The Late Miocene Siwalik giraffids, despite

Transcription

1 Pkistn J. Zool., vol. 44 (3), pp , Hydspitherium (Artiodtyl: Girffide) from the Dhok Pthn Formtion of the Middle Siwliks, Pkistn: New Colletion Zfr Hmeed Bhtti, Muhmmd Akr Khn* nd Muhmmd Akhtr Pleontology Lortory, Zoology Deprtment, Quid-e-Azm Cmpus, Punj University, Lhore, Pkistn Astrt. A few dentl remins from the lte Mioene lolities of the Dhok Pthn Formtion hve een identified s elonging to Hydspitherium f. megephlum. The olletion omprises n upper premolr, severl upper molrs, prt of mxill nd prt of mndile. The nlysis of the dentl hrters shows individul vritions nd ontriutes to the knowledge of the Siwlik lte Mioene girffids. The systemti sttus of the Siwlik speies of Hydspitherium hs een reviewed. Key Words: Siwlik girffids, Hydspitherium, lte Mioene, Dhok Pthn Formtion, Mmmli. INTRODUCTION The Lte Mioene Siwlik girffids, despite their rihness, re still rther poorly known nd their systemtis outdted, with unrelile identifitions (Mtthew, 1929; Gentry, 1997; Gerds nd Gule, 1999). They re minly represented in the Siwliks y the lrge-sized Sivtheriine nmely, Brmtherium, Hydspitherium, Helldotherium, nd Sivtherium (Mtthew, 1929; Colert, 1935; Gentry, 1997; Gerds nd Gule, 1999; Bhtti, 2005; Kostopoulos nd Sr, 2005). Reently, few well preserved lrge girffid speimens were olleted from the villge Dhok Pthn (Lt 'N: Long 'E) nd the villge Hsnot (Lt N: Long E) of the Dhok Pthn Formtion, the Middle Siwliks, Pkistn (Fig. 1). The girffid smple yields set of hrters, morphologilly similr to the lte Mioene Siwlik girffid, Hydspitherium (Mtthew, 1929; Colert, 1935; Khn, 2007). The omprison of the speimens with the known speies of Hydspitherium hs shown lerly tht the speimens elong to speies Hydspitherium f. megephlum. The girffid Hydspitherium megephlum is restrited to the Dhok Pthn Formtion of the Middle Siwlik Sugrou in the northern Pkistn (Sehgl nd Nnd, 2002; Brry et l., 2002; Bdgley et l., 2005). The detiled strtigrphy nd iohronolgy of the sites re given * Corresponding uthor: kr111@yhoo /2012/ $ 8.00/0 Copyright 2012 Zoologil Soiety of Pkistn. in Brry et l. (2002) nd Khn et l. (2009). The reovered girffid mteril is ssoited with remins of Trgoportx punjius, T. slmontnus, Selenoportx vexillrius, Phyportx ltidens, Gzell lydekkeri, Kous porretiornis, Cprotrgoides potwrius, Dortherium mjus, Dorune nthrotherioides, Propotmohoerus hysudrius, Hippopotmodon sivlensis, Sivlhippus theoldi, Sivlhippus perimense, Hipprion sp., Aertherium perimense, Chilotherium intermedium, Stegolophodon ltidens nd Crout rnifex (Pilgrim, 1937, 1939; Heissig, 1972; Bernor nd Hussin, 1985; Akhtr, 1992; Bii, 2007; Khn et l., 2009). The ssemlge fits well with the Siwlik lte Mioene fun, inditing lte Mioene ge (Brry et l., 2002; Khn et l., 2009). Four iostrtigrphi intervl-zones re proposed in the Potwr Plteu of the northern Pkistn in whih the iostrtigrphi studies re lued with mgnetostrtigrphi ges (Brry et l., 1982). These zones re Hipprion sl. Intervl- Zone; Selenoportx lydekkeri Intervl-Zone; Hexprotodon sivlensis Intervl-Zone nd Elephs plnifrons Intervl-Zone. It is notied tht Hydspitherium megephlum ours in the Selenoportx lydekkeri Intervl-Zone (7.4 to 5.3 M) of the Middle Siwlik Sugroup (Brry et l., 1982). The primry gol of this pper is to present new olletion of Hydspitherium from Dhok Pthn nd Hsnot in the northern Pkistn (Fig. 1). The unpulished olletion ould gretly improve our knowledge of the Lte Mioene girffid, Hydspitherium. Gerds nd Güleç (1999) suggest

