There is one row of pigmented teeth on the anterior labium. The beak is keratinized, the upper jaw slightly convex anteriorly and the lower jaw in a strong “V” shape. The oral disc is laterally indented. There is a single row of small, blunt papillae which occur continu- ously along the lateral and posterior edges of the oral disc and are absent from the anterior edge. The ground color is pale buff, heavily speckled with brown on the dorsal surface of the body. The venter is lightly pigmented ante- riorly and heavily speckled posteriorly with brown. The brown speckling on the tail is restricted to the dorsal aspect and the fins are unmarked. Etymology. The specific epiphet is derived from the Greek ery- thros, meaning red, and omos, meaning upper arm, in reference to the distinctive reddish-orange flash marks on the posterodorsal sur- face of the upper arm. Natural History. Dendrobates erythromos is known only from the Centro Cientifico Rio Palenque. The forest there is considered to be tropical wet forest in the sense of Holdrige (1967) although available clmatic data do not seem to match this physiognomy (Dodson and Gentry 1978). The frogs are apparently restricted to a single creek system within this patch of forest. All of the specimens have come from along the banks of Lodo and Sherd Creeks above trail 3a (see Dodson and Gentry 1978:1x for map of station). The forest along these creeks has a somewhat broken canopy with signif- icant amounts of bamboo and other second-growth vegetation mixed in with the tall trees. The relief is rather steep and much of the openness of the canopy appears to be due to tree falls, although there has been some cutting in this area. The creeks flow over rocky or sandy beds and parts of them become dry during the end of the dry season. Dendrobates erythromos has been taken along these stream beds in January, when water levels were still low prior to the onset of the rainy season. Several specimens were collected along a dry portion of the bed of Lodo Creek although the majority were taken near flowing water. The frogs were never observed in the water but came from areas near the stream with good deposits of leaf litter and fairly heavy understory vegetation. They seemed to 6 BREVIORA No. 459 prefer those areas with relatively closed canopies, avoiding the numerous light gaps along the streams. All of the frogs of this species were taken during the day as they were active on the surface of the litter and ground. They were quite wary and difficult to approach and when pursued they sought shel- ter under the litter and in dense thickets of understory vegetation. Calls tentatively associated with this species were soft repetitive chirps. Three other species of dendrobatids are known from the Centro Cientifico Rio Palenque. They are Dendrobates espinosai, D. histri- onicus, and Colostethus infraguttatus. Only the much smaller D. espinosai has been found syntopically with d. erythromos, although D. histrionicus is common in the adjacent forest away from the creeks. DISCUSSION Dendrobates erythromos most closely resembles D. ingeri, a spe- cies known only from a single locality in Amazonian Colombia. The two species are very similar morphologically, both sharing maxil- lary and premaxillary teeth and a light spot in the axilla and upper arm and both lacking lateral stripes. The dorsum of D. ingeri is very granular and it has a light calf spot and is easily distinguished from D. erythromos. The two species are probably closely allied; Silver- stone (1976) placed D. ingeri closest to D. pictus within his pictus group. Although D. pictus does have lateral stripes and some speci- mens lack premaxillary and maxillary teeth, this arrangement does seem to have validity. The tadpoles of D. erythromos resemble those of D. pictus rather closely; those of D. ingeri are unknown. A fourth species, D. parvulus, was also associated with D. pictus by Silver- stone, but it seems to be somewhat more distantly allied with the other three species. Silverstone believed that his pictus group might not be a natural group and recognized three different assemblages within it. The three species he associated with D. pictus, along with D. erythromos, do seem to constitute a group of similar organisms, but further information regarding calls and skin toxins will be necessary before the relationships can be worked out. ACKNOWLEDGEMENTS We thank the following for financial support of our fieldwork: Museu de Zoologia, Universidade de Sao Paulo, Brasil; the Barbour 1979 A NEW SPECIES OF DENDROBATES Wf Fund of the Museum of Comparative Zoology; and Earthwatch and the Center for Field Research of Belmont, Massachusetts. For help in the field we thank Bruce Jayne, Roy McDiarmid, Eugene Schupp, and participants in the Earthwatch programs; for permi- sion to collect we thank Calaway Dodson and Alcides Marmol. We are grateful to Joseph Kronbichler, John Lynch, Charles Myers, José Rosado, Susan Rhodin, and Ernest Williams for assistance of varied sorts: LITERATURE CITED Dopson, C. H. AND A. H. GENTRY. 1978. Flora of the Rio Palenque Science Center. Selbyana, 4(1—6):1-628+ xxx. GosnerR, K. L. 1963. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16:183-190. HotpripGe, L. R. 1967. Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica. 206 pp. Myers, C. W., J. W. DALy, AND B. MALKIN. 1978. A dangerously toxic new frog (Phyllobates) used by Embera Indians of western Colombia, with dis- cussion of blowgun fabrication and dart poisoning. Bull. Amer. Mus. Nat. Hist., 161:307-366. SAVAGE, J. M. 1968. The dendrobatid frogs of Central America. Copeia, 1968: 745-776. SILVERSTONE, P. A. 1976. A revision of the poison-arrow frogs of the genus Phyllobates Bibron in Sagra (Family Dendrobatidae). Nat. Hist. Mus. Los Angeles Co., Sci. Bull., 27:1-53. LIBRARY MAR 1 8 1985 BREVI O'R-A Museum of Comparative Zoology US ISSN 0006-9698 CAMBRIDGE, Mass. JUNE 30, 1980’ NUMBER 460 yi ON THE EVOLUTION OF THE JAW ADDUCTOR MUSCULATURE IN PRIMITIVE GNATHOSTOME FISHES GEORGE V. LAUDER, JR.! ABSTRACT. The primitive condition of the adductor mandibulae musculature in gnathostomes ts a large unsubdivided, fan-shaped muscle mass originating from the palatoquadrate and inserting on the lateral aspect of the mandible. Unspecialized suborbital fibers extend posteriorly from the palatoquadrate to insert laterally on the lower jaw, and an intramandibular adductor division is absent. The Actinopterygii, Actinistia, Dipnoi, and Choanata primitively possess three distinct components of the adductor mandibulae: a suborbital division, a medial division, and a posterolat- eral division, as well as medial intramandibular adductor fibers. The suborbital division of the adductor mandibulae has been lost in teleosts and, independently, in coelacanths and living lungfishes. Devonian lungfishes and early choanates possessed both the suborbital and intramandibular adductor muscle components. INTRODUCTION Towards the end of the nineteenth and in the first half of the twentieth century, a large number of investigators were concerned with the homologies of the jaw musculature between the different vertebrate classes (Allis, 1897, 1917, 1923; Edgeworth, 1935; Kes- teven, 1942, 1943, 1944; Lightoller, 1939; Luther, 1913; Souche, 1932; Vetter, 1874). The musculature of the mandibular arch was the subject of special attention and several attempts were made to precisely link each branchial arch muscle with its serial homologue on the mandibular arch [see Lightoller (1939) for an example of this procedure carried to an extreme]. In recent years, little work has been done on the evolution of the musculature in primitive gnathos- tomes, despite the discovery of significant new fossil material that allows more accurate reconstruction of the musculature in extinct taxa. "The Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138 2 BREVIORA No. 460 Lauder (1980) has proposed that the primitive condition of the jaw musculature in the Actinopterygii (ray-finned fishes) is a tripar- tite division into distinct components: an anterior or suborbital division, a medial division, and a posterolateral division. In this paper, the consideration of adductor muscle components, including the intramandibular division, is extended to other primitive gna- thostome taxa to elucidate the evolution and homologies of the gnathostome jaw adductor musculature. COMPARATIVE ANATOMY Elasmobranchiomorphi The adductor musculature in the primitive living selachian Chla- mydoselachus anguineus has been described by Allis (1923) and Smith (1937). Based on their descriptions and on the comparative anatomy of the adductor mandibulae in other living sharks, the adductor musculature in the primitive Devonian selachian Clado- dus has been reconstructed (Fig. 1A). In living selachians, the adductor musculature is divided into two main components. The adductor mandibulae proper [= quadrato- mandibularis of Kesteven (1942) and Lightoller (1939)] is a single unsubdivided muscle mass having a fan-shaped origin on the palato- quadrate. The fibers converge to a tendinous inscription before fan- ning out ventrally to insert on the lower jaw. The adductor insertion is on the lateral aspect of the mandible and no intramandibularis division is present. A suborbitalis muscle [= pterygoideus of Kesteven (1942), adduc- tor £ of Vetter (1874), levator labii superioris of Allis (1923), preor- bitalis of Luther (1913)] originates tendinously from the “ectethmoidal process” (Allis, 1923) of the cranium anteriorly and inserts onto the upper labial cartilage and the skin of the lip. In Cladodus this muscle is represented as distinct suborbital fibers of the adductor mandibulae originating from the palatoquadrate (Fig. 1A: SO). Some fibers may also have originated from the nasal capsule, as in living sharks, and inserted onto the labial cartilages, when present. 1979 EVOLUTION OF JAW MUSCULATURE 3 Figure |. A: Reconstruction of the adductor mandibulae musculature in a primitive shark (Cladodus). Osteological features after Gross (1938). B. Reconstruc- tion of adductor mandibulae musculature in an acanthodian (Acanthodes). Osteological features after Jarvik (1977) and Miles (1973). Abbreviations: AM, adductor mandibulae; MD, mandible; PQ, palatoquadrate; SO, suborbital fibers of the adductor mandibulae (separated into a distinct muscle in Cladodus). 4 BREVIORA No. 460 In many living sharks, but not in Chlamydoselachus, a small superficial muscle overlies part of the adductor mandibulae. This muscle, the adductor mandibulae superficialis of Edgeworth (1935), adductor y of Smith (1937), Vetter (1874), and Allis (1923), origi- nates from the skin overlying the adductor mandibulae and inserts onto the posterior aspect of the palatoquadrate and mandible. There is currently no evidence to indicate whether the lack of this muscle in Chlamydoselachus is a specialization or if the primitive elasmobranchiomorph condition (absence of this muscle) has simply been retained. This muscle represents a small superficial component of the main adductor muscle, and I have reconstructed Cladodus (Fig. 1) with an unsubdivided adductor. Acanthodii The structure of the adductor mandibulae in acanthodians appears to have been very similar to that hypothesized to be primi- tive for elasmobranchiomorphs (Fig. |). The large adductor mandib- ulae had a fan-shaped origin from the palatoquadrate and a lateral insertion on the mandible. An intramandibular division is hypothe- sized to have been absent since the “adductor fossa” is located on the lateral aspect of the mandible and the medial surface is convex in shape (Jarvik, 1977; Miles, 1973). The adductor mandibulae is proposed to have extended ante- riorly ventral to the orbit (Fig. 1B: SO); the suborbital fibers origi- nating from the palatoquadrate and extending posteriorly to join the other adductor fibers at a tendinous inscription before inserting on the mandible. Actinopterygil In ray-finned fishes, the adductor mandibulae complex may be divided into three separate divisions each inserting on the lower jaw, and an intramandibular adductor component (Fig. 2; Lauder, 1980). Suborbital adductor fibers are present in Amia, Lepisosteus, and Polyodon. In Amia, these suborbital fibers have been called the levator maxillae superioris by Allis (1897) while the homologous adductor component in Lepisosteus is the palatomandibularis minor and major. Polyodon possesses a large separate anterior adductor 1979 EVOLUTION OF JAW MUSCULATURE 5 division originating on the palatoquadrate (Danforth, 1913). The suborbitalis adductor component has been lost in all teleosts (Fig. 2: character 6; also see Winterbottom, 1974). In palaeoniscoid actinopterygians, the suborbital fibers were enclosed in the anterior part of the maxillary—palatoquadrate chamber (Schaeffer and Rosen, 1961) and extended posteroven- trally to insert in the mandibular fossa. The medial adductor division represents one of two components of the primitive unsubdivided postorbital adductor fibers. Homolo- gies of the medial division in actinopterygians are given in Lauder (1980). The medial adductor fibers in Po/ypterus are called the “pte- rygoideus” and “temporalis” muscles by Allis (1922). The “pterygoi- deus” of Allis (1922) is not homologous to the “pterygoideus” of Kesteven (1942), which represents the suborbital division. A distinct posterolateral adductor component is also hypothes- ized to be primitive for actinopterygians (Lauder, 1980). This mus- cle represents the superficial lateral adductor fibers which run dorsoventrally from the palatoquadrate to insert in the adductor fossa. An intramandibular adductor division is present in most teleosts and in Amia, Polypterus, Polyodon, and palaeoniscoids (as inferred by the presence of an adductor fossa). The presence of intramandib- ular fibers is thus proposed to be a primitive actinopterygian charac- ter (Fig. 2: character 4). Actinistia Millot and Anthony (1958) describe three adductor muscles in Latimeria. A superficial adductor arises posteroinferiorly from the quadrate and inserts on the mandible in the mandibular fossa. This muscle appears to represent the posterolateral adductor division. The two remaining adductors arise from the ethmosphenoid por- tion of the cranium posterior to the orbit. Millot and Anthony (1958) note that these muscles appear to correspond to the “ptery- goideus” and “temporalis” adductors of Po/ypterus, and thus are both comparable to the medial adductor complex in primitive acti- nopterygians. No anterior (suborbital) division is present in Latime- ria; all jaw adductors extend dorsoventrally behind the orbit. A large intramandibular adductor occurs in Latimeria (Millot and Anthony, 1958: Fig. 19). 6 BREVIORA No. 460 GNATHOSTOMATA Figure 2. The distribution of various character states of the adductor mandibulae musculature in gnathostomes superimposed on one hypothesis of gnathostome phylogeny. For other characters supporting this phylogeny see Liem and Lauder (in press). Character |: fan-shaped adductor mandibulae with unspecialized suborbital and postorbital fibers and a lateral insertion on the mandible; 2: suborbitalis muscle differentiated from suborbital fibers; 3: medial insertion of the adductor mandibulae on the mandible; 4: intramandibular adductor division present; 5: postorbital adductor component divided into medial and posterolateral divisions; 6: loss of suborbital adductor component; 7: specialization of suborbital fibers into the levator maxillae superioris 3 and 4 muscles (see Allis, 1897); 8: loss of the suborbital adductor component; 9: loss of suborbital component in living forms. Dipnoi While extant dipnoans lack both intramandibular and suborbital adductor divisions, Devonian lungfishes appear to have possessed both. A space between the infraorbital bones laterally and the pala- toquadrate medially may have housed the anterior adductor (subor- bital) division, and an adductor fossa for the intramandibular adductor is found on the medial surface of the lower jaw (see Miles, 1977). 1979 EVOLUTION OF JAW MUSCULATURE 1 The main adductor mass in living lungfishes is divisible into pos- terolateral and medial components. The adductor mandibulae pos- terior (Edgeworth, 1935; = retractor anguli oris of Luther, 1913), corresponds to the posterolateral adductor division, and lies lateral to the ramus maxillaris and mandibularis V. The adductor mandib- ulae anterior (Edgeworth, 1935; = temporalis of Luther, 1913) is located anteromedially to the posterior adductor. In adult lung- fishes the adductor mandibulae anterior is a large muscle with a broad postorbital origin from the skull. In larval lungfishes (Edge- worth, 1935), the anterior adductor is nearly equal in mass to the posterior adductor and lies medial to it, and lateral to the trabeculae. Choanata Olson (1961) and Thomson (1967) both provide partial recon- structions of the adductor mandibulae complex in primitive cho- anates. A suborbital component of the adductor mandibulae was located anteriorly and presumably had its origin from the palato- quadrate. This division was housed in a chamber formed between the palatoquadrate medially and the dermal skull bones laterally. Postorbitally, the lateral wall of the suborbital adductor chamber was formed by the postorbital and jugal bones; while anteriorly the prefrontal, jugal, and maxilla enclosed the suborbital adductor. The posterolateral and medial adductor divisions both appear to have been present (Thomson, 1967) and thus the configuration of the adductor mandibulae in primitive choanates closely resembles that of palaeoniscoid actinopterygians. An intramandibular adductor di- vision is inferred to have been present by the occurrence of an adductor fossa in the lower jaw (see Romer, 1966). DISCUSSION In figure 2, the various configurations of the adductor mandibu- lae complex are superimposed on one hypothesis of gnathostome phylogeny. Other characters supporting this hypothesis may be found in Liem and Lauder (in press). The primitive configuration of the adductor mandibulae in gnathostomes is hypothesized to be a single unsubdivided fan-shaped muscle mass with unspecialized sub- orbital fibers originating anteriorly from the palatoquadrate (Fig. 2: character 1). Elasmobranchiomorphs retain the fan-shaped adduc- tor configuration but possess specialized suborbital fibers (Fig. 2: 8 BREVIORA No. 460 character 2); these fibers originate, in most cases, from the nasal capsule. A fan-shaped adductor muscle is also hypothesized for acanthodians (Fig. 1B), and the main postorbital adductor mass is reconstructed as being unsubdivided. One notable aspect of the adductor mandibulae in elasmo- branchiomorphs and acanthodians is its lateral insertion on the lower jaw. This condition contrasts sharply with the medial adductor insertion in actinopterygians, actinists, dipnoans, and choanates (Fig. 2: character 3). The medial insertion correlates with the presence of an intramandibularis adductor division (Fig. 2: character 4) and the occurrence of an adductor fossa on the medial surface of the lower jaw, in contrast to the lateral fossa of elasmo- branchiomorphs and acanthodians. In the Actinopterygii, Actinistia, Dipnoi, and Choanata, an hypothesis of a tripartite division of the adductor mandibulae into suborbital, medial, and posterolateral components best fits the embryological data and morphological relationships of the mus- culature in the adult (Fig. 2: character 5). Although among living taxa, Polypterus lacks the suborbital adductor fibers and Lepiso- steus the intramandibular division, other primitive actinopterygians (including palaeoniscoids) possess or show evidence of three extramandibular adductor components. Teleost fishes have lost the suborbital adductor division (Fig. 2: character 6) but halecomorphs (Amia) retain specialized derivatives of the suborbital musculature, the levator maxillae superioris 3 and 4 of Allis (1897) (Fig. 2: character 7). The cylindrically-shaped suborbital muscles of Amia are not morphologically homologous to the similarly shaped and located suborbitalis muscle of elas- mobranchiomorphs. Both muscles, however, are derived from the same geographical area of the adductor anlagen, and are thus homologous as suborbital fibers of the adductor mandibulae. Coelacanths retain both the medial and posterolateral adductor components but have lost all suborbital fibers (Fig. 2: character 8). The primitive condition for dipnoans and choanates is the presence of well-developed suborbital, medial, and posterolateral adductor divisions as well as an intramandibular section. Devonian dipnoans and early choanates appear to possess a very primitive configuration of the jaw adductor musculature, similar in all essential features to that of palaeoniscoids. Living lungfishes have lost al! suborbital adductor fibers (Fig. 2: character 9). 