UNIVERSITY OF AMSTERDAM. The Netherlands. Abstract. to the Cervoidea Simpson, singly. as distinctive of a Cervine from a Bovine. animal".

Transcription

1 Beauforta STTUTE OF TAXOOMC ZOOLOGY (ZOOLOGCAL MUSEUM) UVERSTY OF AMSTERDAM Vol. 0 no. 7 December 4, 980 The confguraton of the lacrmal orfces n Pecorans and Tragulds (Artodactyla, Mammala) and ts sgnfcance for the dstncton between Bovdae and Cervdae J.J.M. Lenders & E. Hentz * nsttuut voor Aardwetenschappen, Budapestlaan 4, postbus 80.0, 508 TA Utrecht, The etherlands ** Museum atonaled'hstore aturelle, 8 rue de Buff on, Pars, France Abstract The number and poston of the orfces to the lacrmal duct n the lacrmal bone of rumnants s evaluated. t s concluded that the Bovdae, except the Bovn and Tragelaphn, have mantaned the orgnal one-orfce-nsde-the-orbt confguraton, whereas the two-orfces-on-the-rm-of-the-orbt morphology of the Cervdae was acheved before the M. Mocene n the evoluton of ths famly. The classfcaton of the Antlocaprdae s dscussed and t s concluded that the confguraton of the lacrmal orfces corroborates wth the suggested (Lenders, 979) transfer of the Antlocaprdae Gray, 866 from the Bovodea Smpson, 9 to the Cervodea Smpson, 9. TRODUCTO sngly as dstnctve of a Cervne from a Bovne anmal". n general there s no dffculty n separatng recent Cervdae and Bovdae by the character of ther frontal appendages: cervds beng typfed by the bony, decduous outgrowths from cylndrcal The dstncton between fossl bovds and cervds s even more delcate due to the great dversty, the paucty processes of the frontals, bovds by the persstent horn sheaths coverng bony cores on the frontals. of materal and the restrcted nformaton on the anatomy and habtat of fossl pecorans when compared wth the recent ones. However, the exstence of pecorans n whch the Hentz (970) frontal appendages are wantng, such as the Chnese waterdeer (Hydropotes nerms) and the descrbed the dfferences n dentton and post-cranal skeleton between the Vllafranchan Bovdae and Cervdae of western muskdeer (Moschus moschferus), and of peco- Europe. Some of the lsted dfferences have a rans wth anomalous frontal structures such as the pronghorn (Antlocapra amercana), made many general applcablty, especally that n the morphology of the gully zoologsts search for other dstnctve features of at the anteror sde of the metatarsus (Hentz, 96; Lenders, 979). the Cervdae and the Bovdae (Flower, 875; van One of the features to dstngush between bo- Kampen, O'Gara and 975; Matson, 975). Brooke vds and cervds mentoned by Flower (875) and Brooke (878) s the number and poston of the (878, p. 885) lsted characters whch dstngush the Cervdae from the Bovdae, but he orfces to the lacrmal duct n the lacrmal bone: stated: "none of the above characters can be taken cervds havng two orfces stuated on the rm of 55

