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CHURCHER QOdocoileus salinae and Mazama sp. from the Talara tar seeps, Peru 1962 THE ROYAL ONTARIO MUSEUM Cc. S. CHURCHER is Assistant Professor, Department of Zoology, University of Toronto and Research Associate, Life Sciences Division, Royal Ontario Museum. © Royal Ontario Museum 1962 PRECES wo s00, PRINTED BY THE UNIVERSITY OF, TORONTO PRESS ABSTRACT. Twocervids have been recognized from the tar seeps of Talara, Peru. One has been identified as Odocoileus salinae (Frick, 1937) on the basis of dental and skeletal characteristics. The remains of this species are relatively abundant and form the majority of the specimens recovered. An isolated small detached tine, and a single atlas smaller than any recorded for Odocoileus are assigned to the genus Mazama. Odocoileus salinae (Frick 1937) Pe RODUC TION A representative of the genus Odocoileus has inhabited the South American western equatorial littoral region since Upper Pleistocene times. The first remains of this form were described by Frick (1937) as Palaeoodocoileus salinae n. sp., the locality being given as “the coastal oilfields about Salinas”, Ecuador. This faunal horizon is called the Carolinian, after La Carolina, Ecuador. The type specimen described was an incomplete mandibular ramus (Frick Collection. American Museum of Natural History no. 28284) which is illustrated (Frick, 1937, Fig. 21, p. 207) to show the dentition, but no measurements are given. Six other elements are reported from the same site, four of them (a left antler, a fragmental left frontal and antler base, a left maxilla with M*—M®*, and a right mandible with P;-M;) are referred to P. salinae, while two (a left radius and a juvenile right metatarsus) are said to be “of P. gracilis size” (syn. Odocoileus peruvianus). However, Frick says “This (species) which may be equivalent to the Spillmann P. antonii, does not much exceed in dimensions the Recent Pudu. Several of the Equadorian dentitions are characterized in unworn to moderately worn specimens by the detached antero-inner and posterior lophs of the py (P41) (see fig. 21), the latter condition being partially reminiscent of that in Rangifer. The anterior ps (P4) fossette is closed only in certain worn teeth.” Frick indicates the fourth lower permanent premolar by “p,” using a lower case “‘p”, rather than “P,”, using an upper case “P”’, as will be done in this paper. Hoffstetter (1952) comments upon Frick’s remarks “Mais la contradic- tion entre cette observation et les faits est évidente: elle apparait d’abord dans les dimensions de la série dentaire figurée, qui dépasse de beaucoup celle du Pudu andin; d’autre part Ch. Frick lui-méme signale (p. 213) qu’un radius et un métatarsien immature, trouves dans la méme gisement, ont la méme taille que ceux de “P. gracilis” (c’est-a-dire Odocoileus peruvianus ct. ustus).” He considers the original description inadequate in defining a new species, but as only a single cervid species is found in the deposits of La Carolina and as this species conforms in the other characters described by Frick, he is able to place his material from the same horizon (Carolinian of Ecuador) within Frick’s Upper Pleistocene species Palaeoodocoileus salinae. The genus Palaeoodocoileus Spillmann (1931) is placed in synonymy with Odocoileus Rafinesque 1872 by Simpson (1945) and, therefore, P. salinae will be referred to as Odocoileus salinae (Frick 1937). Remains of Cervidae were obtained from the Talara tar-seeps by an 3 expedition from the Division of Zoology and Palaeontology, Royal Ontario Museum, Toronto, during January and February, 1958. This material was tentatively identified as a small representative of the genus Odocoileus (Churcher, 1959; Lemon and Churcher, 1961). Sufficient material has now been prepared for a more accurate diagnosis and for comparisons with O. salinae, as described by Frick and Hoffstetter, to be made. Remains of Mazama have also been identified, but most of the material can be ascribed to a single form Odocoileus. DESCRIPTION OF THE MATERIAL Skeletal elements from all parts of the body have been found, including well- preserved mandibular, vertebral, podial, metapodial and phalangeal speci- mens. Cranial and proximal appendicular specimens are either absent or very badly damaged and fragmentary. Much of the material derives from young and immature animals, the epiphyses of vertebral centra and appendi- cular shafts being frequently missing or separate, and 14 of the 29 premolar specimens show milk teeth. No complete skull or mandible has been found; incomplete specimens of antlers and frontal bones, premaxillae, maxillae, jugals and mandibles only being recovered. Specimens of the antlers and frontal bones show the region from the orbit to the frontal-parietal suture, but are broken medially before the sagittal suture. Portions of the interior of the orbit and cranium are also visible. The antlers possessed three tines, the first tine pointing posteriorly and dorsally from the shaft, the second posteriorly and horizontally, and the terminal