1979 EVOLUTION OF JAW MUSCULATURE 9 Based on this analysis, the following classification of jaw adductor muscle states is proposed. The primitive gnathostome condition consisted of a jaw adductor composed of unspecialized suborbital and postorbital components. The presence of both suborbital and postorbital fibers defines the “fan-shaped” condition. Suborbital fibers occur in elasmobranchiomorphs as a distinct suborbitalis muscle while the postorbital component is divided into two divisions in actinopterygians, dipnoans, and choanates. The mandibular component represents the third adductor component. It lies lateral to the mandible in elasmobranchiomorphs and acanthodians, and medial to it in all other groups (Fig. 2). ACKNOWLEDGEMENTS During this study I was supported by an NIH Predoctoral Fellowship in musculoskeletal biology (5T32 GM077117) and bya Junior Fellowship in the Society of Fellows, Harvard University. I thank Drs. W. Fink, K. Liem, R. Winterbottom, E. O. Wiley, and H.-P. Schultze for comments on the manuscript and Dr. R. S. Miles for his communication on lungfish anatomy. LITERATURE CITED ALLIS, E. P. 1897. The cranial muscles and cranial and first spinal nerves in Amia calva. J. Morph. 12:487-772. _. 1917. Homologies of the muscles related to the visceral arches of the gnathostome fishes. Quart. J. Microsc. Sci. 62:303-406. 1922. The cranial anatomy of Polypterus, with special reference to Polypterus bichir. J. Anat. 56:180-294. 1923. The cranial anatomy of Chlamydoselachus anguineus. Acta. Zool. 4:123-221. DANFORTH, C. H. 1913. The myology of Polyodon. J. Morph. 24:107-146. EDGEWworTH, F. H. 1935. The Cranial Muscles of vertebrates. Cambridge Univ. Press, London. 493 pp. Gross, W. 1938. Das Kopfskelett von Cladodus wildungensis Jaekel. 2. Teil: Der Kieferbogen. Senckenbergiana 20:123-145. JARVIK, E. 1977. The systematic position of acanthodian fishes. pp 199-225. In Andrews, S. M., Miles, R. S., and Walker, A. D. (Eds.), Problems in Vertebrate Evolution, Academic Press, London. Kesteven, H. L. 1942. The evolution of the skull and the cephalic muscles. Part I. The fishes. Mem. Aust. Mus. 8:1-62. 1943. The evolution of the skull and the cephalic muscles Part I. The fishes (continued). Mem. Aust. Mus. 8:63-132. 1944. The evolution of the skull and the cephalic muscles. Part. II. The Amphibia. Mem. Aust. Mus. 8:133-236. 10 BREVIORA No. 460 LAUDER, G. V. 1980 Evolution of the feeding mechanism in primitive actino- pterygian fishes: a functional anatomical analysis of Polypterus, Lepisosteus, and Amia. J. Morph. /63. LieM, K. F. AND G. V. LAUDER. in press. The evolution and interrelationships of the actinopterygian fishes. Jn Fish Neurobiology and Behavior. R. Davis and R. G. Northcutt (Eds). University of Michigan Press. LIGHTOLLER, G. H. S. 1939. Probable homologues. A study of the comparative anatomy of the mandibular and hyoid arches and their musculature. Trans. R. Soc. Lond. 24: 349-445. LuTHER, A. 1913. Uber die von N. trigeminus versorgte Muskulatur der Ganoiden und Dipneusten. Acta. Soc. Scient. Fenn. 4/:1-72. MiLes, R. S. 1973. Relationships of acanthodians. pp. 63-104. /n Interrelation- ships of Fishes, Greenwood, P. H., Miles, R. S., and Patterson, C. (Eds.), Academic Press, London. 1977. Dipnoan (lungfish) skulls and the relationships of the group: a study based on new specimens from the Devonian of Australia. Zool. J. Linn. Soc. 6/:1-328. MILLoT, J. AND J. ANTHONY. 1958. Anatomie de Latimeria chalumnae 1. Squelette, muscles, et formations de soutiens. Centre National de la Recherche Scientifique, Paris. Otson, E. C. 1961. Jaw mechanisms: rhipidistians, amphibians, reptiles. Am. Zool. 1:205-215. RoMER, A. S. 1966. Vertebrate Paleontology. University of Chicago Press, Chicago, 468 p. SCHAEFFER, B. AND D. E. ROSEN. 1961. Major adaptive levels in the evolution of the actinopterygian feeding mechanism. Am. Zool. /:187-204. SMITH, B.G. 1937. The anatomy of the frilled shark Chlamydoselachus anguineus Garman. Bashford Dean Memorial Volume VI pp 333-505. American Museum of Natural History, New York. SoucHE, G. 1932. Morphologie comparative des muscles élévaterrs de la mandibule chez les poissons. Mem. Soc. Scient. Phy. Nat. de Bordeau 7e Serie 3:1-292. THOMSON, K. S. 1967. Mechanisms of intracranial kinetics in fossil rhipidistian fishes (Crossopterygii) and their relatives. J. Linn. Soc. (Zool.) 46:223-253. VeETTER, B. 1874. Untersuchungen zur vergleichenden Anatomie der Kiemen- und Kiefermuskulatur der Fische. I. Selachien. Jena, Zeitschr. Bd. 8:405-458. WINTERBOTTOM, R. 1974. A descriptive synonymy of the striated muscles of the Teleostei. Proc. Acad. Nat. Sci. Phil. /25:225-317. ee, : —4> : a = a . a ‘ose - 4 5 ey. = > BR E VeboioR A LIBRARY Museum of Com MARAE Z, 0 ology US ISSN 2006 TT ry —NIVERSIT CAMBRIDGE, MaAss. JUNE 30, 1980 NUMBER 461 GEOGRAPHIC VARIATION IN ANOLIS BREVIROSTRIS (SAURIA: IGUANIDAE) IN HISPANIOLA DouGLas L. ARNOLD! ABSTRACT. The nominal Hispaniolan anole species, Anolis brevirostris Bocourt, is considered to be comprised of four distinct parapatric sibling species: 4. breviros- tris, A. caudalis, A. websteri, and A. marron. These species and the subspecies of A. brevirostris are described. INTRODUCTION Anolis brevirostris Bocourt is a rather small and stocky anoline lizard that occurs within an irregular range in xeric habitats on Hispaniola and associated islets. In 1870 Bocourt named Anolis brevirostris from Haiti. The name fell into disuse and obscurity, with specimens of A. brevirostris generally being included with A. dominicensis Reinhardt and Liitken. Cochran (1941) described and named two satellite island subspecies of A. dominicensis: wetmorei from Isla Beata, Dominican Republic; and caudalis Ile de la Gonave, Haiti. Barbour (1937) first combined A. dominicensis with A. distichus Cope. This combination was followed by Mertens (1939) who resurrected brevirostris as a subspecies of A. distichus from the vicinity of Barahona in southwestern Dominican Republic. Ernest E. Williams, at the Museum of Comparative Zoology at Harvard University, recognized that some of the forms in Haiti and the Dominican Republic associated with A. distichus actually per- tain to another similar species, A. brevirostris (the differential char- acters were reported in Schwartz, 1968). Named forms correctly associated with A. brevirostris are caudalis Cochran and wetmorei Cochran. Webster and Burns (1973) used electrophoretic observa- ‘University of Kentucky, Albert E. Chandler Medical Center, P. O. Box 269, Lexing- ton, Kentucky 40536, U.S.A. 2 BREVIORA No. 461 tions on several proteins to assign mainland Haitian populations of “4. brevirostris” to three unnamed sibling species. Webster (1978a) later expanded his electrophoretic studies to include most of the remaining distribution of A. brevirostris. His accidental death left behind two near-final-form manuscripts on A. brevirostris which lack the tables, figures, and numerical data he would have prepared to give his statements full documentation. These manuscripts subse- quently became available in the Third Anolis Newsletter (Williams, editor, 1977). Thus, as understood by Webster, the brevirostris com- plex consisted of three unnamed sibling species referred to as A, B, and C, and two named satellite island subspecies. Williams (1976) referred the mainland Haitian species B to caudalis Cochran and regarded species C as nominate brevirostris. ACKNOWLEDGMENTS This paper is the result of: 1) Collections made by Albert Schwartz and parties in Hispaniola, partly under National Science Foundation grants G-7977 and B-023603 and 2) information gathered by Albert Schwartz, Ernest E. Williams, and T. Preston Webster under National Science Foundation grant B-019801X and previous grants. Webster suggested that Schwartz undertake the formal description of the sibling species presented in Webster and Burns (1973) and a discussion of the ecology and distribution of the entire A. brevirostris complex. In the fall of 1974 I began investiga- tion of the A. brevirostris complex under Schwartz’s direction. | am most grateful to him for his guidance, time, patience, and faith throughout the course of this project. Most of the specimens I have examined are in the Albert Schwartz Field Series (ASFS). A more limited amount of material has been borrowed from the American Museum of Natural History (AMNH), Museum of Comparative Zoology (MCZ), Museum National d’Histoire Naturelle (MNHP), Florida State Museum (UF/FSM), and the National Museum of Natural History (USNM). I am grateful to the following curators and their assistants for the loan of this material: Albert Schwartz, George W. Foley, Ernest E. Williams, Jean Guibe, Walter A. Auffenberg, W. Ronald Heyer, and Ronald J. Crombie. Burt L. Monroe provided me with work space at the University of Louisville and was instrumental in facili- tating loans. 1979 GEOGRAPHIC VARIATION IN ANOLIS 3 METHODS I have not considered it worthwhile to study all the available collected specimens of the brevirostris complex since many of the older specimens are long-preserved and now much faded and since coloration and pattern play such a major role in differentiation of the various species and subspecies. I have attempted to examine all the material which might be assignable to new taxa proposed herein and have examined all specimens designated as holotypes, syntypes, and paratypes. The brevirostris complex needs fine sampling to be understood (Webster, in /itt., 1972); therefore I have restricted the type-locality of A. brevirostris. Lists of associated specimens, not from the type- locality itself, include localities and museum numbers which I have assigned to certain taxa on the basis of my examination. A list of referred specimens in Appendix | includes localities and museum numbers which I| have assigned to taxa on the basis of provenance; specimens so listed have not been examined by myself. The proba- bility is high that almost all lizards listed as referred specimens are correctly designated specifically and subspecifically, since they are delimited by fresh material that I have examined. Exceptional cases or uncertain allocations are noted in the text. Various scale counts and relationships were used by Oliver (1948) to describe Bahamian populations of A. distichus. Schwartz (1968) applied the same counts to non-Bahamian populations. Hoping that an application of these counts to A. brevirostris complex members might reveal differences other than coloration and pattern between various sibling species and their subspecies, I have followed the techniques of Oliver and of Schwartz and applied them to the mate- rial I have examined. The following data were recorded on each specimen: 1) Snout-vent length, in millimeters. 2) Number of scales across the snout at the level of the second canthal scale, reckoned from the anterior border of the orbit. 3) Number of horizontal rows of loreal scales. 4) Minimum number of scales between the supraorbital semi- circles. 5) Number of rows of scales between the interparietal scale and the supraorbital semicircles on each side [this datum written as a fraction] (e.g., 1/1 = one row of scales on each side). 4 BREVIORA No. 461 6) Number of scales bordering the postfrontal laterally (see Oliver, 1948:16, for drawings showing these scales in A. distichus). In a small number of specimens of A. brevirostris, the postfrontals may extend so far laterally as to make contact with one or two of the canthals. In such instances, I have not included the canthal as a scale in contact with the postfrontal, since the condition is anomalous. 7) Number of median (usually azygous) scales posterior to the posterior-most paramedian pair of snout scales, usually restricted to the midline from the anterior border of the postfrontals posteriorly to the anterior border of supraorbital semicircle contact, thus excluding the highly variable preoccipital region. (This count differs from those of Oliver and Schwartz but is made comparable by subtracting | from their counts — the nearly-always present preoc- cipital of A. distichus.) 8) Number of supraorbital semicircle scales in contact with the interparietal scale. This count is partly correlated with (5), the number of rows of scales between the semicircles and the interparie- tals; for instance, if the latter count is 1/1, the number of supraorbi- tal scales in contact with the interparietal will of necessity be 0/0. However, if the count of (5) is 0/0 (i.e., there are no scales between the semicircles and the interparietal) then (8) may have a fairly wide fluctuation. 9) Number of postmental scales. 10) Condition of the preoccipital region categorized as: preoccip- ital present (+); series of small scales bordering interparietal ante- riorly which may be incomplete (ss); preoccipital present but separated from interparietal by a series of small scales (ps); multiple small scales in preoccipital region (ms); preoccipital present, but tiny (t); and preoccipital absent (—), supraorbital semicircles in con- tact with anterior border of interparietal. 11) Condition of nuchal patch, categorized as large, medium, reduced, or absent; dark or faint; with or without white border. A medium-sized nuchal patch is reckoned to approximate the size of the external ear opening. The above counts and notations were taken on 770 specimens from Haiti and the Dominican Republic. Test counts taken on the number of subdigital lamellae on phalanges II and III of the fourth toe of the hind foot and the number of scales in tail verticils were discontinued as no significant variation was observed. Complete 1979 GEOGRAPHIC VARIATION IN ANOLIS 5 scale counts for each specific and subspecific samples are given in Table 1, where frequencies and modes are also shown. The reader is referred to this table for details of scutellar variation. A few scale counts as modes, not as absolute counts, have proven to be useful in defining the species and subspecies. The degree of overlap between various counts for different samples is generally extensive, so that it is impossible to identify the species and subspe- cies of a particular specimen solely on the basis of any set of counts. Therefore reliance must be placed on such features as dewlap pat- tern and coloration, nuchal patch pattern, and coloration and pat- tern of the head and body in general. These additional data were taken from series having field notes on coloration. (As mentioned above, museum specimens as such have very limited utility.) The colors of many dewlaps were noted in the field with reference to Maerz and Paul (1950). THE ANOLIS BREVIROSTRIS. COMPLEX As noted in the Introduction, the brevirostris complex is presently understood to consist of three sibling species. The material on which the name A. brevirostris was based had its provenance merely as “Haiti”. I have examined the three syntypes loaned by the Museum National d’Histoire Naturelle in Paris and find that two are A. distichus. The remaining one is A. brevirostris and has similarities to those lizards which are currently called “Species C”. With the recognition of new species within the brevirostris complex, it is appropriate to restrict the type-locality of A. brevirostris in order to clarify my concept of the species; I hereby designate the vicinity of Fond Parisien, Departement de Ouest, Haiti, as the type-locality of A. brevirostris. It is not unlikely that the original specimens, col- lected by M. Braconnier, actually came from the vicinity of Port-au- Prince, for the Haitian capital has long been a prominent Caribbean seaport. No brevirostris complex anoles have been collected recently from Port-au-Prince, yet sibling species A, B, and C have all been collected within 20 km airline of that city. Therefore I have chosen to designate the type-locality of A. brevirostris in the known distri- bution of Species C. Henceforth in this paper, A. brevirostris will refer only to Species C and “A. brevirostris” will refer to the com- plex in general. No. 461 BREVIORA = | = = pe t I 9 81 17 a L G 6 cl G cl (%€9) TE I (1p) ps (%7S) ITI $9 (%6b) IL C 14 81 97 6l /| fl (%TP) €1 tp I¢ (%6P) 88 97 : — I E a = G € (a iS PT (%7S) 91 PY (26SS) LTI (209) 001 IS (%E€S) LZ tl (29) 78 C6 Sz (2619) 68 = = I I — ¢ = 3 ¢ == I = t iS I P Ol 9 (OLE) 8P OS oI 97 8 9 cP 19 PT cr (2069) €€ (2619) 61 PE (Cotr) O01 %LL) LEI (ors) 6L fe = | _ a a IS If Ie] (ALG 6L1 orl 14aisqam UOAADUL syppnoy 1940] 9M DJOINAaSap STISOMIAIAG y y y q°¥ ‘GK ‘q-¥ JO 1OQUINU=N *xa[dWOd sijsOdiAadg SIJOUP JY) JO SajouR Jo suOTR[Ndod xIs Jo sjUNOD a]eds YBIA 1OJ BIeP NSAP [eqoued -13]Ul pue SO[IITIOTWIS uaamjaq SaJBIS SMOI [e210] jnous ssoloe Sa[BIS N ‘sayeos yeJuaujsod jo Jaquinu Joy ATUO paiedipul aie (X) suBaY ‘sasayUaIed UT PazeoIpUl sI (sapow -11] JO -Iq 10) UONTPUoD [epow ay) sulAvy a[dwes ay} Jo adejusoJad ay) PUR ‘SdI]RI UI JS JIB SAPO, “SUNOS [BOIIIIA BUIMOT[OJ 94) Ul J9]JRI19Y} PIISIPUl satouanbady 9Y} YIM “19}OVIVYO YORI Ul UONPLIBA IY} SAIVSIPUL UTNIOS [BOIVIIA ISIY ay] ‘paulwexa suowidads ‘T 91981 GEOGRAPHIC VARIATION IN ANOLIS 1979 Z = I = = = ¢ I (6 L S Z € bp I G lig 91 S 9I € rl (%Z€) Ol (ore) br 8S (%6€) 69 I€ Z %Se) BI (%7€) OI Se (%€b) O01 (%6£) 69 (%€r) €9 I Sa[Bos S I LI €¢ cf Ze 0 peay ueipsy I = == I == = S/P = == = = I = ¢/Z I = (6 L I Z p/Z — I = Z Z iS p/P S I ¢ SI Z 91 b/€ rl ¢ ZI (%S€) 08 LI (rs) 6L e/€ 6 9 7 St IZ 1€ €/Z 198 ]U09 (%IP) 17 (%9S) 8I (%89) 88 (%S€) 08 (%SL) €€1 01 Z/Z [equosyjsod = = =e = se G Z/I [e1aqe] = I = I ¢ I €/Z “= I b I IZ (6 z/Z — 9 L Ol v7 ¢ Z/I [eqaued L S Ze 67 LE LI 1/1 -19]UT Yum Z b 81 4 LI 81 1/0 19B]U09 (%78) @% (%Sr) v1 (%€S) OL (%LS) 9S1 (%0r) ZL (%69) 001 0/0 UI SaTBIS (panunuos) [| 3qey No. 461 BREVIORA Z li (9p) L9 9¢ 1€ re 4 S Z el ra fb 9 sd 9 I el Sl 9 €Z su %L9) v€ (%s¢S) LI Se (SS) 8ZI (%t@pr) SL (%7S) SL s uolsal Z GE p 9 ll } jeudrios0a1d Z Z b SI 6P 9 jo uonIpuo) SiS 9°¢ Lv ZG 8°¢ Lv x I I S 6 Z Z | 9 9] = 8 li ¢ b PZ €¢ 9 ib (%v7) ZI (467) 6 €Z ZS (2697) 9b rZ 9 (%vZ) ZI (60) 6 tr (%S€) 78 (2%97) 9b tr S (%bZ) ZI S (%0r) 7S 09 o¢ (%Lpr) 89 p tp I b S = Z € , = z Z G s]BJUSUNISOd MI ISGIAM YWUOAADLE S1]DPNDdD IAMOLUTAM DJOIUAISAP STAISOATNAAG 4 Vy i GV TaN Z Ga (panunuos) | aqeyp 1979 GEOGRAPHIC VARIATIION IN ANOLIS 9 “ANOLIS BREVIROSTRIS” NATURAL HISTORY Anoles of the brevirostris complex and A. distichus are usually allotopic, but broadly sympatric. However, in certain localities where the generally xeric habitat of “A. brevirostris” intergrades with the generally mesic habitat of A. distichus, the two are precisely syntopic. In these situations hybridization is known to occur, to varying degrees, producing in the analyzed cases sterile offspring (Webster, 1978b). “4. brevirostris” is cryptic. A generalized, composite head and dorsal pattern for the entire complex consists of: barred limbs and tail, two longitudinal dorsolateral stripes, an interocular bar, inter- parietal U, interocular V, and a series of five to seven dorsomedian chevrons pointed posteriorly. The first chevron is an internuchal patch. The upper longitudinal stripe extends from the angle of the jaw posteriorly to the level of the scapulae. Localized emphasis of this stripe forms the nuchal patch and scapular stripe. The lower longitudinal stripe extends from beneath the jaw across the shoulder to the hindlimbs. The pigmentation for the median dorsal stripe with chevrons is darker than the region below the upper dorsolateral stripe. All the markings are in general, faint with poor contrast, and interrupted outlines. The largest specimens of “A. brevirostris” are from inland north- ern Haiti. In the Vallee de l’Artibonite in the vicinity of Dessalines and Pont Sonde, males reach a snout-vent length of 51 mm and females 47 mm. In general, females of all populations reach a maxi- mum size about 4 to 8 mm less than do males. The smallest speci- mens are from Isla Beata off the coast of the Peninsula de Barahona, where males reach a snout-vent length of only 43 mm and females 38 mm. “4. brevirostris” has an extensive but narrow distribution on Hispaniola. It occupies situations varying from mesic moderate ele- vations (Sierra Martin Garcia, 915 m) to extreme xeric regions (Cul de Sac-Valle de Neiba plain, at or below sea level). Typically an inhabitant of xeric habitats, “A. brevirostris” prefers the larger trees of xeric woods and flourishes in the proximity of springs and in the conspicuously taller Acacia around villages. It is present through much of the low (often dense) Acacia scrub between the more favor- able habitats. 