2 800 Z.H. BHATTI ET AL. Fig. 1. The lotion of Dhok Pthn nd Hsnot in the northern Pkistn where the desried mteril hs een found. Boundry dtes re from Brry et l. (2002) nd Nnd (2002, 2008). provisionl synonymy of Hydspitherium Lydekker, 1876 with Brmtherium Floner, 1845 in the Greo-Irnin provine. Although quite possile, this synonymy is not yet formlly founded nd, following Mtthew (1929) nd Colert (1935), we shll ontinue to use Hydspitherium s vlid txon in the Siwliks. Almost ll speimens were found wethering out from or in situ within the mtrix. Some speimens were exposed on surfe nd ut some re entrenhed prtilly. Piering instruments (Chisels nd Geologil hmmers) were used for mining of prtilly entrenhed fossils. The speimens were tlogued nd given numer whih onsists of yerly nd seril tlogue numer, so figures on the speimen represent the olletion yer nd the seril numer of tht yer (e.g. 67/155). Upperse letters with supersript stnd for upper dentition (e.g. M 1 ) nd with susript numer stnd for lower dentition (e.g. M 1 ). Vrious mesurements of the studied speimens in millimeters were tken with the help of metri vernier liper. Tooth length nd width were mesured t olusl level. Heights were mesured on the mesostyle of the upper molr, the metstylid of the lower molr nd the protoonid of the lower premolr. All the desried fossils re housed in the Pleontologil Colletion of the Punj University, Lhore, Punj, Pkistn (PUPCinstitutionl revition). Arevitions: PUPC, Punj University Pleontologil Colletion; AMNH, Amerin Museum of Nturl History; M, million yers; MN, Europen Neogene Mmmlin Zone; d, deiduous; M, molr; P, premolr; L, lrgest length; W, width; H, height; l, left; r, right. SYSTEMATIC PALAEONTOLOGY Order Artiodtyl Owen, 1848 Suorder Ruminnti Sopoli, 1777 Fmily Girffide Gry, 1821 Sufmily Sivtheriine Zittel, 1893 Genus HYDASPITHERIUM Lydekker, 1876 Type speies Hydspitherium megephlum Lydekker, 1876.

3 HYDASPITHERIUM FROM THE MIDDLE SIWALIKS 801 Arevited generi dignosis Teeth lrge, qudrte, with rugose enmel. Lower teeth roder nd somewht longer. The prstyle of Hydspitherium megephlum is stronger thn the mesostyle nd the metstyle. The nterior fossette is trnsversely ompressed (Mtthew, 1929; Colert, 1935; Bhtti, 2005). Strtigrphi rnge Middle Siwliks (Colert, 1935; Sehgl nd Nnd, 2002; Bhtti, 2005; Khn, 2007). Geogrphi distriution South Asi (Suontinent). The ourrene of the genus is reorded from the vrious lolities of the Potwr Plteu (Middle Siwliks) of Pkistn, Dhok Pthn, Hsnot s well s from Azd Kshmir, Bhimer (Lydekker, 1876; Pilgrim, 1910; Bhtti, 2005; Khn, 2007). It is reported from the lte Mioene nd Plioene sediments of Indi (Mtthew, 1929; Colert, 1935; Sehgl nd Nnd, 2002). Hydspitherium f. megephlum Lydekker, 1876 Type speimen A skull: GSI D150. Arevited dignosis The prstyle of Hydspitherium megephlum is prominent s ompred to the mesostyle nd the metstyle. The nterior medin ri is modertely developed, wheres the posterior medin ri is wekly developed. The enmel is rugose. The stylids re present nd the nterior fossette is trnsversely ompressed. Hydspitherium megephlum generlly is smller thn those of H. grnde nd H. mgnum (Colert, 1935; Khn, 2007). Studied mteril Upper dentition: PUPC 97/20, rp 4 (Dhok Pthn); PUPC 91/23, rm 1 (Hsnot); PUPC 67/155, lm 2 (Dhok Pthn); PUPC 97/16, right mxillry frgment with M 1-2 (Dhok Pthn). Lower dentition: PUPC 95/24, ldm (Dhok Pthn); PUPC 97/17, right mndiulr frgment with prtilly P 3-4 nd omplete M 1-3 (Dhok Pthn). Soure of the omprtive mteril Mtthew (1929), Colert (1935), Bhtti (2005). DESCRIPTION Upper dentition The upper dentition omprises one premolr, two isolted molrs nd one mxillry frgment with first nd seond molrs (Fig. 2). The molrs re modertely hypsodont nd they hve simple olusl pttern. P 4 : PUPC 97/20 is prtilly preserved nd in n erly wer (Fig. 2A). The premolr is greter in length thn in width, nd the tooth is hrterized y strong prstyle, nd n internl posterior swelling (Fig. 2A). The tooth is dmged on the lil side. The three rooted premolr hve thik nd rugose enmel. The ingulum is not developed. The fossette is well developed. The internl side of the ul resent is wekly divided into prone nd metone. The premolrs look suqudrngulr with n ntero-lingul protuerne of the lingul wll (Fig. 2A). M 1 : PUPC 91/23 is in n erly wer (Fig. 2B, D). The enmel surfe is highly rugose. The ingulum is poorly developed. It is slightly developed on the nterior side of the protoone. The entostyle is wek, pling in the trnsverse vlley etween the protoone nd the hypoone. The mjor usps re well developed. The ul usps re higher thn the lingul ones. The protoone is resenti in shpe, ut it is less resenti thn the hypoone. The pre-nd postprotorist run prllel to the longitudinl xis of the molr. The prone is pointed in the middle with the two riste sloping ntero-posteriorly. The postprrist is roder thn the preprrist, whih is somewht nrrow. The preprrist is united with the prstyle wheres the postprrist is touhing with the prehyporist. The metone is slightly higher vertilly thn the prone. It is pointed in the middle with two shrp running pre-nd postmetriste. The hypoone is present posterior to the protoone. The prehyporist ontts with the postprrist nd the posthyporist ontts with the metstyle. The styles re strongly developed. The mesostyle present t the nterior