2 the orbt (Plate, fg. ), whereas bovds and Cervdae Gray, 8 tragulds have only one, nsde the orbt (Plate, fg. and Plate, fg. ). Both authors emphaszed the value of ths character, although they reported several exceptons: Tragelaphus and Antlocapra have two orfces as n cervds, whereas n Moschus the same s found as that confguraton Alces alces 4 Axs axs Blastoceros campestrs Capreolus capreolus 89 Cervus elaphus Cervus rusa Cervus nppon Hyelaphus kuhl Hyelaphusporcnus 7 Mazama amercana 9 Masama gouazoubra Muntacus sp. Muntacus muntjak Muntacus rever generally present n bovds and tragulds. The lacrmal ducts (ductus naso-lacrmals) connect the orbta wth the nasal cavty for the dranng of the surplus of lacrmal flud from the eyes Dama dama Elaphodus cephalophus Elaphurus davdanus Hydropotes nerms Odocoleus vrgnanus Rangfer tarandus 4 Rucervus eld to the nostrls. n cervds the lacrmal duct opens nto two or- The purpose of ths paper s to evaluate the fces, one above the other, on the rm of the orbt. number and poston of the lacrmal orfces for Although the poston of the orfces s slghtly the dstncton between Bovdae and Cervdae. varable (from slghtly on the nternal sde of the MATERAL rm to slghtly on the external sde of the rm), the lower orfce has a more anteror poston than the upper one (Plate, fg. ). n two of the The materal studed s stored n: Museum atonale d'hstore aturelle, Pars; nsttut Royal de skulls of Muntacus muntjak both orfces are Scences aturelles, Bruxelles; Rjksmuseum van stuated just nsde the orbt. One of the skulls of atuurljke Hstore, Leden; Zoologsch Museum Hyelaphus porcnus (Mus. d'hst. at. Pars no. 95-4) shows a very small upper orfce, whch Unverstet van Amsterdam; Taxonomsch nsttuut Rjksunverstet Utrecht; nsttuut voor confguraton s Aardwetenschappen Rjksunverstet Utrecht. consdered anomalous. Two anomalous specmens were found n the seres of 89 skulls of Capreolus = number of skulls examned. stored n the Taxonomcal nsttute n Utrecht. The rght orbt of skull no. MORPHOLOGY OF THE LACRMAL ORFCES RECET has only one orfce on ts rm, whereas the left TRAGULDS AD PECORAS. Traguldae Mlne Edwards, 864 orbt shows the normal cervd two. A smlar phenomenon was observed on skull no. 77 of whch Tragulussp. Tragulus javancus Tragulus memnna Tragulus napu Hyemoschus aquatcus the left orbt has only one orfce and the rght s normal. All tragulds examned have one elongated orfce just nsde the rm of the orbt (Plate, fg. ). Antlocaprdae Gray, 866 Antlocapra amercana 6 Moschdae Gray, 8 Moschus moschferus 5 n Moschus one elongated orfce s present just nsde the orbt, whch resembles that of tragulds. At the medal borderof the orfce there s a small protuberance, however, whch gves the mpresson The lacrmal duct of the sngle recent speces of the antlocaprds opens nto two orfces. They dffer from those n the cervds by the poston of the upper orfce, whch s stuated nsde the rm of the orbt. The lower orfce s placed, as n cervds, more anterorly, near the of the rm. margn that the sngle orfce n Moschus s ether the re- O'Gara and Matson (975) reported that Ant- sult of fuson of two separated orfces or that the locapra has only one orfce as the Bovdae, but orfce n Moschus corresponds ths s nether confrmed by the lterature (Flower, to an early (ntal) stage of transformaton from one nto two, whch s the normal confguraton n cervds (Plate, fg. ). 875; Frechkop, nor 955) by our observatons. For further dscusson see the chapter on the classfcaton. 56