10 BREVIORA No. 461 In the mesic regions along the east coast of the Peninsula de Barahona, “A. brevirostris” is frequently observed on fenceposts in variegated sun and shade situations as well as on trees. In addition, “4. brevirostris” has been collected on vines, shrubs, trees, and ona cliff face 90 m above the ocean. In the extreme xeric region of the Cul de Sac-Valle de Neiba plain it is more frequently associated with Cocos, being found diurnally on the ground under palm logs, stacks of palm fronds, in coconut piles, and in Cocos trash. It is possible that in this region “A. brevirostris” resorts to such situations for nocturnal retreats. Generally, when it is found asleep it is one to three m above the ground on vines, shrubs, or particularly Acacia. SYSTEMATIC ACCOUNT Anolis brevirostris brevirostris Bocourt Anolis brevirostris Bocourt, 1870, Nouv. Arch. Mus. Hist., Paris 6k Lectotype. MNHP 2467B, an immature female collected by M. Braconnier. Type-locality. “Haiti”; restricted here to within 5 km of Fond Parisien, Dept. de Ouest, Haiti. Associated specimens. Haiti, Dépt. de Ouest, Leogane (MCZ 13781); Fond Michelle, 549 m (ASFS V37159); 7.4 km W. Thoma- zeau (ASFS V8156); Terre Rouge (ASFS V24266-94); La Source, Fond Parisien on edge of Etang Saumatre (ASFS V8129-31); Source Fond Parisien, 1.6 km NE Fond Parisien (ASFS V36921- 58): 3.2 km NW Fond Parisien (ASFS V36980-81); Dominican Republic, Independencia Province, Tierra Nueva (ASFS V42367- 88): 10.4 km NE Jimani (ASFS X9506); 5 km N Jimani (ASFS V35476-77); Boca de Cachon (ASFS V4379, ASFS V39719-41); Las Baitoas (ASFS V14310-22); 6 km NW Duverge (ASFS V17142-46), 5 km NNW Duvergé (ASFS X9937); just E Duverge (ASFS V23274): Los Saladillos (ASFS V41744-45). Definition. A sibling species of the A. brevirostris complex char- acterized by: moderate size (males to 48 mm, females to 45 mm snout-vent length); dorsum dark gray to medium gray and brown, dewlap pale, monochromatic, shades of orange, peach, yellow, gray, olive, and brown; nuchal patch variable, from reduced but dark, with no white border to medium and dark, with a white border; modally 1/1 scales between the supraorbital semicircles and the 1979 GEOGRAPHIC VARIATION IN ANOLIS 11 interparietal; modally 0/0 supraorbitals in contact with the interpa- rietal; modally a single row of small scales bordering interparietal anteriorly; modally 3/3 scales in contact laterally with the postfron- tals (Species C of Webster and Burns, 1973). Distribution. Hispaniola: in the Dominican Republic from Inde- pendencia Province (Las Salinas), the Valle de Neiba, west into the Haitian Cul de Sac Plain, Dept. de Ouest (Daspinasse) and an isolated population on the Cap de Leogane, Dept. de l’Ouest (Leogane). Description of lectotype. The lectotype has the following meas- urements and scale counts: snout-vent length 34 mm, tail 55 mm, four scales across snout, four loreal rows, semicircles in contact, | / 1 scales between supraorbital semicircles and the interparietal, 0/0 supraorbitals in contact with the interparietal, a single row of small scales bordering interparietal anteriorly, 2/2 scales in lateral contact with postfrontals, one median azygous head scale, six postmentals. Syntypes MHNP 2467 and 2467A are both A. distichus. Variation. Scale counts for the series of 146 specimens of A. b. brevirostris are shown in Table 1. The dorsum and markings of A. b. brevirostris vary clinally. In the east (Duverge), the dorsum is medium gray-brown. The interoc- ular bar is present. The interparietal U and interocular V are often present. Dorsomedian chevrons are occasionally found. Both dorso- lateral stripes are present in reduced form. The upper remains as a scapular stripe and a nuchal patch is medium and dark, with a white border. The lower dorsolateral stripe is irregularly interrupted. The limbs and tail are barred. The more western the population, the more vague are the head and body markings, and the darker and more uniform gray the dorsum. At the western extreme the interpar- ietal U and the dorsomedian chevrons are absent. The tail and limbs are vaguely barred. The upper dorsolateral stripe is reduced, often to a scapular spot and a reduced, dark, borderless nuchal patch. Three geographical regions of dewlap variation in A. b. breviros- tris are discernible. At the western extreme (Daspinasse to Thoma- zeau) dewlaps are pale, monochromatic in hues of gray, brown, yellow, yellow-gray, cream, peach, and rarely white. In the southern Cul de Sac Plain and Valle de Neiba the dewlaps are predominately pale orange and peach. Dark and bright orange dewlaps have been noted in the vicinity of Fond Parisien and Duverge. Bicolored dew- laps, pale orange with a thin pale yellow margin, are found within 12 BREVIORA No. 461 this region from Las Salinas to 10 km west of Duverge. A third region, the northern Valle de Neiba between Etang Saumatre and Lac Enriquillo, has predominately pale yellow and yellow-brown dewlaps. Hues noted for A. b. brevirostris are: Pl. 916, Pl. 9K5, PI. 14Cl, and Pl. 14HS. The isolated population on the Cap de Léogane is represented by a single specimen (MCZ 13781) collected by G. M. Allen, August 1919. The dorsum is a medium to dark brown with a dark colored and medium sized nuchal patch with a white border. The limbs and tail are barred. The scapular stripe is present, but other markings are obscured by the dark dorsum. Scale counts for the Leogane A. b. brevirostris are: six scales across the snout, four loreal rows, supraorbital semicircles in contact, 1/1 scales between semicircles and interparietal, 0/0 supraorbitals in contact with interparietal, 4/3 scales in lateral contact with postfrontals, four median head scales, four postmentals, and preoccipital present but separated from interparietal by a row of small scales. Anolis brevirostris deserticola, new subspecies Holotype. MCZ 132391, an adult male. Type-locality. 2.1 km S San Jose de Ocoa, Peravia Province, Dominican Republic one of a series taken 19 November 1971, col- lected by natives. Original number ASFS V34034. Paratypes (all from Peravia Province, Dominican Republic). ASFS V34031-33, ASFS V34035-67, same data as holotype; ASFS V724-25, 1.8 km S San Jose de Ocoa, 24 August 1963, A. Schwartz; ASFS_ V15615-25, San Jose de Ocoa, 3 August, 1968, native collectors. Associated specimens. Dominican Republic, La Estrelleta Prov., 2.5 km NW El Llano, 365 m (ASFS V31510-12); San Juan Prov., 6.8 km NW Juan Herrera, 490 m (ASFS V31401); 1.7 km NW Sabaneta, 670 m (ASFS V31394); 10.2 km S Las Matas de Farfan, 550 m (ASFS V31327, ASFS V31468); 7.8 km NW Vallejuelo, 732 m (ASFS V31312); | km W Sabana Alta (MCZ 125547-48); SE Sabana Alta (MCZ 125554-55); 3 km W Guanito (MCZ 128236-47); 15 km SE San Juan (ASFS V412, ASFS V470-86); Azua Prov., Padre las Casas (MCZ 58474-76); Sierra Martin Garcia, Mt. Busu, 610-854 m (ASFS V31214-24); Sierra Martin Garcia, above Barre- ras, ca. 915 m (ASFS V21174-202); Barreras, 10 km SW Puerto Viejo (ASFS V21110-16); Peravia Prov., 3-5 km S San Jose de 1979 GEOGRAPHIC VARIATION IN ANOLIS 13 Ocoa (ASFS V21396-97); 14.2 km S San Jose de Ocoa, 396 m (ASFS V715); 16.2 km S San José de Ocoa, 396 m (ASFS V712); 10 km S Los Ranchitos (ASFS V15627); 19 km NW Bani (ASFS V 15632); Limonal (ASFS V14921-28); Barahona Prov., west side Punta Martin Garcia (ASFS VI1I1-16); 3.2 km NE Fondo Negro (ASFS X9688); 4 km SE Canoa (ASFS V40785); 15 km ESE Canoa (ASFS V40796); 0.8 km NE Cachon, 122 m(ASFS V30749): 7.5 km E Cabral (ASFS X9604-05); ca. 3 km E Cabral (ASFS V20540-41); 11.8 km S Cabral, 701 m (ASFS X9849-53); 12.3 km S Cabral, 640 m (ASFS X9834-37). Definition. A subspecies of A. brevirostris characterized by: mod- erate size (males to 48 mm, females to 45 mm snout-vent length); dorsum medium gray-brown to medium brown; venter white; dew- lap bicolor, red-orange basal spot, margin cream or pale yellow to pale, or monochromatic, peach, salmon, or yellow; nuchal patch large and dark, with white border; modally 0/0 scales between the supraorbital semicircles and interparietal; modally 0/0 supraorbitals in contact with the interparietal; modally one or more rows of small scales bordering interparietal anteriorly; modally 2/2 scales in con- tact laterally with the postfrontals. Distribution. Dominican Republic Valle de San Juan (El Llano, La Estrelleta Prov.), east onto the southern slopes of the Sierra de Ocoa and to the Lianos de Azua (Limonal, Peravia Prov.) and south onto the northern slopes of the Sierra de Baoruco (south of Cabral, Barahona Prov.). Description of holotype. The holotype has the following measure- ments and scale counts: snout-vent length 45mm, tail (broken) 48mm, four scales across snout, four loreal rows, semicircles in contact, 0/0 scales between supraorbital semicircles and interparie- tal, 1/1 supraorbitals in contact with interparietal, preoccipital absent, 2/2 scales in lateral contact with postfrontals, one median azygous head scale, seven postmentals. Variation. Scale counts for the series of 179 A. b. deserticola are shown in Table 1. The dorsum of A. 6. deserticola is medium gray-brown to medium brown. The head markings are distinct and generally include the interocular bar, parietal U, and interocular V. Dorsome- dian chevrons are absent, except for the internuchal chevron which is rare. The lower dorsolateral stripe is absent. The upper is present and extends from the scapula to the hindlimbs in an interrupted 14 BREVIORA No. 461 pattern. The nuchal patch is large and dark, with a white border. The limbs and tail are barred. The venter is white to pale yellow with pale yellow on the underside of the tail. The dewlap is uniformly pale yellow, peach, or salmon to bicolor peach or red-orange with a pale yellow or cream margin; hues noted for the dewlap are: P1. 10D8, Pl. 18F1, Pl. 17E1, with centers of P!. 5B12 and Pl. 6Bir The dorsum is a darker gray and the markings most distinct in specimens from the Sierra Martin Garcia and from the southern slopes of the Sierra de Ocoa. The browner dorsum and least distinct markings characterize specimens from the eastern Valle de San Juan (San Juan). Seven specimens collected west of San Juan are noteworthy. The samples are small (one to three each) and were collected in ravines peripheral to the Valle de San Juan. The preoccipital is present in 71% of the specimens—significantly different from all other A. b. deserticola. The nuchal patch is reduced to absent. Scale counts for specimens from the moderate elevations between Lago de Rincon and the Bahia de Neiba and along the northern slopes of the Sierra de Baoruco are intermediate between those of A. b. deserticola to the north and A. b wetmorei to the south. The scale counts and dewlap more closely favor A. b. deserticola and have thus been included with them. Anolis brevirostris wetmoreit Cochran Anolis dominicensis wetmorei Cochran, 1931, Proc. Biol. Soc. Washington 44:89. Anolis brevirostris wetmorei: Schwartz, 1968, Bull. Mus. Comp. Zool. 137): 257. Holotype. USNM 83881, an adult male, 13 May 1931, collected by A. Wetmore and F. C. Lincoln. Type-locality. Isla Beata, Dominican Republic. Associated specimens. Dominican Republic, Barahona Prov., Barahona (ASFS X9530-60, ASFS V14017-38, ASFS X9456-58); 5 km S Barahona (ASFS V20545-48); 6 km SE Barahona (ASFS V40787-88); 6.6 km SW Barahona, 137 m (ASFS_ V30240-43, V30418-19); 1S km SW Barahona (ASFS V40781-83); 5.3 km NE La Cienaga (ASFS X9379-81); 6.4 km SW La Ciénaga, 46 m (ASFS V39858); 9 km SW La Cienaga, 91 m (ASFS V42710); 2 km NW Paraiso (ASFS V17-24); 1 km NE Paraiso, Rio Nizaito (ASFS 1979 GEOGRAPHIC VARIATIION IN ANOLIS 15 V40762-71); | km W Paraiso (ASFS V40772-80); 7.7 km W Paraiso, 152 m (ASFS V30928-31); 17.9 km NE Caleton (ASFS V30790- 802); Los Patos (ASFS V16926); 6 km N Enriquillo, 427 m (ASFS V42190-209); 2.2 km NE Enriquillo (ASFS V14078-80); Pedernales Prov., 21 km SW Enriquillo (ASFS X9416); 8 km NE Oviedo (viejo) (ASFS X9964-65); 7 km N Oviedo (viejo) (ASFS V23121); 5 km NE Oviedo (viejo) (ASFS V14073-75); 3 km NW Oviedo (nuevo) (ASFS V40678-87, ASFS V40471-82); 7 km N, thence 20 km SE Cabo Rojo, 183 m (ASFS V29769-70); 7 km N, thence 17.6 km SE Cabo Rojo, 152 m (ASFS V30088-89); 2 km E turn to Cabo Rojo (MCZ 128257); behind Cabo Rojo police station (MCZ 143407-08); 21 km NE Cabo Rojo, 396 m (ASFS V16782-85); 21.2 km N Cabo Rojo on Alcoa mine road, 440 m (MCZ 143403-04); L’Eglise, near La Mercedes (ASFS V21510); 10 km N Pedernales, 244 m (ASFS V30115-17); 4 km SE Pedernales (ASFS V16729-30); south center of Pedernales (ASFS V29846); Pedernales (ASFS V2505-06, ASFS V2919); Isla Beata (ASFS V2670-71, UF/FSM 36969-70, MCZ 37520, MCZ 58695-702, AMNH 41414, AMNH 41416-18, AMNH 44854-58); just E Punta Beata (ASFS V17220); Haiti, Dept. de l'Ouest, near Saltrou (MCZ 49274-79); 4 km NNE Marigot, 61 m (ASFS V37372-77). Definition. A subspecies of A. brevirostris characterized by: large size (males to 50 mm, females to 45 mm snout-vent length); dorsum pale grayish tan to gray to tan to brown to mottled greenish tan; venter white to yellow; dewlap color very variable from locality to locality (see below); nuchal patch reduced and gray to brown with no white border to large and dark with white border; modally 1/1 scales between supraorbital semicircles and interparietal; modally 0/0 supraorbitals in contact with interparietal; modally a single row of small scales bordering the interparietal anteriorly; bimodally 2/2 or 3/3 scales in contact laterally with postfrontals. Description of holotype. The holotype has the following measure- ments and scale counts: snout-vent length 43 mm, tail (broken) 62 mm, five scales across snout, four loreal rows, semicircles in contact, 1/1 scales between supraorbital semicircles and interparietal, 0/0 supraorbitals in contact with interparietal, a single row of small scales bordering interparietal anteriorly, 3/3 scales in lateral contact with postfrontals, three median head scales, five postmentals. Variation. Scale counts for the series of 232 A. b. wetmorei are shown in Table 1. 16 BREVIORA No. 461 Distribution. Hispaniola: in Haiti from the Dépt. de l’Ouest (Marigot), the southern coast of the Tiburon Peninsula east across the southern portion of the Dominican Peninsula de Barahona (Enriquillo), Isla Beata, and north along the eastern coast of the Peninsula de Barahona to the city of Barahona. A. b. wetmorei varies geographically. On Isla Beata, whence wer- morei was first described, the dorsum is nearly uniform tan or light gray. In general the head and body markings are faint to absent. The interocular bar is faint and the interocular V is reduced to a short postocular stripe. The dorsomedian chevrons and dorsolateral stripes are absent. The nuchal patch is present and varies from large and dark to reduced and gray, but always with a white border. The limbs and tail are faintly barred. The venter is whitish and the dewlap is pale yellow. On mainland Hispaniola wermorei dewlaps vary from pale yellow to bicolored yellow with an orange central spot to deep orange or red-orange. However, regionally the variation is less extreme, favor- ing either the deep orange, red-orange, and bicolored orange with a thin pale margin (Enriquillo to la Cienaga and Pedernales to Can- ada la Cerca) or the pale yellow to yellow with orange blush basally characteristic of the remainder of the extensive wetmorei distribu- tion. The full range of variation has been observed at the northeast- ern distribution extreme, Barahona, and at the regional interfaces at Pedernales, La Cienaga, and south of Enriquillo. The eastern orange-dewlap wermorei have a dorsum that is pale greenish tan to tan. The markings of the head and body are cryptic to the point that the dorsum has a woodgrain pattern. The interocu- lar bar is regularly present. The interparietal U is wide and dark. Mediandorsal chevrons are occasionally present, although most often they are reduced to dorsomedian spots or are absent. The upper and lower dorsolateral stripes are interrupted to absent. The legs and tail are lightly barred. The nuchal patch is reduced and faint, often gray, tan, or reddish brown, and with no white border. Hues noted for the dewlap in this series are: PI. 1017, Pl. 10G11, Plo SRl2y Pils Bil and Plat ys: The western orange-dewlap wetmorei are similar to the eastern, but there are no reports of any green in the dorsal color. The mark- ings have greater contrast. The scapular stripe is long and there are remnants of the lower dorsolateral stripe. Pl. 10C10 is the only hue noted for the dewlap. 