dorsally and slightly posteriorly, the shaft curving backwards from the frontal boss and changing direction upwards and outwards. The presence of three tines is usual in Neotropical Odocoileus (Cabrera and Yepes, 1940); a restoration of the antler is shown in Fig. 1. The milk and permanent dentitions are well preserved. Table I gives the numbers available and the lengths of the crowns of the teeth. Cranial frag- ments other than of the maxillae and antler-bosses are too small for measure- ment and no conclusions can be reached from their study, other than that they are from a small cervid. Mandibular specimens show the whole of the lower jaw and its dentitions (Fig. 5), with the exception of the anterior symphysial region bearing the incisors and canines, although two lower right third incisors have been recovered. Vertebrae recovered included atlas, axis, posterior cervical, thoracic, lumbar and sacral specimens in varying conditions of preservation. Rib and sternal fragments have also been found, but no certain caudal vertebrae. Elements of the forelimb recovered include damaged scapulae, shafts and distal articulations of humeri, proximal fragments of ulnae, entire and frag- mental radii, trapezoid-magna, scaphoids, lunars, pyramidals, unciforms, metacarpi and phalanges I, II, and III of digits HI and IV. No qualitative distinctions could be made between the phalanges of the manus and pes, nor did quantitative analysis indicate size differentiation. No remains of the vestigial digits IT and V have been found. Elements of the hindlimb recovered include parts of the pelvis, epiphysial regions of the femora, and tibiae, shafts of the tibiae, one fibula, calcanea, 4 ad sN—ed re € 6€ eN—IWN Wy 5! LS ¢ él eW I Cele eGurleeGule wOnSIE (oil! SG VAll 2 Pale ae SBE ESI SE ESL tel OPE ISS We ct IW = = td—zd LO 86 L Ol By (yO rd = =. 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No remains of vestigial digits Il and V have been recovered. COMPARISON OF THE TALARA CERVID MATERIAL WITH Odocoileus salinde RECOVERED FROM LA CAROLINA, ECUADOR, BY FRICK AND HOFFSTETTER, AND WITH THE LIVING LOCAL FORM The status of the living local species of Odocoileus is in some doubt. Hoff- stetter, in comparing the local extant form of Odocoileus ranging throughout the littoral region from the island of Puna to the Santa Elena Peninsula, mentions that this form has been neither described nor named, although he considers that it shows affinities to the Odocoileus gymnotis (included in O. virginianus cariacou; Hershkovitz, 1958) from Venezuela, Northern Brazil, Colombia and the Guianas. Hoffstetter refers to the living littoral form as “Odocoileus sp. X”, Spillmann (1948) refers to it as O. punensis n. sp., but gives an inadequate description. Hoffstetter further considers that O. sp. X and O. salinae might not be specifically different and separate from O. gym- notis, but might be a subspecies of the latter. Hershkovitz (1958) considers all the South American forms of Odocoileus conspecific with O. virginianus, and says (pers. comm.) “The teeth and mandibles of your (fossil) South American deer can be matched closely with those of our specimens of South American Odocoileus virginianus. On the basis of this evidence alone, I would regard the fossil (from Talara) as conspecific with Recent whitetails.” It may be, therefore, that O. salinae is but a subspecies of O. virginianus, but in this paper the name O. salinae will be retained for the fossil form, while the living local species will be referred to as O. sp. X in conformity with Hoffstetter’s usage. CRANIAL MATERIAL The cranial material includes fragments of antlers, incomplete dental series and isolated teeth, both milk and permanent, fragments of the skull, e.g. pre- maxillae, supraorbital ridges, parts of the braincase, occipital condyles and cther less identifiable fragments. Only the antler and dental specimens are complete enough for comparisons to be made. Six antler fragments have been recovered from the Talara tar-seeps, Royal Ontario Museum Palaeontological Specimens number 2080, 2104, and 2711 showing the portion from the orbital margin and braincase to the shaft distal to the junction with the first tine, 2710 from the region of the bony boss, past the first tine to where the rack broadens out to give rise to the second tine, 2714 showing the shaft between the two tines, the junction with tine two and the continuation towards the tip of the rack, and 2808 showing the tip of the antler or terminal tine almost complete. Thus all except the tip of the second tine is known from at least one specimen. (Throughout this paper the specimen numbers of the material in the Royal Ontario Museum will be given without prefixes and all are four-figure numbers. ) Hoffstetter (Fig. 109a) illustrates a fossil antler (V. 1431) of O. salinae, which is similar to specimens 2104, 2710, and 2711, from Talara. When the distance from the rugose base of the antler to the fork of the first tine is compared for the four specimens, those from Talara measure 50.5, 46.3, and 50.3 mm. respectively, while V. 1431 