4 802 Z.H. BHATTI ET AL. A B C D Fig. 2. Hydspitherium f. megephlum, upper dentition. A, PUPC 97/20 rp4; B, PUPC 91/23 rm1; C, PUPC 67/155 lm2; D, PUPC 97/16, right mxillry frgment with M1-2. Views re olusl () lingul () nd lil (). Sle r equls 10 mm.

5 HYDASPITHERIUM FROM THE MIDDLE SIWALIKS 803 side of the metonl loe is nerly s high s the prstyle ut it is omprtively wek nd infleted. The metstyle is weker thn the prstyle nd the mesostyle. The medin ris re firly developed. The nterior ri is stronger thn the posterior one. The nterior ri is rod t the tip nd nrrow t the se (Fig. 2B, D). M 2 : The molrs re nerly qudrte nd rugose (Fig. 2C-D). The ingulum is poorly developed on the nterior side of the protoone s well s on the posterior side of the hypoone. The fossettes seem to e wide nd resent. The prnd metones re higher vertilly thn the protond hypoones. The prone medin ri is stronger thn the metone medin ri. The styles re well preserved nd prominent. The prstyle is omprtively rod nd hevily uild. The prone is higher thn ll the other ones. The protoone is more pointed thn the hypoone. Between the pr nd metones the enmel is muh folded produing strongly developed mesostyle. The metone is olique nteroposteriorly wheres the prone is stright thn eing olique. The hypoone is onneted with the metone through nrrow ridge posteriorly (Fig. 2C-D). Lower dentition The lower dentition inludes one deiduous molr nd mndiulr frgment with two premolrs nd omplete series of molrs (Fig. 3). dm: The deiduous molr onsists of three loes of lmost similr shpe ut they hve vrition in width nd length (Fig. 3E). The nterior loe is muh longer thn the posterior one, primitive hrter found in ll Sivtheriine (Gerds nd Asln, 2003). The rown height is moderte. The lil hlves of the three loes re resent, wheres the lingul hlves re spindle-shped. The nterior most prt of the rown is reltively more worn. The enmel is modertely thik nd rugose. The deiduous molr is resenti-shped with widely divergent rms lilly. The lingul prt of the nterior most prt is strongly A-shped with two nterior nd posterior riste. The medin prt of the tooth is quite similr exept tht the posterior end of its lingul prt is produed into well formed style. The posterior loe of the rown is lest worn. The posterior rist of the lil hlf ers deep vertil groove. There re smll entostyles on the tooth (Fig. 3E). Mndile: PUPC 97/17 is frgile mndiulr frgment roken nteriorly nd posteriorly (Fig. 3F). Posteriorly, the mndile is roken posterior to the tlonid of the third molr nd it is roken nterior to the third premolr nteriorly. The premolrs re semierupted nd there morphologil fetures re visile only on the lingul nd on the olusl sides. The P 3 is more erupted thn the P 4. The semierupted P 4 n e oserved lingully in the mndiulr rmus. P 3 : The pronid is well seprted from the prstylid ut they re losed lingully (Fig. 3F). The metonid extends forwrds, meeting the se of the pronid nd the rest joining the protoonid to the metonid olique kwrds. The elongted metonid is independent from the endoonid. The tlonid is omprtively shortened. The redued endostylid is oliquely settled. Lilly, well developed furrow seprtes the ulgy hypoonid from the strong protoonid. P 4 : It is molriform (Fig. 3F). Its metonid is long nd the prstylid thinner thn in P 3. The rest joins to the metonid nd inorportes the entoonid. The entoonid is independent from the metonid. A well developed furrow seprtes the hypoonid from the strong protoonid on the ul side. The enmel is thik nd rugose. The entoonid is well distint nd olique. M 1 : The molr is in middle wer. The mjor onids re interonneted with eh other t the grinding surfe due to the middle wer (Fig. 3F). The etostylid is wekly developed. The longitudinl vlleys re not muh deep. The vlley etween the nterior ones is shllower thn the posterior ones. The ristids of the protoonid re quite simple. The metonid is firly developed. It is spindle-shped. It is slightly wider in the middle with two nrrow sloping ristids. The entoonid is reltively lower vertilly thn the metonid. The hypoonid is V-shped. The mesostylid is well developed s ompred to the met- nd entostylids. The metonid ri is modertely developed nd the entoonid ri is wekly developed. M 2 : The seond molr is modertely high. The length of the tooth is more thn the trnsverse width (Tle I). The ingulum is present. The