1979 GEOGRAPHIC VARIATION IN 4NOLIS 17 The yellow-dewlap wetmorei are in general more conspicuously marked, except in the vicinity of Oviedo where the dorsum is a darker and grayer version of wetmorei on Isla Beata. The nuchal patch for these populations is highly variable, from large and dark with a white border to reduced and faint with no white border. Additional hues noted for the dewlaps are: Pl. 1OF1, PI. 11F3, P1. PE with Pl TvBIO and Pil; 1107 centrally. Anolis caudalis Cochran Anolis dominicensis caudalis Cochran, 1932, Proc. Biol. Soc. Washington 45:185. Anolis brevirostris caudalis: Schwartz, 1968, Bull. Mus. Comp. Zoos @):251- Holotype. USNM_ 76801, an immature male, March 1929, collected by A. J. Poole and W. M. Perrygo. Type-locality. Nan Cafe, Ile de la Gonave, Haiti. Paratypes. Haiti, Ile de la Gonave, USNM 76799-800, same data as holotype; USNM 77080-82, Pointe a4 Raquettes, “1927”, W. J. Eyerdam; USNM 80388-89, 8.3-16.7 km inland Anse-a-Galets, 23 March 1930, L. H. Parish and W. M. Perrygo; Ile de la Petite Gonave, USNM 80390, 23 March 1930, L. H. Parish and W. M. Perrygo. Associated specimens. Haiti, Dépt. de l'Ouest, Trou Forban (ASFS X1939, ASFS V8204-14); 3.5 km E Trou Forban (ASFS X4001-03); 0.3 km NE Carries, Ouanga Bay Hotel (ASFS V36876- 82); 4km N Arcahaie (MCZ 125008-18); 3 km SE Arcahaie (ASFS V43778-81); 6 km N Duvalierville (MCZ 125022-37); 1.4 km SE Duvalierville (ASFS V36872-74); 6 km SE Duvalierville (ASFS V9834-36); Ile a Cabrit, 6 km airline SE Duvalierville (ASFS V9853-56, ASFS V44806-25); Source Matelas (MCZ 125019-21); Dept. du Sud, Presqwile de Baraderes, vicinity of Grand Boucan (ASFS V26342-48); 4.2km W Jeremie (USNM 59255-56); Ile de la Gonave, Source Picmi, above Picmi (ASFS V26610-15, ASFS V26652); Anse-a-Galets (ASFS V22413-22): Etroits (ASFS X2410- 15). Definition. A sibling species of the A. brevirostris complex char- acterized by: moderate size (males to 48 mm, females to 44 mm snout-vent length); dorsum medium brown to light gray-tan to light gray; venter pale yellow; dewlap highly variable from uniformly 18 BREVIORA No. 461 white, yellow or yellow-gray to bicolor yellow with red or orange centers to bicolor gray-green with rusty centers; nuchal patch medium, dark with white border; modally 1/1 scales between the supraorbital semicircles and the interparietal; modally 0/0 supraor- bitals in contact with the interparietal; modally preoccipital present; modally 2/2 scales in contact laterally with the postfrontals (Species B of Webster and Burns, 1973). Distribution. Haiti: Dept. de POuest, from Trou Forban south along the coast of the Canal de St. Marc to the Baie de Port-au- Prince (Source Matelas):; Ile de la Gonave; Ile de la Petite Gonave; and two isolated populations on the northern shore of the Tiburon Peninsula (Dept. du Sud: Presquile de Baraderes and Jeremie). Description of holotype. The holotype has the following measure- ments and scale counts: snout-vent length 42 mm, tail broken, six scales across snout, four loreal rows, semicircles in contact, 0/0 scales between supraorbital semicircles and interparietal, | / 1 supra- orbitals in contact with interparietal, preoccipital region with five small scales with an incomplete row of small scales bordering inter- parietal anteriorly, 2/2 scales in lateral contact with postfrontals, one median azygous head scale, five postmentals. Variation. Scale counts for the series of 131 A. caudalis are shown im dable: 1: The dorsum of A. caudalis is medium brown to light gray; the lightest specimens examined are from Anse-a-Galets, Ile de la Gonave. The head markings are generally absent. Dorsomedian chevrons are absent. The nuchal patch 1s large and dark with a white border. The scapular stripe is short but distinct. The remainder of the dorsolateral stripes are reduced, interrupted, and vague to absent. The limbs and tail are weakly barred. The venter is pale yellow. The dewlap color is highly variable from uniformly white, yellow, yellow-gray, or gray-green, to bicolored white or yellow with variable orange centrally to bicolored yellow-green or gray-green with yellow, mustard, rusty, or darker gray-green centers. Hues noted for the dewlap color are: PI. 12E1, Pl. 10CI, and Pl. 6K10 centrally with Pl. 10I1 and PI. 17G1 peripherally. At the northern mainland extreme of A. caudalis (Trou Forban) the dewlap is uniformly white or pale yellow to bicolored with a small orange basal spot. There is a clinal increase in the size and intensity of the orange spot the farther south along the coast (Web- ster and Burns, 1973). However, the extent of the orange, even at the 1979 GEOGRAPHIC VARIATION IN 4ANOLIS 19 southern mainland extreme (Source Matelas), is not so great as that of A. websteri. Two exceptions to this general cline are recorded. At Trou Forban, one series (ASFS V8204-14) had dewlaps gray to greenish gray with mustard or darker gray-green centers. On the Ile a Cabrit in the Baie de Port-au-Prince the dewlaps are gray-green. On Ile de la Gonave no such cline exists, and the dewlap color is variable and includes all dewlap colors described above for A. caudalis. The two disjunct populations of A. caudalis are of particular interest. Jeremie caudalis is represented by only two specimens (USNM 59255-56) collected by Henderson and Bartsch, 10 April 1917. The dorsum is a faded gray-tan, with a dorsomedian stripe and short scapular stripe. The nuchal patch is large and dark witha white border. The limbs and tail «1e barred. The head has a faint interocular bar, short ocular stripe, and a broad and dark interpa- rietal U. Scale counts for these two caudalis are: six scales across snout, four loreal rows, supraorbital semicircles in contact, 0/0 and 1/1 scales between semicircles and interparietal, 0/0 and 1/1 supraorbitals in contact with interparietal, 2/2 scales in lateral con- tact with postfrontals, no median head scales, and preoccipital present. The other disjunct population, Grand Boucan caudalis, is repre- sented by seven specimens (ASFS V26342-48) collected by Richard Thomas and natives, 4 August 1971. Of the three known disjunct populations of “A. brevirostris” on the Tiburon Peninsula, only Grand Boucan caudalis have dewlap color notes. The dewlaps are colored “pale (greenish) with faint basal dull yellow smudge”. The dorsum is medium to dark brown with a scapular stripe. The lower dorsolateral stripe is discernible. The interparietal U and interocular bar and V are present. The limbs and tail are barred. The nuchal patch is large and dark with a white border. Grand Boucan caudalis differ from mainland and Gonave caudalis in two modal scale counts: five scales across the snout and three median head scales. The preoccipital, although present, is fragmented into multiple scales in three of seven specimens. Anolis marron, new species Holotype. MCZ 124732, an adult male. Type-locality. Jacmel, Departement de Ouest, one of a series taken 15 June 1970, collected by T. P. Webster. 20 BREVIORA No. 461 Paratypes. MCZ_ 124730-31, MCZ 124733-36, same data as holotype. Associated specimens. Haiti, Dépt. de Quest, Marigot (MCZ 124723-29); 3.2 km W Marigot (ASFS V9771-78); 4.8 km E Cayes Jacmel (ASFS V9711-16); 8 km W Jacmel (ASFS V9696-97): Terre Noire, 19.2 km SW Jacmel (ASFS V37509). Definition. A sibling species of the A. brevirostris complex char- acterized by: moderate size (males to 50 mm, females to 42 mm snout-vent length); dorsum pale gray and tan to medium brown; venter pale yellow to off-white; dewlap uniformly olive-gray to bico- lor tan with reddish center; nuchal patch large and dark with faint white border; modally 0/0 scales between supraorbital semicircles and the interparietal; modally 0/0 supraorbitals in contact with the interparietal; modally one or more rows of small scales bordering interparietal anteriorly; modally 2/2 scales in contact laterally with postfrontals. Distribution. Haiti; the southern coast of the Tiburon Peninsula, from Marigot west to Terre Noire. Description of holotype. The holotype has the following measure- ments and scale counts: snout-vent length 47 mm, tail 62 mm, four scales across snout, four loreal rows, semicircles in contact, 0/0 scales between supraorbital semicircles and the interparietal, 2/1 supraorbitals in contact with the interparietal, a series of small scales bordering interparietal anteriorly, 2/2 scales in lateral contact with postfrontals, one median azygous head scale, five postmentals. Variation. Scale counts for the series of 31 A. marron are shown in table 1. The dorsum of A. marron is pale gray and brown to medium brown. A thin interocular bar is regularly present, but other head markings are vague or absent. The dorsomedian chevrons are reduced to small dorsal spots or are absent in all but the darkest specimens. The thin, dark scapular stripe broadens and becomes less distinct posteriorly. The lower dorsolateral stripe is interrupted and vague. The nuchal patch is large and dark with a white border. The limbs and tail are barred. The venter is pale yellow to off-white. The dewlap is either uniformly dark olive-gray or bicolored tan with a reddish center; hues noted for the dewlap are: PI. 15SH5, P1. 6B10, and PI. 6C10. 1979 GEOGRAPHIC VARIATION IN ANOLIS Dal Anolis websteri, new species Holotype. MCZ 132390, an adult male. Type-locality. 7.4 km NW Dessalines, Departement de I’Artibo- nite, Haiti, one of a series taken 8 July 1974, collected by natives. Original number ASFS V39323. Paratypes. ASFS V39322, ASFS V39324-37, same data as holo- type. Associated specimens. Haiti, Dépt. de Nord Ouest. Mole St. Nicholas (MCZ 63143-53); Dept. de lArtibonite, 6.7 km NW Gonaives (ASFS V39295-302); 1 km NE Pont Sonde (ASFS V39342-50); Bains de Amani-y (ASFS V36900-03, ASFS V43742): 2.6 km NW Montrouis (ASFS V36888-89). Definition. A sibling species of the A. brevirostris complex char- acterized by: large size (males to 51 mm, females to 47 mm snout- vent length); dorsum medium to dark gray and brown; venter bright yellow to yellow-orange; dewlap orange with a thin pale yellow edge; nuchal patch large and dark with no white border; modally 1/1 scales between the supraorbital semicircles and the interparietal; modally 0/0 supraorbitals in contact with interparietal; modally a single row of small scales bordering interparietal anteriorly; modally 2/2 scales in contact laterally with postfrontals. (Species A of Webs- ter and Burns, 1973). Distribution. Haiti; from the Dept. de Nord Ouest (Mole St. Nicholas) and northwestern Dept. de l’Artibonite (Marche aux Poteux; between Gonaives and Ennery), south into the Vallee de lArtibonite (Dessaliens and Pont Sonde) and along the coast of the Golfe de la Gonave to Pointe de Montrouis (Montrouis). Description of holotype. The holotype has the following measure- ments and scale counts: snout-vent length 47 mm, tail 70 mm, five scales across snout, four loreal rows, semicircles in contact, 1/1 scales between supraorbital semicircles and interparietal, 0/0 supra- orbitals in contact with interparietal, a single row of small scales bordering interparietal anteriorly, 3/3 scales in lateral contact with postfrontals, one median azygous head scale, four postmentals. Variation. Scale counts for the series of 51 A. websteri are shown in Table 1. The dorsum of A. websteri is moderate to dark gray or brown. Head and body markings, if present, are generally dark and diffuse, 22 BREVIORA No. 461 often disrupted. The nuchal patch is large and dark, with no white border. The scapular stripe is dark and short. The interocular bar is present in the lighter colored specimens, as is sometimes the case with the interparietal U and interocular V. Dorsal chevrons are always absent. The dorsolateral stripes are disrupted and diffuse. The limbs and tail are vaguely barred. The interparietal is yellow, but appears as a copper flake in preserved specimens. The venters are bright yellow to yellow-orange. The dewlap is uniformly orange or orange with a narrow yellow margin; hues noted for the dewlap are: Pl, 9L7, Pl. 9L8, Pl. 9L10; Pl. 108, and) Pie cig: A series of eight A. websteri from Pont Sonde is notable in the variation in dewlap color and modal scale counts. The dewlaps vary from orange (PI. 10L8) to mustard (P1. 12L7) to pale yellow (P1. 11L4). The venters are not so bright as other websteri. Modal scale counts differences for the series are: six scales across the snout; five loreal rows; and 3/3 scales in contact laterally with postfrontals. Well outside its normal distribution, A. websteri was found in abundance 3.5 km NW Bon Repos, Dept. de Ouest, in young native mahogany (Swietenia mahagoni Jacq.). Electrophoretic evi- dence indicates that it is slightly more similar to northern A. web- steri than to southern specimens of that species. Presumably, A. websteri in this location was a recent introduction by man to an artificial habitat. Recent road improvement has led to the total destruction of the localized habitat (Webster, 1978a). No specimens were preserved. DISCUSSION INTERACTION OF THE “ANOLIS BREVIROSTRIS’— ANOLIS DISTICHUS COMPLEXES Species of the “A. brevirostris” —A. distichus group often present contrasting dewlap colors in areas of contact or proximity. Such differences suggest that dewlap color may be important for species identification and reproductive isolation (Webster and Burns, 1973). Both body size and color appear to be important in Anolis species identification, especially in simple (e.g., two species) faunas (Will- iams and Rand, 1977). The A. brevirostris complex is closely associated with four sub- species of A. distichus: dominicensis, ravitergum, favillarum, and suppar. In areas of sympatry “A. brevirostris” and distichus gener- 1979 GEOGRAPHIC VARIATION IN ANOLIS 23 ally vary in body size, body color, and dewlap color. In brief, the associated subspecies have the following characteristics: dominicen- sis, males to 58 mm, females to 48 mm snout-vent length, dorsum bright marbled green to rich chocolate brown, dewlap usually pale yellow; ravitergum, males to 56 mm, females to 45 mm snout-vent length, dorsum ashy gray to dark tan to pale greenish, dewlap pale yellow to bicolored with an orange basal spot; favillarum, males to 54 mm, females to 47 mm snout-vent length, dorsum dark green, dewlap vivid orange centrally with narrow pale yellow edge; suppar, males to 54 mm, females to 44 mm snout-vent length, dorsum pale green, dewlap pale yellow to yellow-green, at times with a dull yellow-orange basal smudge. For more detailed descriptions, see Schwartz (1968) and Webster (1978b). In general, A. distichus 1s larger than “A. brevirostris” and the green and brown unmarked dorsa of distichus contrast with the gray and marked dorsa of “brevirostris”’. A. caudalis is the only species within the brevirostris complex which is not extensively sympatric with A. d. dominicensis. Quite to the contrary, both mainland and Gonave caudalis appear to exclude dominicensis, except for two specimens, one from Trou Forban and another from nearby Ste. Philomene. At the disjunct populations on the Tiburon Peninsula, Grand Boucan caudalis also appear to exclude dominicensis and the Jeremie population is sympatric with another subspecies, A. d. suppar. No color notes exist for the Jere- mie caudalis, nor can any be assumed. A. caudalis here is the smaller lizard and has two prominent markings. The nuchal patch is large and faint with a white border and the interparietal U is broad and dark. A. d. dominicensis is broadly sympatric with A. websteri and A. marron throughout the latters’ respective ranges. Locally domini- censis 1s precisely syntopic with websteri. A. websteri has the largest snout-vent length for the brevirostris complex and at this region of syntopy is larger than dominicensis. The reduced, borderless nuchal patch and generally vague and diffuse markings of gray-brown web- steri probably indicate an increased reliance on dewlap color for species recognition and reproductive isolation. The orange dewlap of websteri sharply contrast with the typical pale yellow of domini- censis and white to gray-yellow with greenish basally of A. caudalis. The isolaticn is incomplete, for Webster (1978b) reports a low inci- 24 BREVIORA No. 461 dence of hybridization at Montrouis with dominicensis; and (in litt.) some hybridization and introgression (two of five proteins, only one way, only a short distance) with caudalis at Trou Forban. Though populations of A. marron are never far from those of A. d. dominicensis, no areas of syntopy are known. The dark olive-gray or bicolored tan with reddish center dewlap contrast with the pale yellow of dominicensis. The dark scapular stripe and large, dark, white bordered nuchal patch of marron are more prominent dorsal markings than those of A. websteri and may provide additional cues for species recognition and reproductive isolation. A. marron and A. b. wetmorei come in contact near Marigot. The bicolored pale yellow with orange basal spot of A. b. wetmorei contrasts with the marron dewlap. ~ In the eastern Cul de Sac Plain (Thomazeau: Manneville), A. b. brevirostris 1s sympatric (but apparently not syntopic) with 4. d. dominicensis. Of the two species in the area, A. b. brevirostris is distinctly the inhabitant of the xeric scrub and A. d. dominicensis the inhabitant of more mesic situations (oases and cultivated areas). A. b. brevirostris is the widespread lizard in open areas, whereas A. d. dominicensis is restricted to certain less rigorous habitats and is in effect surrounded by A. b. brevirostris. Here, the dewlap color of A. d. dominicensis 1s atypically a deep orange (PI. 4CI1, P1. 4G10, and Pl. 4G11), at times with a faintly brown cast (Schwartz, 1968). This contrasts strongly with the dewlap of A. b. brevirostris, which locally is dully pigmented with no conspicuous pattern in white to pale shades of yellow, gray-yellow, and gray-brown. There is a clinal decrease in the amount of orange in the dewlap of A. d. dominicensis away from A. brevirostris, until at the southern edge of the Cul de Sac and Valle de Neiba plain, pale yellow is again the typical color of A. d. dominicensis dewlaps. It is important to note that in this area (Fond Parisien to Duverge) A. b. brevirostris dewlaps are predominantly pale orange, dull orange, to bright orange. Another subspecies, A. d. ravitergum, 18 sympatric with A. b. brevirostris at the latter’s eastern distribution. Webster (1978b) reports hybridization locally at Balneario la Zurza, 5 km WNW Duverge. Here the dewlaps of A. d. ravitergum range from uni- formly light yellow-orange to strongly bicolored with an orange basal spot. Dewlaps of A. b. brevirostris are all monochromatic 1979 GEOGRAPHIC VARIATION IN ANOLIS DS orange in varying degrees of intensity. Dewlaps do not appear con- sistently to differentiate these populations despite the strong dewlap color contrast. A. b. wetmorei, like A. caudalis, appears to exclude A. distichus but is locally sympatric with two subspecies of A. distichus at three localities. 4. d. dominicensis and A. b. wetmorei are sympatric at Belle Anse (=Saltrou), Dept. de POuest, and Las Mercedes, Peder- nales Province. Here, wetmorei is bicolored with an orange basal spot and a yellow or cream outer margin. A. d. dominicensis dewlap is typically pale yellow but occasional specimens in the Sierra de Baoruco in Pedernales Province have the dewlap very pale yellow to practically white. A. d. favillarum and A. b. wetmorei are sympatric at Hermann’s finca, near Paraiso, Barahona Province. No color notes exist for the single specimen of wetmorei taken from this locality. (These are, however, old records and may not imply real sympatry.) At all three of these widely separated localities, wetmorei is gener- ally the lizard of the lower elevations with distichus occupying the moderate to higher elevations (300-1250 m) of the Massif de la Selle and Sierra de Baoruco. A. b. deserticola is sympatric with one of three subspecies of A. distichus throughout most of its range. A. d. dominicensis and A. b. deserticola are widely sympatric in the Valle de San Juan from 7 km NW Vallejuelo and 15 km SE San Juan westward to the Dominico- Haitian border. The dewlaps of both are pale yellow except at Valle- juelo where deserticola is bicolored with a pale orange blush. A. d. ravitergum 1s sympatric throughout most of the remaining range of deserticola and in local situations is even syntopic (0.5 and 2.5kmE Cachon). Webster (1978b) notes the similarity in dewlaps. Both are bicolored, with a pale margin surrounding a darker or brighter spot of variable size. Corresponding to this high degree of similarity is a higher incidence of hybridization at these localities. A. b. deserticola extends onto the northern slopes of the Sierra de Baoruco, where in the intermediate altitudes and transitional vege- tation it is sympatric and locally syntopic with 4. d. favillarum. The dewlaps are here identical; both are dark orange, but the nuchal patch and gray dorsum of deserticola contrast strongly with the green dorsum of favillarum. The incidence of hybridization here has not been investigated. 