measures 73.5 mm. (Hoffstetter, pers. comm.). It seems therefore that the antlers of the Talara Odocoileus popula- tion were less developed than those from La Carolina, being approximately s smaller. Additional measurements on the Talara specimens are, from the base of the antler to the fork of the second tine, 129.4 (2710), and the total length of the antler along the outside of the curve, estimated 165.0 mm. (2710). The length of the first tines measured from the fork to the tip is 19.1 (2104), 20.0 (2710), and 21.4 mm. (2711). (Throughout this paper all measurements will be given in millimetres and estimated figures followed by-e’:) The antler bases all show a well-developed ring of rugosities, but these are much less strongly developed in the littoral form. Both fossil series show a marked ring (Fig. 1, and Hoffstetter, Fig. 109a and b, p. 358) while the living Odocoileus sp. X shows a rugose thickening of the basal section of the shaft. The angle between the first tine and the shaft in all three Talaran specimens appears to be greater than in O. salinae of Hoffstetter or either of the living species. The angle of the frontal bosses and the bases of the antlers to the skull shows a strong posterior inclination, and the antlers arise from elongated bony peduncles of the frontals, which agrees with the characters ascribed to O. salinae by Hoffstetter. The dentitions of the Talara specimens agree in general with those of O. salinae as described by Frick and elaborated by Hoffstetter. Specimens 2418 and 2728 containing a moderately worn full P.-M; and M,—Ms; dentition respectively and 2513 with a slightly less worn P;—Ms3 row, agree in size and most details of external styles, cingula, etc., with the lateral occlusal view illustrated by Frick (Fig. 21, p. 207). The characters of Ps quoted from Frick (vide supra) and also illustrated (Frick, Fig. 21, p. 207) hold in general for the Talara specimens, although some variation in the continuity of the protoconid-metaconid lophid is apparent. The paraconid (antero- internal cusp) is free from the protoconid in three of the four specimens recovered, although in some cases a medial projection of the protoconid meets the paraconid at the base of the cusp within the fossette and posterior to the maximum height of the paraconid cusp (Fig. 5). As wear proceeds this medial projection joins the paraconid to the protoconid by a posterior transverse ridge. The hypoconid (posterior loph) is always free in unworn P,’s, but with wear becomes attached to the posterior extremity of the proto- conid, forming a continuous W-shaped ectolophid. The metaconid-proto- conid lophid is continuous in all four P, specimens, and the anterior P, fossette is closed anteriorly, but opens posteriorly to the lingual side between the paraconid and metaconid in three of the four specimens. Some variation in the position and development of the lateral styles is noted, particularly of the buccal style lying between the protoconid and hypoconid of Py. This style may or may not be present, or may show varying degrees of development. The lower and upper milk dentitions and permanent lower and upper 7 molar dentitions exhibit no qualitative peculiarities that can be observed (Figs. 3, 4, 5, and 6). Table I gives measurements of the length of the crowns of the upper and lower milk and permanent dentitions. Comparison of these measurements with those for O. salinae (after Hoffstetter) and with the local living Odo- coileus sp. X (after Hofistetter ), shows marked similarity between the ranges of variation and the means given for all three populations. Agreement between the measurements of O. salinae and the Talara specimens is seen in 16 of the dimensions given in Table I; in M’ and M; there is approximate agreement, but since the posterior molars are subject to a wide degree of variation, this is not considered significant, and the variation in the crown length of M'—M° is probably partly due to the variation in the length of M’. The conclusion may therefore be drawn from the dental evidence that the Talaran cervid is identical with Odocoileus salinae (Frick 1937), such differences as can be observed lying within the possible range of variation for a single species. The postcranial material from Talara is more abundant than that reported by either Frick or Hoffstetter. VERTEBRAL COLUMN The vertebrae are similar to those of the species of Odocoileus, but are smaller than those of any of the species with which they have been compared. Cervical Thirteen specimens of cervical vertebrae have been recovered, including four partial atlases, two broken axes, two Cervical III’s, two Cervical IV’s, one Cervical V, two Cervical VI’s, and one Cervical VII. Plate I, Figs. 2 and 7, illustrate the most entire atlas, 2565, and the most complete axis, 2585. Measurements of the atlases 2565, 2564, and 2724 are: length along the dorsal midline 26.0, 24.5e, and 23.0; length along the ventral midline 26.1, 24.9, and 24.6; width across the posterior wings 65.2, 57.6e, and —; width across the condylar articulations 45.4, 43.6, and 44.1; dorsoventral depth 32.4, 29.8, and 30.0 respectively. Measurements of the axes 2585 and 2082 are: length along the dorsal midline 47.2, —; width of the odontoid spout 19.1, 21.6; and width of the odontoid collar 42.1 and 43.8 respectively. Measurements of Cervical III 2602 are: length along the dorsal midline 29.3; length from pre- to post-zygapophyses 50.7; width across the prezygapo- physes 29.2; width across the postzygapophyses 29.2; and dorsoventral maximum depth 34.2. Similar measurements of Cervical IV 2603 are: 30.2, 50.1, 33.4, 32-1; and 34.5: for Cervical V 2604 are: 28-le, 53:8) 40. 