6 804 Z.H. BHATTI ET AL. A B Fig. 3. Hydspitherium f. megephlum, lower dentition. A, PUPC 95/24 ldm, B, PUPC 97/17, right mndiulr frgment with prtilly P3-4 nd omplete M1-3. Views re olusl (), lingul () nd lil (). Sle r equls 10 mm. postmetristid is overlpping with prentoristid. The entoonid is well developed nd spindleshped. It is pointed in the middle with two sloping ristids. The hypoonid is perfetly V-shped. The mesostylid is well developed s ompred to the met- nd entostylids (Fig. 3F). M3: The third molr well preserved with the hypoonulid (Fig. 3F). The ingulum is well developed on the nterior side of the tooth. The protoonid is well developed nd V-shped. The pex of the V is rod t the se eoming pointed nd nrrower towrds the distl end due to the middle stge of wer. The preprotoristid is nrrower thn the postprotoristid. The vlley enlosed y the posterior onids is ompressed trnsversely. The prehyporistid nd the posthyporistid re simple. The omprtive mesurements re provided in Tle I. COMPARISON Being squred nd tetr tuerulted teeth, it

7 HYDASPITHERIUM FROM THE MIDDLE SIWALIKS 805 Tle I.- Comprtive mesurements of the heek teeth of the Siwlik Hydspitherium. *The studied speimens. Referred dt re tken from Mtthew (1929) nd Colert (1935). Tx Numer Nture/Position Length Width W/L rtio H. f. megephlum PUPC 97/20* rp PUPC 91/23* rm PUPC 97/16* rm rm PUPC 67/155* lm PUPC 97/17* rm rm rm PUPC 95/24* ldm H. megephlum AMNH P M M BW 9 dm AMNH lm lm lm PUPC 95/24 lm H. grnde Ind. Mus. B135 M M Ind. Mus. B135 M H. mgnum GSI B514 M M PUPC 67/195 rm rm n e referred to some herivorous mmmlin group. Sine the ones/onids re resenti in outline, it n sfely e inluded in Ruminnti nd Tylopod (Romer, 1974; Zittel, 1925). Tylopod inludes Cmelide hving teeth with finely rugose enmel. In ddition, the Cmelide is not reported from the Siwlik lte Mioene (Mtthew, 1929; Colert, 1935). Ruminnti omprises four fmilies nmely, Trgulide, Cervide, Bovide nd Girffide. The trgulids nd ervids re smll nd medium size ruminnts (Gentry et l., 1999). The ovids re ommonly medium size ruminnts however some forms re lrge ut without rippled enmels (Pilgrim, 1937, 1939; Thoms, 1984). The girffids re lrge-sized ruminnts hving teeth with rippled enmel. The studied premolrs nd molrs with rippled enmel re lrge enough to inlude in girffids. They hve typil girffid s fetures, inluding lrge size nd finely rippled enmel (Mtthew, 1929; Colert, 1935; Gerds nd Asln, 2003; Kostopoulos nd Sr, 2005). The Siwlik girffids my e divided into two groups, the Middle Mioene Pleotrgine nd the Lte Mioene Sivtheriine (Colert, 1935; Srwr nd Akhtr, 1987). The Siwlik middle Mioene smll forms inlude the gener Girffokeryx nd Girff, while the lte Mioene lrge forms inlude the gener Brmtherium, Hydspitherium nd Sivtherium (Bhtti, 2005). The reovered teeth re lrge-sized s in Brmtherium, Hydspitherium nd Sivtherium (Colert, 1935; Gerds nd Gule, 1999). In ddition, hrters s the flt entoonid wll nd rhydonty present in Eursin pleotrgines, re sent in the studied smple. Aording to Floner nd Cutley (1861) Sivtherium differs from Hydspitherium hving less pointed loes nd wek prstyle. Brmtherium differs from Hydspitherium hving L-shped protoone (Khn nd Srwr, 2002). In the desried speimen the nterior fossette is