26 BREVIORA No. 461 The sample examined in this study represents the known distribu- tion of the “A. brevirostris” complex. Three of the six populations recognized by scale counts correspond to the informal siblings A, B, and C of Webster and Burns (1973). A. brevirostris is restricted to Species C and the name A. caudalis is assigned to Species B as suggested by Williams (1976). The distribution of A. b. wetmorei of Isla Beata is expanded to include the Peninsula de Barahona and part of the southern coast of the Tiburon Peninsula. Species A (A. websteri), a new sibling species (A. marron), and a new subspecies (A. b. deserticola) are described. Scale counts were taken as supple- mentary evidence of geographic variation along with that in dewlap color within the brevirostris complex, and to provide a basis for the formal description of parapatric siblings. A. caudalis and each of the three subspecies of A. brevirostris overlap in dewlap color variation. Dewlaps bicolored yellow with orange centers and/or uniformly pale yellow are found to varying degrees in the distribution of each species. A. caudalis is, by modali- ties of scale counts, the most readily distinguishable of these species and of the entire brevirostris complex. Modally, has six scales across the snout (all others four), preoccipital present (all others a single row of small scales bordering the interparietal anteriorly), and two median scales (all others modally one or bimodally one and two). Webster (1978a) noted the presence of shared alleles of A. cauda- lis with A. marron and A. b. deserticola. The scale counts of the latter two populations differ from the former not only by the unique modalities of caudalis noted above, but further by the modes shared by marron and deserticola: scales between interparietal and supra- orbital semicircles 0/0 (all others 1/1); postmentals, bimodes five and six, with the highest mean values deserticola 5.8 and marron 5.6 (others, mode four or five, or trimode four, five, and six; means 4.7 to 5.3). Despite the wide geographic separation (ca. 120 km airline across the Sierra de Baoruco and Massif de la Selle at the closest points), these two populations have essentially the same scale counts. The modalities for the scale counts taken are the same, the percentages of the modalities are within three to 17% of each other, and the distribution of the non-modal counts is similar. Further, these are two of the three populations that are extensively sympatric with A. distichus. A. marron differs from A. b. wetmorei and the 1979 GEOGRAPHIC VARIATION IN ANOLIS 27 entire brevirostris complex by its unique dewlap color (dark olive- gray to tan with reddish center). A. b. brevirostris differs from other populations in the “A. brevi- rostris” complex by the unique modes of 3/3 scales in contact with postfrontals laterally (all others modally 2/2 or bimodally 2/2 and 3/3) and four postmentals, with 4.7 the lowest mean value (others modally five, bimodally five and six or trimodally four, five, and six; means 4.7 to 5.8). A. b. wetmorei differs from other populations in the brevirostris complex by the unique bimode of 2/2 and 3/3 scales in contact with postfrontals laterally. A. b. wetmorei differs from the other subspe- cies of A. brevirostris in having modally five postmentals (breviros- tris modally four and deserticola bimodally five and six), but share this characteristic with A. caudalis. Variation in scale counts and dorsum occur within 4. b. wetmorei. At Enriquillo there is a dra- matic change in vegetation and geography from the mesic slopes of the Sierra do Baoruco in the north to the arid plain that forms the southern tip of the Peninsula de Barahona. This change appears to effectively inhibit genetic exchange between A. b. wetmorei popula- tions demonstrated by Webster’s observation of different frequen- cies for variants of three proteins and the modal shift in loreal rows and the dorsum coloration differences for the two regions docu- mented by the present study. On the eastern coast of the Peninsula de Barahona (Barahona to Enriquillo) wetmorei modally has five loreal rows and a greenish tan dorsum in association with the orange, red-orange, and bicolored orange with a thin yellow margin dewlap color. To the south and west (Oviedo to Marigot) wetmorei modally has four loreal rows and no greenish dorsa have been recorded. A. websteri differs from other populations in the brevirostris complex by the unique trimode of four, five, and six postmentals (others modally four or five, or bimodally five and six). A. websteri and each of the three subspecies of A. brevirostris overlap in dewlap color variation. Dewlaps uniformly orange and bicolored orange with a thin yellow margin are found to varying degrees in the distri- bution of each species. The subspecies of A. brevirostris are further distinguishable from A. websteri by their unique modal scale counts noted above. 28 BREVIORA No. 461 The distribution presented in this paper follows closely that of Webster and Burns (1973), but varies from that of Webster (1978a). Webster noted electrophoretic similarities between A. caudalis and the Jacmel-Marigot populations and between A. caudalis and the San Juan-San José de Ocoa populations. He implied that the Jacmel-Marigot populations be referred to caudalis. Here the southwestern Tiburon populations are designated as a new species, A. marron. For the brevirostris complex, marron varies greatly from caudalis in modal scale counts, dewlap color, and in relation- ships to A. distichus. Modal scale count variations are: scales across snout (four versus five), loreal rows (five versus four), scales betwen supraorbital semicircles and interparietal (0/0 versus 1/1), median head scales (bimodes one and two versus mode two), postmentals (bimodes five and six versus mode five), and preoccipital condition (modally a series of small scales bordering interparietal anteriorly versus preoccipital modally present). The dark olive-gray to tan with reddish centered dewlap of marron distinguishes it not only from caudalis but is unique in the whole brevirostris complex. A. caudalis excludes A. distichus dominicensis where A. marron and A. distichus are sympatric. Webster also noted shared alleles of marron with A. b. wetmorei at Bell Anse. The northern Dominican populations having electrophoretic sim- ilarities to A. caudalis are designated as a new subspecies, A. b. deserticola. A. b. deserticola and A. marron have the same scale count modes, and thus the same substantial difference from cauda- lis. Webster (1978b) noted variation in hybridization of A. distichus with A. b. brevirostris from that with A. b. deserticola. However, neither this variation nor the electrophoretic similarities discour- aged Webster from associating deserticola with brevirostris. A. marron would then appear to be an extension of “A. breviros- tris’ from the Peninsula de Barahona, a pattern found in other reptilian species (Ameiva leberi, Ameiva lineolata privigna, Ameiva taeniura vulcanalis). APPENDIX’ 1 1) Anolis brevirostris brevirostris: Haiti, Dépt. de l-Ouest, Source Trou Caiman (MCZ_ 124737-49); Thomazeau (MCZ 13769-70, MCZ 59388-89, MCZ 124705); between Thomazeau and Manne- ville (MCZ 59381); Manneville (MCZ 59382-87, MCZ 63154-58, 1979 GEOGRAPHIC VARIATION IN 4NOLIS 29 MCZ 121777-78, MCZ 123125-55) Eaux Gaillées (MCZ 59943-48, MCZ 63159-71, MCZ 118790-92, MCZ 121096-101); Ganthier (MCZ 124706-22); Dominican Republic, Independencia Prov., Las Lajas (AMNH 41456, AMNH 41458-59, AMNH 41461); La Descu- bierta (AMNH 41433, AMNH 41437-38); Las Baitoas (AMNH 50149-50, AMNH 50199, AMNH 50210-16); 8.8 km W Duverge (MCZ 125541); 2 km SW Duverge (UF/FSM 42738). 2) Anolis brevirostris deserticola: Dominican Republic, Azua Prov., 4 km SW Sabana Yegua, 420 m(UF/ FSM 42739): Barahona Prov., Cabral (MCZ 58511-25, MCZ 58526-36); 5 km S Cabral (MCZ 58509-10); La Cueva (MCZ 58425, MCZ 58429, MCZ 58432, MCZ 58436); lower part of road to Polo (MCZ 125551-53, MCZ 128219-25); 7.5 km N Canoa (MCZ 125549-50); Sierra Martin Garcia, 1540 m (UF/FSM 36971); Peravia Prov., 2 km N San José de Ocoa (MCZ 107079-80); 1 km N San Jose de Ocoa (MCZ 128255-56); | km S San Jose de Ocoa (MCZ 126012-44); 2 km S San Jose de Ocoa (MCZ 128248-54); Rio Limon, 8 km S San José de Ocoa (MCZ 107081-82); 5 km N Bani, 110 m (UF/FSM 36972). 3) Anolis brevirostris wetmorei: Dominican Republic, Barahona Prov., Barahona (AMNH 41307-15, AMNH 41317, AMNH 41322, AMNH 51430, AMNH 51446, AMNH 51533, MCZ 58694, MCZ 93106, UF/ FSM 42733-35, USNM Field No RIC 040037-43); avia- tion field, Barahona (AMNH 51508-22, AMNH 51525, AMNH 51527); Hotel Guarocuya, Barahona (MCZ 107083-86); 1.7 km S Barahona (MCZ 9311); SW Barahona (AMNH 51535-40); Paraiso (AMNH_ 51630-31); Hermann’s finca, near Paraiso (AMNH 516145); Los Patos (MCZ 58559-60); Pedernales Prov., Oviedo (viejo) (MCZ 58537-38); 2 km E turn to Cabo Rojo (MCZ 128215- 18, MCZ 128258-59); 1 km SW Las Mercedes, 380 m (UF/FSM 42736); 2 km E Las Mercedes, 250 m (UF/FSM 42737); Haiti, Dépt. de l’Ouest Lan Banane, near Saltrou (MCZ 68685-90). 4) Anolis caudalis: Haiti, Dépt. de Quest, 6.3 km N Duvalierville (MCZ 123072-99); Ste. Philomene (USNM 123351-56); Ile de la Gonave, no other locality (MCZ 13783-89); Anse-a-Galets (MCZ 29046-50, MCZ 37507-16, MCZ 85299-309): Nan Café (MCZ 85225-45, UF/FSM_ 12286-1-9, UF/FSM_ 12287-1-9, UF/FSM 12288, UF/FSM 12290); Pointe a Raquettes (MCZ 25509-18, MCZ 80613-38, UF/FSM 12291-1-2); vicinity of Pointe a Raquettes (MCZ 80581-86); Ti Roche, 9.5 km Pointe a Raquettes (MCZ 80587-612); Nan Palmiste, 4 km from Pointe a Raquettes (MCZ No. 461 BREVIORA 30 ‘IVT BY 0} JaSO[O ae INQ ‘MJOIIIM4asap “Gq “¥ pure 1as0Wjam “g ‘PY UIIMIIG aeIPAWIIIU! Ie 1eY) susuioads juasaidas jeiqed JO YyINos OONIORY ap B1IIIG 941 Jo sadojs usay OU ayI ZuIpnjour pue ‘uooury op ode] puke euoyeieg UsIMIAq Sa[dII9 uado ou] ‘ouRB09T 1¥ STUJSO4IAadg “G ‘f JO 99U9I1INIIO dy} SUIMOYS SMOLIL ay) puke ‘uvonog puesH pure aiaiof¢ 1B SDN “f JO} SaTITTROO] SUIMOYS B[NsutUsd UOINGI] IYI Jo UOTIION [BISIP 94} 0} SMOLIL OM] OY) OSTP DION ‘pauayory]g Jey ISI YIM SapoIII—sasowjam “q ‘PY :pauaydelq J[BY 1a] YIM SafoIO— D/OIM1AaSaP “q “f ‘S2]PI19 PIOS— SHISOdIAadg “q “YW ‘SB PalROIPUl ae STAISOdIAaIg ‘PY JO Satsadsqns aary) BY] “se[suRIy ‘aisqan ‘suodexoy ‘WOLD ‘salenbs ‘siDPNDI ‘Sa]IIO ‘S1d4ISOAIAIAG -aue sarsads 10] sjoquiAg ‘dnoid s1ajsosiaasg SOUP IY) JO SIOQUISU INOJ 3YI JO UOTINQIISIP 24) Suimoys ‘ejotuedsiy jo dep "| oinsi4 » ov oe oz oO 1979 GEOGRAPHIC VARIATION IN ANOLIS 31 80639-58); Nan Saline (MCZ 85169-74, MCZ 85175-81); Téte Source (MCZ 85182-205); Platon Yeye (MCZ 85206-24); Lan Coupe (MCZ 85281-98); Ile de la Petite Gonave (AMNH 49745, AMNH 49747-48). 5) Anolis websteri: Haiti, Dept. de l’Artibonite, Marche aux Poteaux (MCZ 125117-25); Passe Reine, between Gonalves and Ennery (MCZ 63141-42); Pont de l-Estere (MCZ 63135-40); bridge over Riviere de l’Artibonite, St. Marc road (MCZ 59355-80); Lafond (MCZ 118783-89, MCZ 123101-24, MCZ 125110-16); St. Marc (AMNH 42593, AMNH 49696, AMNH 49699-700, AMNH 49702-04, AMNH 49708-09, AMNH 49711-12, AMNH 77566, USNM 59219); Montrouis (MCZ 125126-69). LITERATURE CITED Barspour, T. 1937. Third list of Antillean reptiles and amphibians. Bull. Mus. Comp. Zool., 82(2):77-166. CocuRAN, D.M. 1941. The herpetology of Hispaniola. Bull. U.S. Nat. Mus., 177: vii + 398 pp. MAERZ, A., AND M. R. PAu. 1950. A dictionary of color. New York, McGraw- Hill, Book Co., N.Y., vii + pp. 1-23 and 137-208, 56 pls. MERTENS, R. 1939. Herpetologische Ergebnisse einer Reise nach der Insel His- paniola, Westindien. Abh. senckenberg. naturf. Ges. (449):1-84. Ouiver, J. A. 1948. The anoline lizards of Bimini, Bahamas. Amer. Mus. Novitates (1383):1-36. SCHWARTZ, A. 1968. Geographic variation in Anolis distichus Cope (Lacertilia, Iguanidae) in the Bahama Islands and Hispaniola. Bull. Mus. Comp. Zool., 137 (2):255-309. WessTeR, T. P. 1978a. Geographic variation in “Anolis brevirostris”: evidence from proteins and a consideration of dewlap color. Third Anolis Newsletter: 153-164. 1978b. Hybridization of Hispaniolan lizards in the Anolis distichus species group. Third Anolis Newsletter: 166—170. AND J. Burns. 1973. Dewlap color variation and electrophoretically detected sibling species in a Haitian lizard Anolis brevirostris. Evolution, 27: 368-377. Wituiams, E. E. 1976. West Indian anoles: a taxonomic and evolutionary summary. I. Introduction and species list. Breviora, 440: 1-21. AND A. S. RAND. 1977. Species recognition, dewlap function and faunal size. Amer. Zool., 17: 261-270. rs MUS. COMP, ZOOL LIBRARY BREV- LOR A UNIVERSITY Museum of Comparative Zoology US ISSN 0006 9698 CAMBRIDGE, MaASss. JUNE 30, 1980 NUMBER 462 ELEUTHERODACTYLUS EREMITUS, A NEW TRANS-ANDEAN SPECIES OF THE LACRIMOSUS ASSEMBLY FROM ECUADOR (AMPHIBIA: LEPTODACTYLIDAE) JoHN D. LYNcH! Abstract: Eleutherodactylus eremitus 1s named from cloud forests along the Rios Mindo, Pilaton, and Saloya, in Provincia Pichincha, Ecuador, between 1540 and 2100 m. The new frog is most closely related to E. bromeliaceus from the Amazonian versant of the Andes in southern Ecuador. Lynch and Duellman (1980) included the following four species (all cis-Andean) in the /acrimosus assembly of the wnistrigatus group of Eleutherodactylus—E. bromeliaceus Lynch, E. lacrimosus (Jimenez de la Espada), FE. mendax Duellman, and EF. petersi Lynch and Duellman. Eleutherodactylus lacrimosus 1s distributed below 1000m from eastern Ecuador east to the mouth of the Amazon river (Lynch and Schwartz, 1972; Lynch, 1979), whereas the other three species occur on the eastern slopes of the Andes—FE. petersi, from the headwaters of the Rio Magdalena (Colombia) south to the val- ley of the Rio Pastaza (Ecuador); FE. bromeliaceus, south of the Pastaza Trench to the Cordillera de Zamora in southern Ecuador; and £. mendax, from central Peru (Huanuco) south into adjacent Bolivia. In 1970 while collecting on the Pacific slopes in Andean Ecuador, I found a small, green frog in a bromeliad. Subsequent study of the collections of the late James A. Peters provided two additional examples from nearby localities. Additional specimens were obtained by William E. Duellman, Kenneth Miyata, and myself in the course of field work between 1975 and 1978. Sufficient material is now available to provide a description of the first trans-Andean representative of the /acrimosus assembly. ‘School of Life Sciences, The University of Nebraska, Lincoln, Nebraska 68588 D) BREVIORA No. 462 Eleutherodactylus eremitus sp. nov. Holotype: MCZ 92103, a gravid female, taken on the La Palma- Chiriboga road (hwy 28), 25.7 km above (NE) La Palma, Prov. Pichincha, Ecuador, 1820 m, on 12 July 1976 by Kenneth Miyata. Paratypes (all from Prov. Pichincha, Ecuador). Topotype, MCZ 92104; 2 km W Campamento Silante (La Palma-Aloag road), 2100m, KU 140878; 5 km ESE Chiriboga, Quebrada Zapadores, 2010m, KU 179085-86, 180248; km 58, + 8 km W Chiriboga, USNM 211209; 3.5 km NE Mindo, 1540m, KU 165884; 6.2 km E Tandapi, 1750m, MCZ 92105; Tandayapa, USNM 211208. Diagnosis. 1) skin of dorsum finely areolate, that of venter coarse- ly areolate; no dorsolateral folds; no anal sheath; 2) tympanum partially visible, round, its length 2/5—1/2 eye length; 3) snout sub- acuminate in dorsal view, rounded or weakly protruding in lateral profile (papilla at tip); canthus rostralis moderately distinct; 4) upper eyelid bearing a conical tubercle, narrower than IOD; no cranial crests; 5) vomerine odontophores oval in outline, elevated; 6) males with vocal slits, vocal sac, and non-spinous nuptial pads; 7) first finger shorter than second; all fingers with broad discs, pads on II-IV expanded; numerous supernumerary palmar tubercles; 8) fin- gers bear lateral fringes; 9) 2-3 small, subconical ulnar tubercles; 10) small tubercles on knee, heel, outer edges of tarsus; 11) two metatar- sal tubercles, inner oval, 4 times size pungent outer; numerous supernumerary plantar tubercles; 12) toes bear lateral fringes, broad discs, dilated pads; pads smaller than those of fingers; 13) cream with faint brown stippling; vague canthal-supratympanic and flank stripe; colorless areas in groin and on posterior surfaces of thighs; 14) adults small, males 17.2-20.0 (x=18.9, N=6) mm, one adult female 27.6 mm SVL. Eleutherodactylus eremitus most closely resembles E. mendax but differs in having areolate skin on the dorsum and more prominent vomerine odontophores (odontophores absent or indistinct in E. mendax). Both have larger tympana than do E. bromeliaceus, E. lacrimosus, and E. petersi. Description. Head broader than body (except in gravid female), wider than long; HW 35.1-38.4 (x=36.4, N=8) per cent SVL; snout subacuminate in dorsal view with conical papilla at tip, in profile rounded or truncate but with papilla, sometimes protruding; snout moderately long, E-N in males 81.5—-88.5 (x=85.4, N=4) per cent 1979 ELEUTHERODACTYLUS EREMITUS SP. NOV. 3 eye length, in females 86.4—100.0 (x=95.6, N=4) per cent; nostrils weakly protuberant, directed dorsolaterally; canthus rostralis mod- erately distinct, straight; loreal region weakly concave, sloping grad- ually to lips, lips not flared; upper eyelid width 70.4-93.4 (x=81.5, N=8) per cent IOD, bearing one conical tubercle in center (Fig. la—b) and several low warts (areolations) posterolaterally; no cran- ial crests; tympanum moderately distinct, annulus visible through thin skin, directed dorsolaterally, round, its length 42.3-50.0 (x= 47.2, N=8) per cent eye length, separated from eye by distance equal to tympanum length; supratympanic fold indistinct amidst areola- tions; large, subconical, postrictal tubercles posterolateral to tympanum. Choanae round, large, not concealed by palatal shelf of maxillary arch; vomerine odontophores elevated, median and posterior to choanae, oval in outline, separated on midline by distance equal to I-1/2 odontophore widths, each bearing transverse row of 4-5 teeth; tongue longer than wide, bearing a shallow notch along poste- rior border or not, posterior 2/5 not adherent to floor of mouth; males with vocal slits posterolateral to tongue; vocal sac median, subgular, external. Skin of dorsum finely areolate (most obvious paravertebrally and on upper flanks), venter coarsely areolate; throat areolate in females, no dorsolateral folds or anal sheath; larger warts posterior and posterolateral to vent; discoidal fold well anteriad to groin. Three to four small, subconical ulnar tubercles; palmar tubercle bifid, much larger than oval thenar tubercle; palm areolate, but I-2 more pungent supernumerary tubercles evident at base of each fin- ger; subarticular tubercles pungent, round (or slightly broader than long); KU 179086 has a bifid basal subarticular tubercle on finger III; fingers bear prominent lateral fringes; similar fringe on outer edge of IV continuing along most of palm (Fig. 1); fingers bear broad discs, pads (except on thumb) moderately large, apically rounded; first finger shorter than second; thumbs of males swollen, bearing white, non-spinous nuptial pads. Small (but prominent), subconical tubercles on knee, heel, outer edge of tarsus; less distinct tubercle on inner edge of tarsus; inner metatarsal tubercle twice as long as wide, relatively flat; outer sub- conical, 1/4 size of inner; numerous supernumerary plantar tuber- cles (some more prominent than others, in rows); subarticular tubercles round to slightly longer than wide; toes bear prominent No. 462 BREVIORA ‘wu Z jenba sour] “980621 MM “100J pue puey (GD) ‘C0176 ZOW ‘peoy (qv) sniUasa snjAiopposay inal °| 21n314 TS bh Jar 1979 ELEUTHERODACTYLUS EREMITUS SP. NOV. 5 lateral fringes, no webbing; toe pads smaller than those of fingers, apically rounded, bearing broad discs; heels of flexed hind limbs overlap; shank 47.1—55.0 (x=51.9, N=8) per cent SVL. Most individuals are cream with diffuse brown stippling above and less below; pigmentless areas