2e5 3 and 40.8; no measurements could be obtained from Cervicals VI and VII. Thoracic Ninety-seven specimens of entire or partial adult or juvenile thoracic verte- brae are present. These specimens represent ail the thoracic vertebral ele- ments when compared with a column from living Odocoileus, but are smaller than any of the elements with which they have been compared. A small Thoracic VII or VIII 2607 is illustrated in Plate I (Fig. 8), the measure- 8 ments for which are: length along the dorsal midline 28.6, proximal length of the neural spine 62.0e, width across the prezygapophyses 40.8, and width across postzygapophyses 10.8. Lumbar Eleven specimens of entire or partial adult or juvenile lumbar vertebrae are present. These are mainly from young individuals, as are all the vertebral elements, since most of the vertebrae lack their central discs. Measurements for Lumbar V? 2083 (Fig. 24) and Lumbar VI? 2084 are: length along the dorsal midline 25.8, —; length from pre- to post-zygapophysis 47.0, —; width across prezygapophyses 36.4, —; width across postzygapophyses 31.5, 40.5; and length of centrum 37.0e, 33.7 respectively. Sacrum A single example of an adult sacrum has been recovered, 2085 which is illu- strated in Plate III (Fig. 25). Measurements for this specimen are: length along the dorsal midline 55.0e, width across the prezygapophyses 53.5, length from prezygapophysis to distal lateral margin on the right side 75.6, and depth from dorsum of prezygapophysis to ventrum of centrum 36.6. Caudal Only two fragmentary and juvenile vertebrae that may be caudals have been recovered. Ribs Fragments of many ribs have been recovered, but none of them was entire or adult so far as can be judged. Other fragments of vertebrae or ribs have also been recovered but were too damaged or fragmentary to be identified in any respect. FORELIMB Scapula The scapulae are mainly represented by the glenoid, coracoid and spinal regions and again are similar to those of living Odocoileus except for a reduced size. Measurements of eight scapular specimens, 2153, 2383, 2384, 2721, 2744, 2745, 2746, and 3438 are: dorso-ventral diameter of the elenoid tossa 16.8, 20:3,'21.35 21-9 116.6, 22.9; ‘20.5; and 20-4: coracoid process-glenoid fossa diameter —, 31.5e, —, 31.4, —, 34.2, 33.2, and 31.5; maximum width of the neck of the scapula at the level of the spine 14.1, 15.5, —, 17.8, 13.6, 20.4, 18.8, and 18.7 respectively. No further measurements were possible as the blades of the scapulae are incomplete; specimens 2153 and 2744 are probably from immature animals as the absence of coracoid processes may indicate an unfused condition at the time of death. Humerus Entire humeri have not been recovered. The remains of humeri are mainly of the distal articulation with varying lengths of attached shaft (Fig. 9). 2 Specimens 2767, 2768, 2770, 3434, and 3435 measure 29.6, 31.8, 35.4, 31.4, and 32.5 across the articulation and 2767, 2768, and 3434, 12.8, 14.4, and 15.5 by 15.5, 16.8, and 18.6 for the minimum width and depth of the shaft respectively. Radius One entire adult radius, 2762, four almost adult radii lacking only the distal epiphyses, 2764, 2896, 3422, and 3438, two entire juvenile radii lacking both epiphyses, 2765 and 3424, ten fragmentary specimens of both adult and juvenile radii, six distal and one proximal detached radial epiphysis have been recovered. Measurements of the proximal ends of the fragmentary or nearly adult specimens, 2896, 3422, 3438, and 2510, 2713, 2763, 3427 are: maximum width 31.6, 29.6, 29.6, 28.2, 27.2, 26.5, 27.7 and maximum depth 18.0, 17.4, 17.4, 16.9, 16.4, 16.9, 16.7 respectively. The entire speci- men 2762 (Fig. 15) measures 28.5, 18.3, for the proximal width and depth respectively, 17.1 and 10.9 for the width and depth of the middle of the shaft, 26.2 and 17.6 for the width and depth of the distal end respectively and the total length of the shaft is 158.1. The largest detached distal epi- physis, 2725, measures 28.7 by 22.6 for its width and depth respectively. The only detached proximal epiphysis, 3432, measures 41.9 by 41.3 for the same respective dimensions. Ulna Only proximal ends of ulnae have been recovered. Adult specimens 2729 (Fig. 23) and 3428 measure 32.2 and 33.2 in maximum depth, 37.6 and 40.0 in dorsal length, and 46.4 and 46.5 in the length from the middle of the articulation to the extremity of the olecranon process; the depth of the shaft of the ulnae anterior to the radial facets is 15.5 and 15.4, and, at the