8 806 Z.H. BHATTI ET AL. H. f. megephlum H. megephlum H. grnde H. mgnum Width Upper First Molr Length Width Upper Seond Molr Length Width Lower Seond Molr Length Width Lower Third Molr Length Fig. 4. Bivrite plots showing dentl proportions of the Siwlik Hydspitherium. Referred dt re tken from Mtthew (1929) nd Colert (1935). ompressed trnsversely, the styles/stylids re present, the nterior medin ri is prominent nd the enmel sulpture is orse (Figs. 2-3). These hrteristis re found in the genus Hydspitherium (Pilgrim, 1911; Mtthew, 1929; Colert, 1935; Bhtti, 2005). The studied smple shows the mximum resemlne with Hydspitherium mong the known Siwlik Sivtheriine. The studied mteril s dimensions lose to speies H. megephlum (Fig. 4). The desried molrs re lerly smller thn those of H. mgnum (Tle I; fig. 4). The desried speimen nd the referred speimens of H. megephlum pper sme in size (Fig. 4) nd rown struture (Pilgrim, 1911; Colert, 1935; Bhtti, 2005). Dimensions of the teeth re very similr to those of the H. megephlum holotype nd lso to those of the lredy reovered speimens from the Siwliks (Mtthew, 1929; Colert 1935). The smll differene in size my e onsidered s intrspeifi vriility. Consequently, the mteril is ssigned to H. f. megephlum s the mteril is insuffiient for definitive speifi determintion. DISCUSSION AND CONCLUSIONS Lydekker (1876) desried the genus Hydspitherium, sed on olletion from the different lolities of the Siwliks. Further olletion of Hydspitherium hd een mde y Pilgrim (1910), Bohlin (1926) nd Mtthew (1929). These olletions omprise rnil nd postrnil elements. Four speies of Hydspitherium were desried in the Siwliks: H. megephlum Lydekker, 1876, H. grnde Lydekker (1878), H. mgnum Pilgrim, 1910 nd H. irmnium Pilgrim, 1910 (Lydekker, 1876; Pilgrim, 1910; Bohlin, 1926; Mtthew, 1929; Colert, 1935). The numer of these determined speies hs lredy een onsidered s exggerted, nevertheless the txonomy nd the vlidity of the girffid speies in the Siwliks hs not een fully resolved yet