in groin and on posterior surfaces of thighs (bordered by more dense brown stippling); no labial or limb bars; indefinite canthal-supratympanic stripes, continuing as a diffuse band onto anterior flanks. KU 140878 and 179086 have dark brown dorsolateral stripes continuous with canthal-supratympanic stripe (Fig. 2). KU 179085 is golden brown above with a brown blotch in the center of the back and prominent canthal-supra- tympanic stripes. In life, E. eremitus is green above with a brown to reddish brown head and white below (throat stippled with brown). The groin and posterior surfaces of the thighs are pale yellow. The iris is bright copper with brown flecks. Measurements of holotype inmm. SVL 27.6; shank 13.0; HW 9.7; head length 8.5; upper eyelid width 2.8; IOD 3.0; tympanum length le: sever lenoth 3.1: ESN 23-1. Etymology. Latin, meaning lonely or solitary, in allusion to its relatives which are all cis-Andean. Natural history. KU 140878 was found during the day in a large terrestrial bromeliad beside hwy 30, before the habitat was severely altered by road construction. Specimens from the Quebrada Zapa- dores were collected at night as they sat on broad leaves in the forest beside the streams. Eleutherodactylus eremitus is either quite uncom- mon or several collectors have failed to discover its preferred micro- | A Pe % Dia 5 a Figure 2. Eleutherodactylus eremitus. (A) KU 179085, male, 19.8 SVL; (B) KU 179086, juvenile female, 23.0 mm SVL. 6 BREVIORA No. 462 habitat in the cloud forests (on only one occasion has more than one specimen been found [Miyata secured two on a single night in the type-locality]). Remarks. Frogs of the /acrimosus assembly have been discovered slowly. Eleutherodactylus lacrimosus was named in 1875 but not reported again until 1952 (Lutz and Kloss, 1952). Duellman (1978), Lynch (1979), and Lynch and Duellman (1980) each named species found on the Amazonian slopes of the Andes from Colombia to Bolivia, but the first specimens of these were not obtained until the late 1950’s or early 1960's. In eastern Ecuador, these small frogs are especially common in arboreal bromeliads. In July 1977 at Mera (Prov. Pastaza), at least a dozen examples of E. /acrimosus and E. petersi were found in one bromeliad in remaining patches of forest along the Rio Pastaza. The bromeliad microhabitat is occupied by a variety of frogs on the Pacific versant. At the base of the Andes, E. subsigillatus (Boulenger) normally is found only in arboreal brome- liads. In high cloud forests, E. celator Lynch, E. phoxocephalus Lynch, and E. thymelopsoides Lynch are found almost exclusively in bromeliads. I suspect E. eremitus is actually more abundant on the Pacific versant between 1500 and 2100 m than is suggested by the ten specimens now available. Too infrequently I have failed to exploit the bromeliad microhabitat betwen 800 and 2400 m, in part because frogs were abundant at night and I collected infrequently by day. Inspection of only three bromeliads in July 1977 at Tandapi (Prov. Pichincha) yielded one example each of E. crucifer (Bou- lenger) and FE. parvillus Lynch. Eleutherodactylus bromeliaceus seems to be the closest known relative of E. eremitus, with E. mendax being less closely related. Eleutherodactylus lacrimosus and E. petersi are considered more closely allied to one another than either is to the other three species. ACKNOWLEDGMENTS Specimens were loaned by: William E. Duellman, Museum of Natural History, University of Kansas (KU); W. Ronald Heyer, National Museum of Natural History (USNM); and Ernest E. Willi- ams, Museum of Comparative Zoology (MCZ). Field work in Ecua- dor was supported by funds from the American Philosophical Society (Penrose Fund) and the University of Nebraska Research Council. David Cannatella and Marsha Lynch aided me in field work. 1979 ELEUTHERODACTYLUS EREMITUS SP. NOV. 7 FRE RADRURESCIRED DUELLMAN, W. E. 1978. Three new species of Eleutherodactylus from Amazonian Peru (Amphibia: Anura: Leptodactylidae). Herpetologica 34: 264-270. Lutz, B. & Koss, G. R. 1952. Anfibios anuros do alto Solimoes e Rio Negro apontamentos sobre algumas formas e suas vicariantes. Mem. Inst. Oswaldo Cruz 50: 626-78. Lyncu, J. D. 1979. Leptodactylid frogs of the genus Eleutherodactylus from the Andes of southern Ecuador. Misc. Publ. Mus. Nat. Hist. Univ. Kansas (66): I- 62. Lyncu, J. D. & DUELLMAN, W.E. 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Misc. Publ. Mus. Nat. Hist. Univ. Kansas, in press. Lyncu, J. D. & ScHWarRTz, A. 1972. Taxonomic disposition of some 19th century leptodactylid frog names. J. Herpetol. 5: 103-114. th oe, - a 5 pa ens ay a +e “te ‘ 4 = ne ; ” ake * BRE V POR A MAR 1 8 4985 Museum of Comp A TARVE Ro C0105) US ISSN 0006 UNI VERBITY CAMBRIDGE, MASS. AuGustT 15, 1980 NUMBER 463 JAW MUSCULATURE OF THE WEST INDIAN SNAKE ALSOPHIS CANTHERIGERUS BROOKSI ; (COLUBRIDAE, REPTILIA). KENNETH V. KARDONG! ABSTRACT The West Indian colubrid snake Alsophis cantherigerus brooksi is presently assigned to the subfamily Xenodontinae (sensu Maglio, 1970). It is a member of an assemblage of related snakes from continental stocks that have spread to and diversi- fied within this Caribbean archipelago. The head myology proves to be representa- tive of other xenodontine snakes of the West Indies and thus is a convenient reference for comparison to other subspecies and may be useful in later comparisons to other species’ assemblages of the region. Its jaw musculature conforms basically to that of other generalized colubrid snakes. Special attention is given to the internal fascicular bundles within each muscle. The adductor mandibulae externus superficialis passes posteroventrally from its origin on the skull, inserting mostly by a broad aponeurosis except, for a rostral slip that forms its anterior edge and inserts directly on the mandible. The adductor mandibulae externus profundus consists of anterior and posterior wedges of muscle delineated by a vertical suture between them. Adductor mandibulae externus media- lis is separable into superficial and deep slips. One significant difference in general muscle anatomy from some other colubrids concerns the protractor quadrati. This muscle inserts on the retroarticular process of the mandible which differs from the genus Opheodrys in which it is reported to insert on the quadrate. Lateral jaw musculature proves to be very similar within the subspecies of A/sophis cantherigerus. The only detectable difference occurs in the deep division of the depressor mandibulae. In most subspecies (adspersus, brooksi, cantherigerus, cayma- nus, schwartzi), this deep division is a single slip of the depressor arising from a single site. However, in fuscicauda, pepei, and ruttyi it splits lengthwise so as to arise from two sites of origin. One instance of individual asymmetry was identified in adspersus wherein the insertion of the superficialis differs on left and right sides. The conformity of lateral jaw musculature among subspecies related to brooksi emphasizes the uniform nature of this group. Other authors also found this unifor- mity in features of the integument, skull osteology, and hemipenial characteristics. ' DEPARTMENT OF ZOOLOGY WASHINGTON STATE UNIVERSITY PULLMAN, WASHINGTON 99164 2 BREVIORA No. 463 INTRODUCTION Examination of the West Indian colubrid snakes of the subfamily Xenodontinae by Maglio (1970) served to emphasize the special island biogeographic features of these species. Compared to conti- nental species, West Indian insular members of this subfamily have well delineated distributions and their probable origin, namely from mainland genera, seems better established. Routes of initial coloni- zation by and diversification within the different species assem- blages through this Caribbean archipelago were proposed by Maglio as well. With a view to testing this and related proposals on the phylogeny of this group, I examined the jaw musculature of members from the various assemblages, following any changes through the various species on the different islands. This work and its implications will be the subject of later reports. The present paper is restricted to the species Al/sophis cantherigerus and, in particular, the subspecies A. c. brooksi. This subspecies 1s found only on the Swan Islands, located almost 200 kilometers north of Honduras. It deserves special attention for several reasons. It is part of the assemblage of closely related popu- lations that spread to and diversified over the islands. Its jaw mus- culature is representative of the condition in other West Indian xenodontine species. Its jaw anatomy can thus serve as a central point of reference in later analysis of these island snake species. It is also a rare subspecies even in museum collections. MATERIALS AND METHODS All descriptions of the head anatomy of Alosphis cantherigerus brooksi are based on a single alcohol preserved female specimen from the Museum of Comparative Zoology (MCZ) 11979. The spec- imen, taken on Swan Islands and donated to the collection in 1916, measured 85.5 cm snout-vent length. The tail, missing its severed tip, measured 23.4 cm. The cranial musculature was exposed initially by cutting through the integument along the lips and reflect- ing the skin backwards. This revealed that the specimen had sus- tained a deep cut in the neck musculature that also severed the vertebral column through the fourth cervical vertebra and conse- quently limited the description of structures in the immediate vicin- ity. This study centered upon the jaw musculature and therefore often required the disruption of nerves to first confirm muscle at- 1979 Jaw musculature of Alsophis cantherigerus brooksi 3 tachments with certainty. As a result, some of the peripheral distri- butions of the nerves were lost before they could be traced with confidence. The blood vessels proved impractical to follow for the same reasons. Consequently, only those pathways and distributions of nerves and blood vessels that could later be confirmed are included in the descriptions. No bilateral asymmetry in jaw muscu- lature was detected. Muscle and ligament terminology follows Haas (1973), Kardong (1973, 1974), and Pregill (1977); interpretation of cranial nerves relies upon Auen and Langebartel (1977); and cephalic gland nomenclature is based on Taub (1966). One depar- ture in nomenclature concerns the muscle running between the neck and dorsal end of the quadrate, which is often referred to as M. retractor quadrati especially in descriptions of venomous snakes (e.g. Dullemeijer, 1956). Others employ this same term for a quite different muscle (e.g. Kochva, 1962). To avoid confusion herein, the term M. cervicoquadratus is used instead. The other subspecies of Alsophis cantherigerus examined include: adspersus (MCZ 68727), cantherigerus (MCZ 13288), caymanus (MCZ 44886), fuscicauda (MCZ 44875), pepei (MCZ 13289), ruttyi (MCZ 44877), and schwartzi (MCZ 56430). GENERAL DESCRIPTION Cephalic Glands The nasal glands, like other lateral cephalic glands, are bilaterally paired. Each rests in an anterior depression on the anteromedial face of the prefrontal and is bounded medially by a dorsal process of the respective septomaxilla and by the nasal capsule. The Harderian gland (Figs. 2, 3a, 4b) is found ventral to and extends behind the eye. Beneath the eye the gland lies within the orbit and is com- pressed between eyeball and the ligamentum orbitale interioris (sep- tum interorbitale of Dullemeier, 1956) which forms the ventral floor of the orbit. Passing posteriorly out of the orbit it enlarges, occupying the space lateral to the levator pterygoidei muscle and medial to the postorbital bone and the adductor mandibulae exter- nus superficialis muscle. The infralabial glands (Fig. 4c) extend pos- teriorly along the lateral aspect of the dentary and compound bones to approximately the level of the anterior insertion of the adductor mandibulae externus superficialis muscle. The supralabial glands 4 BREVIORA No. 463 extend posteriorly slightly farther. However, they lie along the upper lips and pass lateral to the superficialis muscles and maxillae. Pressed into the side of each supralabial gland and residing behind the eye is the large Duvernoy’s gland (Fig. 4c), which in the pre- served state appears lighter in color and more distinctly lobed. The small, unpaired premaxillary gland lies beneath the upper lip on the superficial face of the premaxillary bone. In the chin, the sublingual gland (Fig. 6a, b) is represented by three parts, a single medial and paired lateral glands. The medial sublingual gland is stationed along the ventroanterior end of the tongue and anchored to its connective tissue sheath. The lateral sublingual glands insert into the oral epithelium and serve as attach- ment sites for the pars glandularis of the transversus branchialis and the protractor larygeus muscles. Ligaments A number of small, unnamed ligaments link the cranial elements together, but only the prominent or most commonly cited in the literature are discussed here. Two primary ligaments serve to check the motion of the supratemporal relative to the braincase. One is the parieto-supratemporal ligament that runs from the dorsal edge of the supratemporal to the dorsal surface of the exoccipital. It also serves as the site of origin for parts of the adductor mandibulae externus medialis and depressor mandibulae muscles. The other is the prooto-supratemporal ligament that attaches to the ventral edge of the supratemporal and to the adjacent region of the braincase. Rotation of the prefrontal is controlled in part by the fronto- prefrontal ligament (Fig. 2) from the ventromedial corner of the prefrontal to the adjacent parasphenoid. Several prominent ligaments are associated with the palato- maxillary arch (pterygoid, ectopterygoid, maxilla, and palatine). Within the arch, the intramaxillary ligament (Fig. 2) reaches from the medial ectopterygoid process of the maxilla forward to attach jointly on the palatine process of the maxilla and maxillary process of the palatine. The short, fan-shaped quadrato-pterygoid ligament runs from the narrow posterior end of the pterygoid to a wider medial attachment site along the mandible that includes the retroar- ticular process and extends below the articular notch. The strong, cord-like maxillo-postorbital ligament (Fig. 4) reaches from the 1979 Jaw musculature of Alsophis cantherigerus brooksi 5 downward directed tip of the postorbital to the lateral side of the maxilla just anterior to its articulation with the ectopterygoid. The quadrato-maxillary ligament (Fig. 4) begins along the laterodorsal edge of the retroarticular process, passes forward above the lateral condyle of the quadrate, and divides. The smaller division attaches to the nearby dermis beneath the most posterior supralabial scale, whereas the larger division continues forward to attach to the pos- terolateral region of the maxilla. Cranial Nerves Figs, 2=3a-d> 4a-¢ Trigeminal Nerve (V). The trigeminal is one of three major nerves supplying the jaw musculature. Trigeminal branches that supply the jaw musculature exit from the cranium through two openings in the prootic bone, the maxillary and mandibular foramina. Five branches could be found passing through the maxillary foramen. The largest, the maxillary division (V.), passes lateral to the pseudo- temporalis then bends forward continuing in an anterior path dorsal to the origin of the pterygoideus. A second nerve divides lateral to the pseudotemporalis with one branch passing laterally to glandular tissue along the upper lip and the second swinging dorsally, superfi- cial to the facial vein, to enter the ventral surface of the adductor mandibulae externus superficialis. Two other nerves also course dorsally after exiting from the foramen. Both pass deep to the facial vein, but one enters the medial face of the adductor mandibulae externus medialis (deep division) and the other enters the medial face of the adductor mandibulae externus superficialis. The fifth nerve leaving this foramen passes laterally. Through the mandibular foramen exit six branches of the trigemi- nal nerve. The largest branch is the mandibular division (V3) that passes posteriorly and then turns ventrally following a route that carries it between the adductor mandibulae externus profundus and the adductor mandibulae posterior. Eventually it enters a foramen in the mandible located at the anterior end of the mandibular fossa. A second branch of the trigeminal courses posteriorly, then swings dorsally, bifurcates, and enters the medial face of the adductor man- dibulae externus profundus and medioposterior face of the adduc- tor mandibulae externus medialis (superficial division). A third nerve passes posteriorly deep to the mandibular division. It sends 6 BREVIORA No. 463 one branch between medial and lateral divisions of the adductor mandibulae posterior, another between the pars minimus and the lateral division of adductor mandibulae posterior, and several branches into the medial surface of the adductor mandibulae exter- nus profundus. The remaining three nerves of the trigeminal that exit through the mandibular foramen pass ventrally—one disap- pears between the pterygoideus and the levator pterygoidei, one enters the lateral face of the pseudotemporalis, and the last runs laterally into the glandular tissue along the upper lip. Facial Nerve (V11). Two divisions of the facial nerve exit through the mandibular foramen. The larger is the hyomandibular nerve that passes posteriorly over the columella, under the quadrate, and enters the depressor mandibulae. A slender communication arising from the base of the hyomandibular travels posteriorly and merges with craniocervical trunk. The second division of the facial nerve is the palatine nerve. It courses ventrally along the underside of the braincase, is joined by the cranial sympathetic nerve, and then enters the posterior Vidian canal in the ventral aspect of the basi- sphenoid bone. Craniocervical Trunk. The glossopharyngeal (IX), vagus (X), and hypoglossal (XII) cranial nerves combine to form a single large nerve or plexus, the craniocervical trunk (Auen and Lagebartel, 1977). In addition, a slender communication runs between the base of the hyomandibular and this trunk. Finally, a spinal nerve (pre- sumably the ventral ramus of the first spinal nerve) emerges from between the rectus capitus anterior pars dorsalis and longissimus dorsi (ventral head) cervical muscles to join with the craniocervical trunk. The trunk sweeps around the side of the body at about the angle of the jaws and supplies various members of the hypobran- chial musculature. Lateral Jaw Musculature M. cervicomandibularis (cm) Figs. 1, 4c The epimysium of the spinalis-semispinalis complex near the dor- sal midline serves as the surface of origin. This origin extends along a line even with neural spines 3 to 8 in the nape of the neck and 1979 Jaw musculature of Alsophis cantherigerus brooksi 3 tachments with certainty. As a result, some of the peripheral distri- butions of the nerves were lost before they could be traced with confidence. The blood vessels proved impractical to follow for the same reasons. Consequently, only those pathways and distributions of nerves and blood vessels that could later be confirmed are included in the descriptions. No bilateral asymmetry in jaw muscu- lature was detected. Muscle and ligament terminology follows Haas (1973), Kardong (1973, 1974), and Pregill (1977); interpretation of cranial nerves relies upon Auen and Langebartel (1977); and cephalic gland nomenclature is based on Taub (1966). One depar- ture in nomenclature concerns the muscle running between the neck and dorsal end of the quadrate, which is often referred to as M. retractor quadrati especially in descriptions