centre of the articulation, 21.3 and 20.7, respectively. Carpalia Five trapezoid-magna, four scaphoids, two lunars, three pyramidals, five unciforms and three sesamoid carpals have been recovered. A left scaphoid, 21S Ere: 14), anda right lunar, 2774 (Fig. 20), are illustrated in Plate I. The scaphoids, 2775, 3445A and B, measure 14.4, 13.6, and 15.6 in dorso- plantar depth, compared with 11.6, 12.3, and 13.7 reported by Hoffstetter (1952) for O. salinae. Measurements for the other elements are given in Table II. Metacarpus The metacarpal is represented by three entire and one fragmentary adult specimen and by six entire juvenile specimens all of which, except one, lack the distal epiphyses, and by five fragmentary juvenile specimens. Compati- son of the adult metacarpals, 2560 (Fig. 13), 2414, and 2845, with that shown by Hoffstetter (Fig. 106d, V. 683, p. 353) shows them to be simi- larly built, and to most nearly resemble O. salinae rather than O. peruvianus, either fossil or living. 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PHALANGES OF THE MANUS AND PES Hoffstetter (1952) gives some dimensions and measurements for the phalanges of both feet of O. salinae, but does not indicate whether the phalanges of the manus differ qualitatively from those of the pes. Since he cites measurements for only seven specimens, two phalanx I’s, four phalanx II’s and one phalanx III, his problem of deciding from which foot they were derived was relatively easy. As the phalanges of the manus are generally considered to be smaller than those of the pes, the seven recovered were in all probability divided into large and small groups. The Talara tar-seeps have yielded 28 adult phalanx I’s (Fig. 18), 28 adult phalanx II’s (Fig. 17), and 18 adult phalanx III's (Fig. 16) from adult animals. They have all been examined for possible qualitative differences that would enable a separation into manus or pes, but without success. They have all been measured for their maximum length, maximum proximal width and maximum proximal depth, and these figures plotted for a bimodal dis- tribution. In all nine distributions there is no absolute indication of a bimodal curve, the curves approximating normal distributions in all cases. All the specimens may be considered to be fully adult as the proximal epiphyses are well fused to the shafis in all cases. In addition to the adult phalanges dis- cussed here, 19 juvenile shafts and nine detached epiphyses of phalanx I’s, 15 juvenile shafts and one detached epiphysis of phalanx II’s and one juvenile phalanx III have been recovered. Table II gives the comparable figures for the Talara specimens and those for O. salinae and O. peruvianus (= O. virginianus ) ustus (after Hoffstetter, 1952). The figures given by Hoffstetter for O. salinae lie closer to those for the Talara specimens than do those for O. peruvianus, and, in fact, in all but one case fall within the range of variation of the Talara specimens. There is thus no method of distinguishing between the Talara phalanges and those of O. salinae recorded by Hoffstetter, and it is not possible to support Hoffstet- ter’s identification of manus and pes elements, as his figures lie close to one another or to the upper limit of the range, and therefore should be considered to be possibly from the same foot and possibly from the pes, though such conclusions are speculative. HIND LIMB Pelvis Portions of 13 halves of pelves have been recovered but only specimens 2151 (Fig. 21), 2730, and 2731 are sufficiently complete for measurement. The maximum width of the iliac process is 24.0, 22.4, and 23.5, the maximum width of the ischial process is 24.6 (2151 only), the iliac-ischial diameter of the acetabulum is 28.6, 30.0, and 29.0, and the maximum width of the pubic process is 13.8, —, and 13.4 respectively. TZ 13 & ox CG OG rc SiGe aunc OES — sae yysugy [POL LIT XUepedqd G 6 61 LOZ CAG il laa — — yydep jeurxoid WNwWIxe [Y G 6° FI 8ST ZV i ell — — yIpIM peurxord WUNWIXe JA a 6 Ge 0°S& 6 GS I G 0G <7, ee yysug [PIOL See etl 6 Oo OL 0: Ue ¢’ SI if 1°91 ass oe yydop yeumxoid winturxe [Yl 6 IE 0 91 8 FI it c GL = = y pia peurxoid WNWIXe | 6 Zi om 0 Si Oe Ly I 0 CF a = yysug] [P10 L j xuereyd aAWITGNIH SI | 6 SI 38 Gl = ran = aa = = — = yidep jewrxoid [euUlxe |] rol 6 8 Tl 0 2 - ie ae ac = a == — YPM jeurxoid WNUIXe Jl 81 8° Gz G' 8G 8 1G G 6 VE Gece Ose cl OS as yy sus] [ROL III xueredd eater 9 C1 OL = Zoli OL Oe Oe = 0 6t) 44 ON yydap yeurxoid winuixe |] SG ge | Gn 8° 6 OL Grail Gel EVAL ¢ — aia GO y pM pewrxo.d WIN UWIX® JA] 8G 8 VG G 86 0 16 Ol V 66 Seite, 0 86 & a 0 16 & 9G yy sug] [POL Ph Serle del 8G 8 CT Csi G Vi 9 OnZl 0 61 ey i! € St = = yydop yeurxosd wnuixe |] 8G G Gl v Si 8 Ol 9 cae Gaal 8 sl I S Gl = ae: yy pm peurxoid wWNwrxe [Ay 8G 8° 9e L'0V 6 && ¥) G OV LGV V 6& \! G°6& aa ane yr sug] [20 L, pees tal ANITANO +] iro. a eS N Xx EIN TN N x FEW CONT N x SIN INL suawloads viele |, (193790}SJJOH) (19}}03S]JOH) snupinnsad $naj10j0pO aDUIDS SNA]L0IOPO “SOIJOUNT[IU UL UAATS 1v syUstUsINsvaW [[Y “Quuy Toys 0} HI USISsse JOU seop oY SB quuT[pUIH{, Pue quiet J, YOq JopuN VIM UDAIS ST HI ‘19}193SJOH Aq paysodes [j{] xuevjeyd ATU 9y} Jo vsvo oY} UT ‘1n3y oy} SUIMOT[OJ .