9 HYDASPITHERIUM FROM THE MIDDLE SIWALIKS 807 (Mtthew, 1929; Gentry, 1997; Gerds nd Gule, 1999). The desription of H. irmnium Pilgrim, 1910 ws simply sed on single speimen, GSI B517, right upper molr (Mtthew, 1929; Colert, 1935). Hydspitherium grnde Lydekker (1878) nd H. mgnum Pilgrim, 1910 were only ereted to disply minor vritions in size (Mtthew, 1929; Colert, 1935). In our opinion the systemti position of H. irmnium Pilgrim, 1910 is doutful nd it my e n invlid speies. Similrly, H. grnde Lydekker (1878) hs the sme hrteristis s H. mgnum Pilgrim, 1910 nd oth of them re lrger in size (Fig. 4) thn H. megephlum Lydekker, 1876 (e.g., see Mtthew, 1929, p. 543). Morphologil nd metril dentl hrters (Tle I) of H. grnde Lydekker (1878) nd H. mgnum Pilgrim, 1910 indite tht the differenes mong the speimens do our, ut do not exeed ordinry intrspeifi vriility (Mtthew, 1929; Colert, 1935). The differene proly reflets sexul dimorphism, sujet to intrspeifi vriility in the lte Mioene Siwlik Sivtheriine. This is espeilly true if one onsiders the rnge of vrition oserved within speies of extint Eursin sivtheriines (e.g., see Gerds nd Gule, 1999; Gentry et l., 1999). Erlier uthors (Mtthew, 1929, p. 243; Colert, 1935) lredy proposed lose ffinities mong H. grnde Lydekker (1878), H. mgnum Pilgrim, 1910 nd H. irmnium Pilgrim, Moreover, there is no pprent reson to keep them seprte; the ommon strtigrphi origin of three speies from the Dhok Pthn Formtion, strengthens suh deision. At present, it is onvining to sy tht there re two speies of Hydspitherium in the Siwlik lte Mioene: smll speies H. megephlum nd lrge speies H. grnde. Following nomenlture rules, it is ovious tht H. grnde Lydekker (1878) hs priority over oth H. mgnum Pilgrim, 1910 nd H. irmnium Pilgrim, In our view, the generi ttriution of Hydspitherium is open. However, further work is needed, pending the reovery of more dignosti mteril of Hydspitherium from the suontinentl Siwliks. Hydspitherium is the most ommon girffid t Dhok Pthn nd Hsnot, nd the equivlent sites dted etween the lte Mioene nd erly Plioene (e.g. for iohronology see, Brry et l., 1982, 2002; Bdgley et l., 2005). The lte Mioene smple provides new evidene of the norml vriility of girffid teeth whih seems to onfirm the previous hypothesis of teeth vriility in the lte Mioene sivtheriines (Gerds, 1989; Gerds nd Asln, 2003; Khn, 2007). REFERENCES AKHTAR, M., Txonomy nd distriution of the Siwlik ovids. Ph.D. dissert. (unpulished), University of the Punj, Lhore, Pkistn. BADGLEY, C., NELSON, S., BARRY, J., BEHRENSMEYER, A. K. AND CERLING, T., Testing models of funl turnover with neogene mmmls from Pkistn. In: Interpreting the pst: Essys on humn, primte, nd mmml evolution in honor of Dvid Pilem (eds. D.E. Lieermn, R.J. Smith nd J. Kelley). Brill Ademi Pulishers, Boston. BARRY, J. C., LINDSAY, E. H. AND JACOBS, L. L., A iostrtigrphi zontion of the middle nd the upper Siwliks of the Potwr Plteu of northern Pkistn. Pleogeogr., Pleolimt., Pleoeol., 37: BARRY, J., MORGAN, M., FLYNN, L., PILBEAM, D., BEHRENSMEYER, A. K., RAZA, S., KHAN, I., BADGELY, C., HICKS, J. AND KELLEY, J., Funl nd environmentl hnge in the lte mioene Siwliks of Northern Pkistn. Pleoiology, 28: BERNOR, R.L. AND HUSSAIN, S. T., An ssessment of the systemtis phylogeneti nd iogeogrphi reltionships of Siwlik Hipprionine Horses. J. Vere. Pleontol., 5: BHATTI, Z. H., Txonomy, evolutionry history nd iogeogrphy of the Siwlik girffids. Ph.D. Dissert. (unpulished), University of the Punj, Lhore, Pkistn. BIBI, F., Origin, pleoeology, nd pleoiogeogrphy of erly Bovini. Pleogeogr., Pleolimt., Pleoeol., 248: BOHLIN, B., Die Fmilie Girffide. Pleont. Sini Pekin, C4: COLBERT, E. H., Siwlik mmmls in the Amerin Museum of Nturl History. Trns. Am. Phil. So., N. S., 26: FALCONER, H. AND CAUTLEY, P.T., Sivtherium gignteum, new fossil ruminnt genus from the vlley of the Mrknd in the Siwlik rnh of the Su- Himlyn Mountins. Asi. Res., 19:1-24. GENTRY, A. W., Fossil ruminnts (Mmmli) from the Mnong Vlley, Tnzni. In: Neogene pleontology

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