of venomous snakes (e.g. Dullemeijer, 1956). Others employ this same term for a quite different muscle (e.g. Kochva, 1962). To avoid confusion herein, the term M. cervicoquadratus is used instead. The other subspecies of Alsophis cantherigerus examined include: adspersus (MCZ 68727), cantherigerus (MCZ 13288), caymanus (MCZ 44886), fuscicauda (MCZ 44875), pepei (MCZ 13289), ruttyi (MCZ 44877), and schwartzi (MCZ 56430). GENERAL DESCRIPTION Cephalic Glands The nasal glands, like other lateral cephalic glands, are bilaterally paired. Each rests in an anterior depression on the anteromedial face of the prefrontal and is bounded medially by a dorsal process of the respective septomaxilla and by the nasal capsule. The Harderian gland (Figs. 2, 3a, 4b) is found ventral to and extends behind the eye. Beneath the eye the gland lies within the orbit and is com- pressed between eyeball and the ligamentum orbitale interioris (sep- tum interorbitale of Dullemeier, 1956) which forms the ventral floor of the orbit. Passing posteriorly out of the orbit it enlarges, occupying the space lateral to the levator pterygoidei muscle and medial to the postorbital bone and the adductor mandibulae exter- nus superficialis muscle. The infralabial glands (Fig. 4c) extend pos- teriorly along the lateral aspect of the dentary and compound bones to approximately the level of the anterior insertion of the adductor mandibulae externus superficialis muscle. The supralabial glands 4 BREVIORA No. 463 extend posteriorly slightly farther. However, they lie along the upper lips and pass lateral to the superficialis muscles and maxillae. Pressed into the side of each supralabial gland and residing behind the eye is the large Duvernoy’s gland (Fig. 4c), which in the pre- served state appears lighter in color and more distinctly lobed. The small, unpaired premaxillary gland lies beneath the upper lip on the superficial face of the premaxillary bone. In the chin, the sublingual gland (Fig. 6a, b) is represented by three parts, a single medial and paired lateral glands. The medial sublingual gland is stationed along the ventroanterior end of the tongue and anchored to its connective tissue sheath. The lateral sublingual glands insert into the oral epithelium and serve as attach- ment sites for the pars glandularis of the transversus branchialis and the protractor larygeus muscles. Ligaments A number of small, unnamed ligaments link the cranial elements together, but only the prominent or most commonly cited in the literature are discussed here. Two primary ligaments serve to check the motion of the supratemporal relative to the braincase. One is the parieto-supratemporal ligament that runs from the dorsal edge of the supratemporal to the dorsal surface of the exoccipital. It also serves as the site of origin for parts of the adductor mandibulae externus medialis and depressor mandibulae muscles. The other is the prooto-supratemporal ligament that attaches to the ventral edge of the supratemporal and to the adjacent region of the braincase. Rotation of the prefrontal is controlled in part by the fronto- prefrontal ligament (Fig. 2) from the ventromedial corner of the prefrontal to the adjacent parasphenoid. Several prominent ligaments are associated with the palato- maxillary arch (pterygoid, ectopterygoid, maxilla, and palatine). Within the arch, the intramaxillary ligament (Fig. 2) reaches from the medial ectopterygoid process of the maxilla forward to attach jointly on the palatine process of the maxilla and maxillary process of the palatine. The short, fan-shaped quadrato-pterygoid ligament runs from the narrow posterior end of the pterygoid to a wider medial attachment site along the mandible that includes the retroar- ticular process and extends below the articular notch. The strong, cord-like maxillo-postorbital ligament (Fig. 4) reaches from the 1979 Jaw musculature of Alsophis cantherigerus brooksi 5 downward directed tip of the postorbital to the lateral side of the maxilla just anterior to its articulation with the ectopterygoid. The quadrato-maxillary ligament (Fig. 4) begins along the laterodorsal edge of the retroarticular process, passes forward above the lateral condyle of the quadrate, and divides. The smaller division attaches to the nearby dermis beneath the most posterior supralabial scale, whereas the larger division continues forward to attach to the pos- terolateral region of the maxilla. Cranial Nerves Figs. 2, 3a-d, 4a-c Trigeminal Nerve (V). The trigeminal is one of three major nerves supplying the jaw musculature. Trigeminal branches that supply the jaw musculature exit from the cranium through two openings in the prootic bone, the maxillary and mandibular foramina. Five branches could be found passing through the maxillary foramen. The largest, the maxillary division (V5), passes lateral to the pseudo- temporalis then bends forward continuing in an anterior path dorsal to the origin of the pterygoideus. A second nerve divides lateral to the pseudotemporalis with one branch passing laterally to glandular tissue along the upper lip and the second swinging dorsally, superfi- cial to the facial vein, to enter the ventral surface of the adductor mandibulae externus superficialis. Two other nerves also course dorsally after exiting from the foramen. Both pass deep to the facial vein, but one enters the medial face of the adductor mandibulae externus medialis (deep division) and the other enters the medial face of the adductor mandibulae externus superficialis. The fifth nerve leaving this foramen passes laterally. Through the mandibular foramen exit six branches of the trigemi- nal nerve. The largest branch is the mandibular division (V;) that passes posteriorly and then turns ventrally following a route that carries it between the adductor mandibulae externus profundus and the adductor mandibulae posterior. Eventually it enters a foramen in the mandible located at the anterior end of the mandibular fossa. A second branch of the trigeminal courses posteriorly, then swings dorsally, bifurcates, and enters the medial face of the adductor man- dibulae externus profundus and medioposterior face of the adduc- tor mandibulae externus medialis (superficial division). A third nerve passes posteriorly deep to the mandibular division. It sends 6 BREVIORA No. 463 one branch between medial and lateral divisions of the adductor mandibulae posterior, another between the pars minimus and the lateral division of adductor mandibulae posterior, and several branches into the medial surface of the adductor mandibulae exter- nus profundus. The remaining three nerves of the trigeminal that exit through the mandibular foramen pass ventrally—one disap- pears between the pterygoideus and the levator pterygoidei, one enters the lateral face of the pseudotemporalis, and the last runs laterally into the glandular tissue along the upper lip. Facial Nerve (VII). Two divisions of the facial nerve exit through the mandibular foramen. The larger is the hyomandibular nerve that passes posteriorly over the columella, under the quadrate, and enters the depressor mandibulae. A slender communication arising from the base of the hyomandibular travels posteriorly and merges with craniocervical trunk. The second division of the facial nerve is the palatine nerve. It courses ventrally along the underside of the braincase, is joined by the cranial sympathetic nerve, and then enters the posterior Vidian canal in the ventral aspect of the basi- sphenoid bone. Craniocervical Trunk. The glossopharyngeal (IX), vagus (X), and hypoglossal (XII) cranial nerves combine to form a single large nerve or plexus, the craniocervical trunk (Auen and Lagebartel, 1977). In addition, a slender communication runs between the base of the hyomandibular and this trunk. Finally, a spinal nerve (pre- sumably the ventral ramus of the first spinal nerve) emerges from between the rectus capitus anterior pars dorsalis and longissimus dorsi (ventral head) cervical muscles to join with the craniocervical trunk. The trunk sweeps around the side of the body at about the angle of the jaws and supplies various members of the hypobran- chial musculature. Lateral Jaw Musculature M. cervicomandibularis (cm) Figs. 1, 4c The epimysium of the spinalis-semispinalis complex near the dor- sal midline serves as the surface of origin. This origin extends along a line even with neural spines 3 to 8 in the nape of the neck and 1979 Jaw musculature of Alsophis cantherigerus brooksi 7 partially overlaps the anterior origin of the neuromandibularis. This broad muscle travels ventrally, passing over the deep cervico- quadratus, and narrows abruptly to insert on the lateral epicondyle of the quadrate and on the adjacent part of quadrato-maxillary (dermal) ligament, which passes posteriorly to its own attachment on the dorsolateral retroarticular process. M. depressor mandibulae (dm) (M. occipito-quadrato-mandibularis) Figs. 1, 4 a-c There are two parts of this muscle, superficial and deep, divided by the cervicoquadratus that passes between these two parts en route to its own insertion. Branches of cranial nerve VII also pass through the belly of the deep division, then along the medial side of the superficial division. The superficial division has an origin that runs from the parieto- supratemporal ligament next to the adductor mandibulae externus medialis, across the supratemporal to the anterodorsal corner of the quadrate, and extends down along the surface epimysium of adduc- tor mandibulae externus profundus near its own origin. It inserts directly on the lateral rim of the mandibular retroarticular process, overlapping the attachment site of the quadrato-maxillary ligament. The deep division arises from the posterior tip of the supratem- poral and the posterior, dorsomedial edge of the quadrate. It is a parallel muscle that inserts on the dorsomedial surface of the retro- articular process. M. cervicoquadratus (cq) Figs. 1, 4 a-c This is a long, tapering muscle that takes origin in the dermis of the lateral integument posterior to the belly of the neuromandibula- ris over the intercostal muscles. It gradually narrows as it courses forward, slipping under the more superficial cervicomandibularis, becoming a cord-like tendon that passes between the superficial and the deep divisions of the depressor mandibulae to insert laterally on the proximal end of the quadrate just below its posterior corner. 8 BREVIORA No. 463 Figure |. Lateral view of the skull and ventral view of the quadrate and lower jaw showing sites of muscle attachments. Figure 2. Ventral view of head. The bones of the left palato-maxillary arch have been removed to expose deep structures. On the right, the M. pterygoideus (pg) has been cut and partially reflected and the mandible rotated outward to better expose some of the underlying structures. Abbreviations: am M. adductor mandibulae externus medialis ap M. adductor mandibulae externus profundus apo M. adductor mandibulae posterior as M. adductor mandibulae externus superficialis br transverse ridge on basisphenoid Cc columella cm M. cervicomandibularis cp choanal process of palatine cq M. cervicoquadratus ct craniocervical nerve trunk (IX, X, XII) dm M. depressor mandibulae gg M. genioglossus gt M. geniotrachealis Hd Harderian gland ima M. intermandibularis anterior ipa M. intermandibularis posterior, pars anterior ipp M. intermandibular posterior, pars posterior Ifp ligamentum fronto-prefrontale lim ligamentum intramaxillare Imp ligmentum maxillo-postorbitale loi ligamentum orbitale inferioris Ip M. levator pterygoidei Iqm ligamentum quadrato-maxillare nm M. neuromandibularis pg M. pterygoideus pga M. pterygoideus accessorius pp M. protractor pterygoidei pq M. protractor quadrati pst M. pseudotemporalis red M. rectus capitus anterior, pars dorsalis cv M. rectus capitus anterior, pars ventralis rp M. retractor pterygoidei IV M. retractor vomeris Vo Trigeminal nerve, maxillary division V; Trigeminal nerve, mandibular division 1979 Jaw musculature of Alsophis cantherigerus brooksi >> : SS, 2 ead 10 BREVIORA No. 463 M. adductor mandibulae externus superficialis (as) Figs lcd 40a A temporal crest, prominent posteriorly, becomes lower as it passes forward along the dorsolateral side of the skull. Anteriorly it curves outward to terminate on the postorbital process of the parie- tal. The superficialis originates directly from the anterior half of this temporal crest and from the base of the postorbital bone. It passes posteriorly and downward, partially overlying the adductor mandi- Figure 3. Lateral view of head showing progressively more structures (A-D) in place. Abbreviations: am M. adductor mandibulae externus medialis, deep (d) and superficial (s) divisions ap M. adductor mandibulae externus profundus apo M. adductor mandibulae posterior, lateral division (1), medial division (m), pars minimus (mi) as M. adductor mandibulae externus superficialis c columella ct craniocervical nerve trunk (IX, X, XII) ec ectopterygoid f frontal Hd Harderian gland if infralabial nerve foramen (posterior) ile M. iliocostalis Id M. longissimus dorsi Ip M. levator pterygoidei mf mental foramen mx maxilla n nasal pf prefrontal pg M. pterygoideus pga M. pterygoideus accessorius po postorbital pp M. protractor pterygoidei pq M. protractor quadrati pst M. pseudotemporalis q quadrate cv M. rectus capitus anterior, pars ventralis ssp M. spinalis-semispinalis st supratemporal tc temporal crest Vo Trigeminal nerve, maxillary division V3 Trigeminal nerve, mandibular division 1979 Jaw musculature of Alsophis cantherigerus brooksi 11 is . igi eS pif Ha pr. | pals ‘KEZ} 1 fF AK 12 BREVIORA No. 463 1979 Jaw musculature of Alsophis cantherigerus brooksi 13 bulae externus medialis. Its anterior fibers arch around the corner of the mouth without forming any attachments to the buccal mem- brane and pass to the mandible, where they directly insert on the compound bone just posterior to the end of the dentary tooth row and lateral to the anterior insertion of the adductor mandibulae externus profundus. The remaining fibers of this flat muscle termi- nate superficial to the profundus in a broad aponeurosis that spreads across the surface epimysium of the profundus and can be followed to an insertion extending in a narrow line from the lateral epicondyle of the quadrate along the mandible, ventral to the inser- tion of thé profundus, and forward to a point not quite reaching the directly inserting anterior fibers described above. M. adductor mandibulae externus profundus (ap) Figs dis ced: 4sa-¢ This is a large, triangular block of muscle divisible into two mus- cle wedges that separate cleanly along a vertical cleft between them. The anterior wedge takes origin from the lateral, anterodorsal corner of the quadrate. The origin of the posterior wedge also begins on this corner of the quadrate beneath the attachment of the ante- rior wedge. However, the origin of the posterior wedge extends Figure 4. Lateral view of skull showing progressively (A~C) more structures in place. Abbreviations: am M. adductor mandibulae externus medialis ap M. adductor mandibulae externus profundus as M. adductor mandibulae externus superficialis cm M. cervicomandibularis cq M. cervicoquadratus dm M. depressor mandibulae, deep division (d), superficial division (s) Dv Duvernoy’s gland Hg Harderian gland ig infralabial gland ipp M. intermandibularis posterior, pars posterior Imp ligmentum maxillo-postorbitale Ip M. levator pterygoidei lqm ligamentum quadrato-maxillare nm M. neuromandibularis pg M. pterygoideus sg supralabial gland 14 BREVIORA No. 463 distally along the anterolateral edge of the quadrate reaching the lateral epicondyle. Both parts form a large, thick muscle that inserts directly on the lateral side of the compound bone in a low depres- sion from below the quadrato-mandibular articulation forward to the posterior infralabial nerve foramen. M. adductor mandibulae externus medialis (am) Bigs, 1h 3 ¢-d.4-a—c This muscle takes origin from the parieto-supratemporal ligament and the posterior half of the temporal crest, filling a depression and area adjacent to the crest. As the muscle passes ventrally its fibers gather into two divisions—superficial and deep. Dorsally a few fib- ers intermingle between both but as they pass downward they become more discrete. The deep division takes origin along the posterior temporal crest only. This division descends as a wide, thin sheet that inserts directly to the compound bone medial to the adductor mandibulae posterior. This insertion then extends forward and upward to the dorsal edge of the compound bone reaching a point posterior to the end of the dentary tooth row. The superficial division contains most of the fibers of this muscle and originates from both the posterior temporal crest and the parieto-supra- temporal ligament. It narrows and gives rise to a short tendon that inserts on the dorsal, posterior end of the dentary. M. adductor mandibulae posterior (apo) Figsei. aucad This muscle, composed of parallel fibers, is triangular in shape and concealed beneath the adductor mandibulae externus profun- dus. It passes to both sides of the prearticular crest and so is divided naturally into two major parts—lateral and medial. In addition, a distinctive broad, flat muscle sheet forms the most lateral division, here referred to as the pars minimus. The pars minimus takes origin at the base of the proximal end of the quadrate and then extends along its entire anterior, lateral edge to a point above the lateral condyle. It inserts along the dorsal edge of the surangular crest forward to a point just posterior to the adjacent mandibular nerve. The lateral division of the muscle takes origin along the anteromedial length of the quadrate. Its parallel fibers pass ventrally and forward to insert directly on the floor of the mandibular fossa and lateral side of the prearticular crest. 1979 Jaw musculature of Alsophis cantherigerus brooksi 15 The medial division of this muscle takes origin along the medial side of the quadrate but more posterior in position than the lateral division. A few central fibers arise from a small, short, common tendon, but most are parallel and all collectively form a sheet that passes in an anteroventral direction eventually inserting directly on the dorsal edge of the prearticular crest and along a low depression on its medial face. The anterior part of this insertion passes lateral to the posterior insertion of the deep division of adductor mandibu- lae externus medialis. M. pseudotemporalis (pst) Figs, 1, 3° b-d This muscle arises beneath the anterior part of the adductor man- dibulae externus medialis and the posterior part of the adductor mandibulae externus superficialis along the mid-temporal crest. It is a ribbon-like muscle of parallel fibers that courses ventrally, passing deep to the maxillary division of the trigeminal (V) nerve, the facial carotid artery, and jugular vein. It attaches directly to the com- pound bone medial and slightly anterior to the front of the deep division of the adductor mandibulae externus medialis. M. pterygoideus (pg) igss iby 2. 