¢, AQ PezeoIPUl 1e BST} ‘saInBY $,10}}9}SHYOH JO ouos OF S}S!Xo AVUIe}AIOUN JWIOG *(SeINsSy ,qUITeIOJ, IY} ayisoddo) dnois opsuls & se UdAIZ ore sasuRjeYyd prAjoo uvIEeL, 94} [IP 40} soinsy ay} A;JuUenbesuod ‘sad pue snurw ojuUr payeredas aq OU PfNoo PIAtIo wire], oy} Joy sasuvyeyd ey] ‘apurps ‘OC OY TRUS PIAdao eleyey, ay} JO esoy} YIM (ZG61 ‘FaOISJOP] 1037e) (SNUMIMIG1I2 ‘CE =) snuniniad ‘CQ pur apurps SNajvoI0PQO JO sod pur snueur ay} jo sasueyeyd oy} Jo sjuateinsvau oAtereduiod “[[] ATaV Femora No entire femora have been recovered, although a number of ends and detached epiphyses are represented in the collection. The distal epiphyses of the femora are represented by five entire and two fragmentary specimens and four chips. The three largest, 2748B, D, and E, measure 41.8, 46.3, and 41.0 respectively across the condyles and 53.3, 55.0, and 53.1 respectively on the anteroposterior diameter. No other measurements of femoral fragments are available. Tibiae The tibiae are represented by five proximal and eleven distal epiphyses, one adult proximal fragment with part of the shaft attached, 2751, three frag- mentary immature shafts lacking the distal epiphyses and much of the proximal regions, two adult distal fragments, three adult shaft fragments, one with the distal end, and two adult specimens lacking only the proximal articular regions, 2081 (Fig. 22) and 2753 and one immature shaft lacking both epiphyses. From specimens 2081, 2751, and 2753 the following dimen- sions were measured: Total length 230.0e, —, 237.0e, proximal width 45.0e, 51.6, —, proximal depth 45.0e, 45.7, —, maximum diameter in the middle of the shaft 16.5, —, 20.1, minimum diameter in the middle of the shaft 12.6, —, 15.6, maximum width of the distal articulations 28.5, 31.2, 31.2 and, minimum depth of the distal articulations 20.8, 22.6, 22.5 respectively. Fibula Three proximal fragments of fibulae have been recovered. The largest, 3439 (Fig. 34) measures 111.6 in total length, 15.5 anteroposterior diameter of the proximal end and 8.7 width of the proximal end. TARSALIA Twenty-seven whole or fragmentary specimens of adult and juvenile cal- canea, 35 astragali, 18 navicular-cuboids to one of which the cuneiform is fused, three free cuneiforms, and one malleolus, have been recovered from the Talara tar-seeps, but few undoubted mature bones can be recognized. Calcaneum The calcanea are variable in development, 22 of them are undoubtedly im- mature, and only three, 3100J, 3099A and G of the entire specimens and four fragmentary specimens, 3099M, 3100C, K and M have the posterior tuber calcis region coossified with the body of the calcaneum. Comparison of the certainly adult calcanea, e.g. 2799 (Fig. 10), with that illustrated by Hoffstetter (Fig. 108 c, V. 690, p. 356) shows that the Talara specimens are similarly slim and similarly shaped to his O. salinae specimen. Measurements for this tarsal bone and for the other tarsalia are given in Table IV. Astragalus Only nine of the 35 astragali recovered can be considered to be adult and fully developed (Fig. 11). Measurements for these adult specimens only are given in Table IV. 14 Wy €°9¢ 896 3G & & 9G PSG = 8 GS 9 SZ G 86 = L GZ YPM Ie[Apuod [eqsIq ¢ 6 bo £96 I & p96 & SZ are TE ai 1 &@ re b 6 NDE rena 9 G chee wy Gach nd 3 I ¥I 8 él ia! 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ON Xx “xe “UL (409394syo}H]) SNAHWIOddS VAWIVL ADUYDS $N9]L0I0P() ‘SOIJIWI[IU UT dIv sJUBWBINSvaU! [TW “(ZEGT ‘JoVesOH Joye) snyesesjse oy} pure wumnouroyeo dy} JOJ A[UO BIGLIIVAR d1v aDUI]DS Snaj1oI0PE JO} SaInsy IATyeIedwoos sy], “PAI URIETe] ay} JO IsteyeJIPY pue eI[eSIB] dy} JO s}UdWINseI| “A] ATAV];, Ip Navicular-cuboid Of the 18 navicular-cuboids recovered, ten are undoubtedly juvenile and eight seemingly adult. One of the probable adult specimens is fused with its neighbouring cuneiform (Fig. 19), which supports its identification as adult since such abnormalities are more common in old (or injured) animals. Measurements for the navicular-cuboids and for the single fused cuneiform are given in Table IV. Cuneiform The three free cuneiforms are small and probably immature. No measure- ments for these are given. Metatarsus The metatarsal is represented by two entire and six fragmentary adult speci- mens and by three entire almost adult specimens lacking the distal epiphyses, by three smaller entire juvenile shafts also lacking the distal epiphyses and by some 14 fragmentary specimens of various ages. The metatarsal elements are longer than the metacarpal elements as is expected. An adult metatarsus 1980, is illustrated in Plate II (Fig. 12) in comparison with an adult meta- carpus, 2560 (Fig. 13). Measurements for the adult metatarsi are given in Table IV. The measurements given in Tables I, III, and IV support the qualitative conclusions, for the appendicular skeleton as a whole, although the few measurements given by Hoffstetter suggest that the Talaran cervid may be slightly shorter in the leg than that from La