23nc-d. 4 ab. SeaaD The pterygoideus takes origin from the lateral maxillary process of the ectopterygoid, primarily by a prominent tendon. The tendon remains superficial and passes ventrally reaching the middle of the muscle. Most muscle fibers arise along the length of this prominent tendon although a few directly arise from a small area on the ecto- pterygoid immediately medial to the origin of this tendon. The fib- ers sweep backward forming, on approach to the insertion, the swollen belly of the muscle that projects below the compound bone. The fibers curve upward to insert on the ventral surface of the mandibular retroarticular process. Specifically, this insertion includes the ventroposterior surface of the retroarticular process and extends a short distance forward along the ventrolateral edge of the process to a point even with the articular notch. There are two further noteworthy anatomical features of this muscle. First, along its anterolateral surface the epimysium is drawn up into a loose fascial connection that joins it with the anterior edge 16 BREVIORA No. 463 of the nearby adductor mandibulae externus medialis. Second, fib- ers composing the anterodorsal part of the muscle form a separate fascicle. Though anatomically an integral part of the pterygoideus, these fibers insert, via a wide aponeurosis, on the ventrolateral edge of the retroarticular process. Most fibers of this slip arise directly from the prominent ventral tendon near its anterior end, but a few arise directly from the lateral maxillary process of the ectopterygoid medial to this tendon. M. pterygoideus accessorius (pga) Figs, 15, 25 3b This muscle arises directly from the ventral surface of the ptery- goid and base of the ectopterygoid. Specifically, this origin includes the ventral surface of the lateral projection of the pterygoid, its lateroventral groove, and lateroposterior base of the ectopterygoid. The muscle passes backward to insert on the inner side of the retroarticular process along its medial curvature, parallel with but medial to the insertion of the pterygoideus. M. protractor quadrati (pq) Figs. 2\\3a—b This muscle arises from a tendon, shared with its contralateral partner, that originates from the midventral basioccipital. It is a flat muscle, passing horizontally in a posterolateral direction over the dorsal surface of the protractor pterygoidei. A few fibers insert on the medial epicondyle of the quadrate, but most insert directly to the dorsomedial side of the mandibular retroarticular process, dorsal to the origin of the pterygoideus accessorius. M. protractor pterygoidei (pp) Rigs: e223 This muscle takes origin from the basisphenoid, specifically from a mid-transverse ridge, and forward parasagittally along the basi- sphenoid to a point just past the anterior Vidian foramen. It passes posteriorly, becoming fusiform in shape, and inserts across the pos- terodorsal end of the pterygoid with an especially firm attachment to the caudal tip of the bone. 1979 Jaw musculature of Alsophis cantherigerus brooksi iy M. levator pterygoidei (Ip) Bisse ly 2, 3.a=d; 4a A low depression in the posterior, ventral face of the postorbital process of the parietal serves as the surface from which this muscle takes origin. It passes ventrally, widening along an anteroposterior axis to form a long insertion on the pterygoid. This insertion begins posteriorly on the dorsolateral side of the pterygoid, runs forward to the ectopterygoid-pterygoid articulation, across the base of the ectopterygoid, and ends on the adjacent outer side of the pterygoid short of the pterygo-palatine articulation. The insertion thus lies along the shaft of this bone occupying the dorsolateral fossa. M. retractor pterygoidei (rp) Fig: 2 This muscle arises directly from the anterior slope of the trans- verse ridge on the basisphenoid, adjacent to the origin of the pro- tractor pterygoidei. It passes forward to insert directly on the anterodorsal surface of the pterygoid and posterodorsal surface of the palatine, and by a broad aponeurosis to the posterior edge of the choanal process of the palatine. M. retractor vomeris (rv) Fig. 2 This muscle arises from the sphenoid, lateral to the anterior end of the origin of the protractor pterygoidei. It passes forward as a spindle-like muscle bounded laterally by the retractor pterygoidei. Anteriorly, its fibers converge into a slender cord-like tendon that accounts for half the total muscle length. This tendon inserts on posterodorsal corner of the vomer. Throat Musculature M. neuromandibularis (nm) Figs. 1, 4c, 5 a-b, 6b Beginning in the neck, this broad, flat muscle sweeps downward and forward around the side of the body, passing under the cervico- quadratus and over the axial musculature, to insert at several sites BREVIORA BANS A ANS / A Niaa' My ba) No. 463 1979 Jaw musculature of Alsophis cantherigerus brooksi 19 along the mandible via aponeuroses. It has been variously treated by other authors, some dividing it into several parts (Langebartel, 1968; Kardong, 1973; Haas, 1973) while others include it as part of a composite muscle (Albright and Nelson, 1959; Cundall, 1974). For descriptive purposes, it is here treated as a single unit. It originates from the epimysium of the spinalis-semispinalis muscles adjacent to neural spines 6-11. The anterior part of this origin lies under the posterior part of the cervicomandibularis. As it passes into the throat, it is crossed by two inscriptions. A separate muscle, the costomandibularis arises by three separate slips from the ventrolat- eral tips and Costal cartilages of ribs 2-4. From here the costomandi- bularis passes forward to join and most fibers terminate on the medial face of neuromandibularis along the posterior inscription. Farther anteriorly, the neuromandibularis is joined by a few addi- tional fibers originating from the lingual process and anterior end of the ceratobranchial of the hyoid; these fibers contribute to the medial border of the muscle. Insertion on the mandible is by a broad aponeurosis, but along three specific sites. One site of insertion, via an aponeurosis, is along the ventral edge of the angular and posteroventral edge of the sple- nial. A second insertion, also via an aponeurosis, begins along the posterolateral edge of the dentary where it articulates with the com- Figure 5. Ventral view of throat musculature. At bottom (B), a section of each intermandibularis posterior, pars posterior (ipp) has been removed. The left neuro- mandibularis (nm) has been cut and reflected to expose the underlying structures, including a prominent blood vessel (bv). Abbreviations: cc M. constrictor colli cos M. costomandibularis ht M. hyotrachealis hy hyoid laa M. intermandibularis anterior, pars anterior lag M. intermandibularis anterior, pars glandularis lap M. intermandibularis anterior, pars posterior ig infralabial gland is M. interscutali ipa M. intermandibularis posterior, pars anterior ipp M. intermandibularis posterior, pars posterior nm M. neuromandibularis tb M. transversus branchialis th M. transversus hyoideus 20 BREVIORA No. 463 pound bone. This insertion extends forward along the lateral face of the dentary, passing below the anterior mental foramen and termi- nates behind the intermandibularis anterior. Thirdly, the epimysium along the superior face of the neuromandibularis forms an aponeu- rosis that passes dorsal to the origin of intermandibularis posterior, pars anterior to insert adjacent to it on the mandible. This forms a sleeve through which intermandibularis posterior, pars anterior passes from its origin forward toward its insertion (5b, 6b). M. intermandibularis posterior (ipa, 1pp) Figs. 1, 4c, 5 a-b, 6b The intermandibularis posterior, pars posterior takes origin beneath the insertions of the adductores mandibulae profundus and superficialis along the ventral surface of the compound bone. It is a strap-like muscle that passes forward superficial to the neuroman- dibularis to join its contralateral partner in inserting in the dermis at the central midline, just posterior to the origin of transversus branchialis. The origin of intermandibularis posterior, pars anterior is also along the ventral edge of the mandible, but anterior to the insertion of the pars posterior. Forming into a strap-like muscle, it passes through a sleeve formed by the insertion of the neuromandibularis and then runs medial to the mandibular ramus. It inserts in the posteroventral epimysium of the transversus branchialis. M. intermandibularis anterior (iaa, lap, 1ag) Figs, 1, 5 a—b) 6a-b The muscle originates from the anteroventral tip of the dentary as the bone narrows and bends inward. This is lateral to the origins of the genioglossus and geniotrachealis. Though not anatomically separate, two regions of this muscle can be recognized—an anterior region (pars anterior), whose fibers pass inward to insert on the interramal pad, and a posterior region (pars posterior), that also passes inward to insert in dermis with its contralateral partner ona midventral raphe. Anterior fibers of the intermandibularis anterior, pars glandularis (=constrictor of Langebartel, 1968) are fan-shaped and attach to the lateral edge of the interramal pad. The narrow posterior part of the muscle wraps around the ventral aspect of the lateral sublingual gland and attaches along the posteroventral surface of this gland. 1979 Jaw musculature of Alsophis cantherigerus brooksi 21 M. transversus branchialis (tb) igse: 5-a—b; 6..a—D This muscle originates from two heads, glandular and mucosal (pars glandularis and pars mucosalis respectively, of Albright and Nelson, 1959). The glandular head arises on the posterior end of the lateral sublingual gland. It passes as a ribbon-like band, arching posteriorly around the more medial genioglossus and geniotrachea- lis, then ventromedially to form the anterior part of the muscle. It is joined by fibers of the mucosal head arising from the dermis of the buccal epithelium midway between mandible and trachea to form the middle and posterior parts of the muscle. It inserts in the dermis of the chin integument at the ventral midline, between insertions of intermandibulares anterior and posterior. M. genioglossus (gg) Figs. 1, 6 a-b Posteriorly this paired muscle is entwined around the tongue. Anteriorly, it originates from two heads. The medial head arises from the posterior tip of the interramal pad, then passes caudally around the lateral face of the medial sublingual gland. The lateral head receives a few fibers from the dorsal surface of the intermandibularis anterior, but most arise from the anterior tip of the dentary along its inward inflection ventral to the geniotra- chealis and dorsal to the intermandibularis anterior. It slants inward as a narrow cord-like muscle to join with the medial head near the middle of the medial sublingual gland. Just posterior to this gland, the muscle passes to the tongue sheath along which it runs caudally. About half way along the tongue, right and left genioglossus mus- cles change sides. The right muscle passes across the ventral surface to the left side and, at the same point, the left muscle passes across the dorsal surface to the right side. They stay in these reversed positions until reaching their posterior insertions on the hyoglossus, at the point of termination of the tongue sheath. M. geniotrachealis (gt) Figs’ 1, 5b; 674-0 This muscle inserts along the side of the trachea beginning just posterior to the intrinsic laryngeal musculature (Kardong, 1972) spanning 9 cartilaginous rings. It passes forward as a ribbon-like No. 463 BREVIORA 22 1979 Jaw musculature of Alsophis cantherigerus brooksi 23 muscle to its origin along the medial curve of the anterior dentary dorsal to and expanding across a wider area of origin than the genioglossus. M. hyotrachealis (ht) Figs. 5b, 6 a-b This slender, ribbon-like muscle inserts along the side of the tra- chea dorsal to the anterior origin of geniotrachealis. This insertion begins immediately posterior to the intrinsic largyngeal muscles (M. dilatator laryngis) and spans three cartilaginous rings. The muscle Figure 6. Ventral view of deep throat musculature. At bottom (B), the left neuromandibularis (nm) and right transversus branchialis (tbm) are reflected. The middorsal surface of the neuromandibularis develops a short attachment to the nearby mandible that forms a sheath-like (sh) channel for the origin of the intermandibularis posterior, pars anterior (ipa). The label for the hyoid (hy) here lies on the lingual process. The intermandibularis anterior (iaa and iap) has been sectioned along its midventral insertion and pulled slightly to one side. At top (A), most of the muscles and infralabial gland have been removed to reveal the course of the transversus branchialis (tbm and tbg), intermandibularis anterior, pars glandu- laris (iag), and geniotrachialis (gt). The anterior part of the tongue (tg) has been removed to expose the trachea (tr) and a few muscle attachments to it. Abbreviations: gg M. genioglossus gt M. geniotrachealis gt M. hyotrachealis hy hyoid laa M. intermandibularis anterior, pars anterior lag M. intermandibularis anterior, pars glandularis lap M. intermandibularis anterior, pars posterior ig infralabial gland im intrinsic musculature of larynx ipa M. intermandibularis posterior, pars anterior ir inter-ramal pad nm M. neuromandibularis pl M. protractor laryngeus sh sleeve formed by neuromandibularis about origin of intermandibularis posterior, pars anterior sll sublingual gland, lateral slm sublingual gland, medial tbg M. transversus branchialis, glandular head tbm M. transversus branchialis, mucosal head tg tongue tr trachea 24 BREVIORA No. 463 passes posteriorly along the floor of the buccal cavity coursing next to the trachea and reaches its origin on about the middle of the hyoid cornu, which lies, at this point, on the dorsal face of the neuromandibularis. M. protractor laryngeus (pl) Fig. 6 a-b This short, flat muscle takes origin from the posterior tip of the interramal pad and adjacent lateral head of genioglossus. It passes medially, running in the dermis of the buccal integument. Its ante- rior fibers meet their contralateral partners at the midline posterior to the orifice of the tongue sheath. The posterior fibers insert on the anterior tip of the glottis. DISCUSSION Jaw musculature of Alsophis cantherigerus brooksi conforms closely to the descriptions of several other colubrids (Albright and Nelson, 1959; Varkey, 1973; Haas, 1973; Cundall, 1974). The inter- nal divisions of the adductor mandibulae externus medialis, adduc- tor mandibulae externus profundus, adductor mandibulae posterior, and depressor mandibulae that were noted are often not included in these earlier papers. This probably does not indicate species differ- ence, but rather differences in investigator emphasis in selecting subtle internal detail of muscle to include in the descriptions. There does, however, seem to be an actual anatomical difference in pro- tractor quadrati. As suggested by the name, this muscle inserts on the quadrate in Opheodrys (Cundall, 1974), but here in A. ©. brooksi, Elaphe (Albright and Nelson, 1959), and Nerodia (Varkey, 1973) it inserts on the retroarticular process of the mandible. As in most other colubrids, a definitive M. /evator anguli oris is absent, but a weakly developed M. constrictor colli is present. In regard to lateral jaw musculature, the subspecies of Alsophis cantherigerus are a uniform morphological group (sensu Bock, 1963). As in A. c. brooksi, the adductor mandibulae externus super- ficialis arises from the anterior temporal crest and inserts as a broad aponeurosis on the lateral side of the mandible. A slip of fibers 1979 Jaw musculature of Alsophis cantherigerus brooksi 25 persists at the anterior edge of the muscle, continues around the corner of the mouth, and inserts directly to the mandible. The only exception is found in A. c. adspersus where this muscle inserts not on the mandible but to the epimysium of the adductor mandibulae externus profundus. However, this condition occurs only on the right side, the left being like other subspecies, and thus likely repre- sents an individual, rather than a subspecific difference. In all subspecies, the triangular adductor mandibulae externus profundus consists of anterior and posterior wedges of muscle separated by a vertical cleft that is often tendinous and forms a site of origin for-some fibers. Similarly in all, the adductor mandibulae externus medialis is separable into superficial and deep divisions, each with different sites of insertion. The adductor mandibulae pos- terior always possesses three parts—two major parts, lateral and medial, that pass to the respective sides of the prearticular crest and a third part, the pars minimus. In all, the protractor quadrati arises from a tendon shared with its partner and inserts along the medial retroarticular process. The general attachments of the depressor mandibulae are similar in all subspecies, and the muscle is divisible into superficial and deep portions divided by the insertion of the cervicoquadratus. However, the deeper part tends to split length- wise in fuscicauda, pepei, and ruttyi into two parallel divisions, one arising from the posterior corner of the quadrate, the other from here and from the posterior end of the supratemporal. Both, how- ever join to insert directly on the retroarticular process. In none of the subspecies examined did the lateral jaw muscula- ture differ fundamentally as to specific bones involved in the attach- ment sites. The observation that the only differences are in the nature of internal muscle anatomy again emphasizes the uniform nature of these island subspecies. ACKNOWLEDGMENTS I am grateful to Dr. Ernest E. Williams (Museum of Comparative Zoology) for the loan of the specimens of A/sophis examined in this paper. My very special thanks go to Drs. David L. Cundall and Alan Savitsky for critically reading and offering thoughtful sugges- tions upon the manuscript. 26 BREVIORA No. 463 LITERATURE ChreED ALBRIGHT, R. G. AND E. M. NeLson. 1959. Cranial kinetics of the generalized colubrid snake Elaphe obsoleta quadrivittata. 1. Descriptive morphology. II. Functional morphology. J. Morph., 105 (2): 193-239; 241-291. AuEN, E. L. AND D. A. LANGEBARTEL. 1977. The cranial nerves of the colubrid snakes Elaphe and Thamnophis. J. Morph., 154(2): 205-222. Bock, W. J. 1963. Evolution and phylogeny in morphologically uniform groups. Amer. Nat., 97: 265-285. CUNDALL, D. L. 1974. The cranial osteology and myology of the green snakes, genus Opheodrys. Unpublished Ph.D. dissertation, University of Arkansas, Fayetteville & Little Rock. University microfilm 74-28, 078. DULLEMIJER, P. 1956. The functional morphology of the head of the common viper, Viper berus (L.). Archives Neerlandaises de Zoologie, 11: 387-497. Haas, G. 1973. Muscles of the jaws and associated structures in the rhyncho- cephalia and squamata. /n C. Gans and T. S. Parsons (Eds.), Biology of the Reptilia, 4: 285-490. KARDONG, K. V. 1972. Morphology of the respiratory system and its musculature in different snake genera. (Part 1) Crotalus viridis oreganus and Elaphe obsoleta quadrivittata. (Part Il) Charina bottae. Gegenbaurs morph. Jahrb., 117(1972): 285-302; 364-376. 1973. Lateral jaw and throat musculature of the cottonmouth snake Agkistrodon piscivorus. Gegenbaurs morph. Jahrb., 119(3):316-335. 1974. Kinesis of the jaw apparatus during the strike in the cottonmouth snake, Agkistrodon piscivorus. Forma et functio, 7: 327-354. Kocuva, E. T. 1962. On the lateral jaw musculature of the Solenoglypha with remarks on some other snakes. J. Morph., 110:277—284. LANGEBARTEL, A. 1968. The hyoid and its associated muscles in snakes. Illinois Biological Monographs, 38:1-—156. MaG.io, V. J. 1970. West Indian xenodontine colubrid snakes: their probable origin, phylogeny, and zoogeography. Bull Mus. Comp. Zool., 141(1): 1-53. PREGILL, G. K. 1977. Axial myology of the racer Coluber constrictor with emphasis on the neck region. Trans. San Diego Soc. Nat. Hist., 18(11): 185— 206. TausB, A. M. 1966. Ophidian cephalic glands. J. Morph., 118: 529-542. VARKEY, A. 1973. Comparative cranial myology of North American natricine snakes. Unpublished Ph.D. dissertation, Louisiana State University and Agricultural and Mechanical College, Baton Rouge. University microfilms, Ue ke ACME BO@KBIN” -!: CO. INC. 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