Carolina. Hoffstetter mentions that his material is scarce and that some of his specimens are undoubtedly from a large adult. The discrepancy in the proportions of the limb-bones may be, therefore, a partial result of too small a sample, together with the accident of a large specimen on the one hand compared with two younger or smaller specimens on the other, perhaps respectively male and female. Mazama sp. A single entire atlas, 2703 (Fig. 35), and a detached right antler, 3151 (Fig. 36), have been recovered from the Talara tar-seeps. These remains do not fit the characters of Odocoileus salinae and consequently are described here separately and ascribed to Mazama sp., possibly M. gouazoubia. The atlas is almost entire, lacking only the posterior portion of the left wing. It has been compared with the atlas from an individual of Mazama gouazoubia, (American Museum of Natural History Specimen no. 130459) and found to resemble it both morphologically and quantitatively. Compara- tive measurements for the various dimensions of the Talara atlas and that of M. gouazoubia are respectively: width across the wings—44.9 and 44.7; length of the wings—39.8 and 35.8; width of the condylar articulations— 38.5 and 33.7; width of the axial articulations—33.le and 30.6; dorsoventral depth—25.5 and 27.7. While these figures are not in absolute agreement, it is evident that the Talara specimen is of approximately the same size and proportions as is the Recent specimen. 16 The detached right antler is almost entire also, having been split off at the base just distal to the bony boss. The curvature and cross-sectional propor- tions are similar to that found in many specimens of Mazama from South America, while the rugosities on the surface indicate that the antler had been but recently renewed and probably the velvet had not yet been rubbed off. The specimen measures 56.6 in total length. DISCUSSION AND CONCLUSION The more abundant form from Talara numbers some 18 or 20 individuals as the minimum number of juvenile and adult animals from which the recovered elements could be derived. The uncertainty as to the actual minimum number exists because of the difficulty in matching partial juvenile specimens of nearly similar ages. The remains of the less abundant form could have been derived from a single individual. The more abundant fossil form from Talara strongly resembles Odo- coileus salinae in its dental characters and measurements. In its post-cranial measurements there is less agreement, perhaps because of too small samples and too many juvenile specimens recovered. The characters of the antlers agree qualitatively, but there is a disagreement in the size, those of O. salinae reported by Hoffstetter being about 50 per cent larger than those from Talara (73.5 cf. 46.3-50.5). This difference would indicate a different population at Talara, similar in general appearance with O. salinae from the Carolinian of Ecuador, but with smaller antlers and a slightly reduced stature, and with otherwise similar proportions. Since variation in the antlers of deer is known to be great, dependent upon age and nutritional factors, especially within the tropics, it is probable that this divergence is not signifi- cant at the specific or subspecific level. The Talaran Odocoileus may be con- sidered conspecific with Odocoileus salinae (Frick), though differences in the skeletal and antler measurements may indicate some separation at the population or subspecific level. The larger antlers in the Carolinian popula- tion may indicate a more easily available source of calcium, and possibly a cooler climate, and thus the divergence between the two populations may reflect differing environmental factors rather than genetic differences. The larger size of O. salinae reported by Hoffstetter, may be considered tenta- tively unrepresentative of the population, as Hoffstetter considers it to be a large adult. The less abundant form is assigned to Mazama (Rafinesque 1817) cf. gouazoubia on the basis of the similarity in size and shape of the atlas, which is the only reliable skeletal element recovered which gives some indication of the size of the animal, and this identification is supported by the antler specimen which bears a strong likeness to the antlers of some living indivi- duals of Mazama. 17 ACKNOWLEDGEMENT The author wishes to record his thanks and appreciation to Dr. W. E. Swinton, Head of the Life Sciences Division of the Royal Ontario Museum, for allowing the use of the facilities at his disposal and also to other members of the staff of the institution for their comments and suggestions during the course of this work. 18 REFERENCES CABRERA, A. AND J. YEPES . Historia Natural Ediar. Mammiferos Sud-Americanos, p. 275, Buenos Aires: Compania Argentina de Editores, 1940. CHURCHER, C. S. Fossil Canis from the Tar Pits of La Brea, Peru. Science, 130: no. 93:75, 564—5,, 1959. FRICK, C. Horned Ruminants of North America. Bull. Amer. Mus. Nat. Hist., 692 205-214; 1937. HERSHKOVITZ, P. The Metatarsal Glands in White-tailed Deer and related forms of the Neotropical Region. Mammalia, 22: no. 4, 537-546, 1958. HOFFSTETTER, R. Les Mammiféres Pléistocéenes de la République de Equateur. Mem. Soc. Géol. de France, no. 66, 346-369, 1952. LEMON, R. R. H. AND C. S. CHURCHER Pleistocene Geology and Paleontology of the Talara Region, North- west Peru. Amer. J. Sci., 259: 410-429, 1961. SIMPSON, G. G. The Principles of Classification and a Classification of Mammals. Bull. Amer. Mus. Nat. Hist., 85: 155, 1945. SPILLMANN, F. Die Saugetiere Ecuadors im Wandel der Zeit, t. I, 112 p., Quito: Unive Central, 1931. SPILLMANN, F. Beitrage zur Kenntnis eines neuen gravigraden Reisensteppentieres (Eremotherium carolinense gen. et sp. nov.), seines lebensraumes und seiner lebensweise. Palaeobiologica, 8: 231-279, 1948. 19 PLATE I Cranial and Axial Skeletal Elements of Odocoileus salinae. Right Antler, anterior aspect; restored from 2710 and 2711. Atlas, right lateral aspect; 2565. Upper milk dentition, left p?—p+; a—above, in buccal aspect and b— below, in occlusal aspect; 2069. Lower milk dentition, right p.—p,; a—above, in occlusal aspect and b— below, in buccal aspect; 2004 and 2512. Lower permanent dentition and mandible; a—above, left P.—M. in occlusal aspect and b—below, left P,—-M. in buccal aspect in situ in mandible. Dentition and ramus from 2418, symphysial region from 2733 and condyle from 2513. Upper permanent dentition, left P4‘-M*; a—above, in buccal aspect and b—below, in occlusal aspect; 2511. ) Axis lacking neural spine, left lateral aspect; 2585. Thoracic Vertebra VII or VIII, left lateral aspect; 2607. eye: GiGe Sive G5ld 31Sa+ YOO? SO alle | BUR alle! € é \ {0} SWO - 3|DIS LO “8 Old £ ‘g bid G9GS PLATE, If 22 Appendicular Skeletal Elements of Odocoileus salinae. Right humerus, distal half, posterior aspect; 2768. Right calcaneum, medial aspect; 2799. Right astragalus, dorsal aspect; 2804. Right metatarsus, anterior aspect; 1980. Right metacarpus, anterior aspect; 2560. Left scaphoid; a—left, lateral aspect and b—right, medial aspect; 2775. Right radius, anterior aspect; 2762. Phalanx III, external aspect; 2800. Phalanx I, external aspect; 2784. Phalanx I, external aspect; 2802. Right navicular-cuboid with fused cuneiform, anterior aspect; 2805. Right lunar; a—left, proximal aspect and b—right, lateral aspect; 2774. Right innominate, fragmentary, lateral aspect; 2151. ¢ Og ‘614 bide 6622 ‘Ol 'b'4 KGae ie able 23 2082 ‘8| ‘614 O08zZ ‘9| ‘bI4 GJ2mS “pl Bi er 09Sz2 ‘¢| ‘bi4 sans v iS Z ] fa) SWO- 8)/D9S bOse ‘1) 6l4 O86] ‘2| 614 PEADE. Tit 24 35 36 Postcranial Skeletal Elements of Odocoileus salinae. Left tibia lacking proximal epiphysis; anterior aspect; 2081. Left ulna, medial aspect of proximal portion only; 2729. Lumbar Vertebra V?, lacking neural and lateral processes, right lateral aspect; 2083. Sacrum in damaged condition, dorsal aspect; 2085. Left unciform; a—left, proximal aspect and b—right, lateral aspect; 3441. Left trapezoid-magnum; a—left, proximal aspect and b—right, medial aspect; 3440. Left pyramidal; a—left, lateral aspect and b—right, medial aspect; 3442. Left navicular-cuboid, lateral aspect; 2816. Right cuneiform, proximal aspect; 3234. Right humerus, proximal articulation: 3436. Left tibia, distal articulation; 2081. Left malleolus, lateral aspect; 2838. Right fibula, lateral aspect; 3439. Skeletal Elements of Mazama sp. Atlas, right lateral aspect; 2703. Antler fragment, lateral aspect; 3151. ¢ 31V1d 4 I SW - 8|D9S 9Erbe ‘Ie bI4 6eve ‘pe'bi4 8¢ese ‘ee bid veese OF bis ONO COG Ge Dig eye ‘82 6l4 Ippe ‘92 B14 092 PLATE IV 26 37-38 39 40 41 42 43 44 45 46 47 48 49-50 Photographs of Selected Skeletal Elements of Odocoileus salinae and Mazama sp. Odocoileus salinae. Right forefoot assembled from individual ele- ments, anterior aspect. Wrist composed of pyramidal 3444A, lunar 2774A, scaphoid 3445B, trapezoid-magnum 3447A and manus of metacarpal 2560, Phalanx I’s 2358N and 3793Q, Phalanx II’s 2364C and 2737D and Phalanx IIP’s 2413B and 2930H. O. salinae. Damaged atlas, ventral aspect; 2565. O. salinae. Right antler lacking tip of tine 1 and all of tines 2 and 3, anterior aspect; 2710. O. salinae. Right radius, anterior aspect; 2762. Mazama sp. Damaged atlas, ventral aspect; 2703. Mazama sp. Damaged antler broken distal to the boss; 3151. Odocoileus salinae. Damaged right innominate lacking ends of iliac, ischial and pubic radii, acetabular aspect; 2151. O. salinae. Immature left tibia lacking proximal epiphysis and dam- aged in that region, anterior aspect; 2081. O. salinae. Left ulna lacking shaft, medial aspect; 2729. O. salinae. Left antler boss showing complete tine | and superior orbital margin, anterior aspect; 2711. O. salinae. Assembled mandible with P,—-M. and damaged incisive and coronoid regions, lateral aspect; 2733 (mesial element) and 2513 (distal element). O. salinae. Left hindfoot assembled from individual elements, anterior aspect. Phalanges of pes omitted as those of manus shown in Fig. 37-38. Left metatarsus 3462; ankle composed of calcaneum 3100J, astragalus 2804L, navicular-cuboid 28161 and cuneiform 3234A. 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