3 Graffdae Gray, 8 ths general rule, however. There s consderable varaton, especally n the Tragelaphn. On the Okapa johnston Graffa camelopardals 8 7 examned skulls of Tragelaphus scrptus we observed the followng confguratons: The Okapa skulls show one very small orfce Two orfces on the rm separated by a very thn nsde the orbt. Of the eght skulls of Graffa bone bar. fve have no lacrmal orfce at all, the other three an orfce smlar to that of Okapa. Two orfces on the rm about 7 mm. from each other. Bovdae Gray, 8 Two orfces on the rm wth the upper more anterorly than the lower one. One orfce on the rm. Alcelaphus major Antdorcas marsupals Anllope cervcapra 5 Madoquasaltana aemorhedus sp. aemorhedus goral One orfce nsde the orbt. Two orfces nsde the orbt. Ammotragus lerva eotragus pygmeus Two orfces nsde the orbt; a large and a small Budorcas taxcolor esotragus sp. one. Capra aegarus Capra caucasca Capra falconer Oreamnos amercanus Oureba oureba Oreotragus oreotragus 4 7 Two orfces nsde the left orbt, one orfce nsde the rght orbt of the same skull. Capra hrcus Capra sberca Caprcorns sumatrenss Oryx sp. Oryx tao Oryx besa 5 Smlar varaton s found n the skulls of the other Tragelaphn. Cephalophussp. Ovbos moschatus On the skulls of the Bovn the followng confgu- Cephalophus dorsals Cephalophusturens Cephalophusmaxwell Connochaetes gnu 4 Ovs ares Ovs musmon Pantholops hodgson Pelea capreolus 9 ratons have been observed: Two orfces on the rm (Bson bson, Bson bonasus, one specmen of Bos banteng). Damalscus albfrons Damalscus korrgum Gazella dorcas 4 Procapra gutturosa Pseudos nayaur Raphcerus campestrs Two orfces nsde the orbt (one skull of Bubalus bubalus). Gazella ruffrons Raphcerus melanots One orfce on the rm of the orbt (Bos ndcus, Gazella soemmerng 7 Raphcerus sharpe two skulls of Bos taurus). Gazella dama Gazella grant Gazella thomson Gazella subgutturosa 0 4 Redunca redunca Rupcapra rupcapra 4 Saga tartarca 6 Sylvcapra grmma One orfce nsde the orbt (all other skulls). REMARKS O THE CLASSFCATO Hemtragus:sp. Hppotragusequnus 5 OF PECORAS AD TRAGULDS Hppotragusnger Kobus kob 8 Boselaphn Smpson, 945 Boselaphus tragocamelus Tetracerus quadrcorns Accordng to Flower (875, p. 86) the presence of one lacrmal orfce nsde the orbt s the Tragelaphn Sokolov, 95 Boocerus eurycerus Bovn Smpson, 945 orgnal confguraton n artodactyls, present n early forms lke Caenotherum and Xphodon. t s Taurotragus sp. Taurotragus oryx Tragelaphussp. Anoa sp. Anoa depresscorns Bson bonasus 6 generally accepted that bovds and cervds have a tragulod ancestry. Tragulds and bovds (except Tragelaphusgratus Bson bson the Bovn and Tragelaphn) stll have one orfce Tragelaphus mberbs Bos banteng nsde the orbt, whereas the two-orfces-on-the- Tragelaphusscrptus 7 Bos ndcus Tragelaphusspek Bos gaurus rm confguraton n cervds may be nterpreted as Tragelaphusstrepcceros 4 Bos taurus 7 a new development durng the early evoluton of Ltocranus waller Bubalus bubalus 5 ths famly. Madoqua sp. Syncerus caffer T Gnsburg and Hentz (966) proposed a new famly (Dremotherdae) for the paleogene antler- n most bovds one crcular orfce to the lacr- less cervods, whch they consdered ancestral to mal duct s found, stuated nsde the orbt (Plate, fg. ). n the trbes Bovn and Tragelaphn the morphology of the orfces s not accordng to the Cervdae and Graffdae. Accordng to Smpson (945) Prodremotherum s a tragulod. However, Jehenne (977) showed that Prodremothe- 57

4 rum s very close to Dremotherum and mght be vdae before the Mddle Mocene. One may also conclude that the closed gully on the metatarsus consdered the ancestor of the latter genus. Dremotherum has the orgnal one-orfce-nsde-theorbt confguraton (Sgogneau, 968). was acheved earler n the cervd evoluton than the two lacrmal orfces on the rm of the orbt. Hydropotes, on the other hand, ncontestably The oldest Cervdae of whch we have nformaton on the lacrmal orfces s Heteroprox from belongs to the Cervdae (Garrod, 877; Forbes, 88), whch s n agreement wth the two orfces the Aragonan (M. Mocene) of Stenhem (Germany). Although ts antlers show no well devel- on the rm of ther orbt (Plate, fg. ). oped pedcle, they already mght have been dec- The Antlocaprdae are generally consdered to duous (Gnsburg & Crouzel, 976). Heteroprox be related to the Bovdae whch fnds ts expresson n the classfcaton of both famles n one has two orfces on the rm of the orbt (Hezmann, pers. comm.), ndcatng that ths character superfamly: the Bovodea. Thenus (969) and s present n the Cervdae at least from M. Mo- Lenders (979), however, suggested that the ant- cene onward. locaprds are an offshoot from early amercan cer- t s generally accepted that Moschus and Hy- vods and for ths reason Lenders (979) pro- dropotes, although not closely related to each other, posed are the most prmtve genera among the recent cervods. n both genera antlers are wantng, whch to shft the Antlocaprdae from the Bovodea to the Cervodea. The presence n the pronghorn (Antlocapra amercana) of two orfces to s compensated by large sabre-lke upper cannes. These two genera Dremotherum (Sgogneau, 968). show certan resemblances wth the lacrmal duct, although postoned more nternally than n cervds (Plate, fg. ), supports the dea of a cervod ancestry of the Antlocaprdae. Moschus shows a mxture of bovd and cervd On the other hand, O'Gara & Matson (975) characterstcs (Flower, 875; Sgogneau, 968), and accordng to Thenus ( 969) ths genus may concluded from the smlarty n the horn-formng process that Antlocapra s very close to the Bov- be a remnant of a pre-pecoran stage of the ru- dae and they proposed to lower the rank of the mnant phylogeny, whch justfes ts classfcaton pronghorn and ts fossl relatves from the sepa- n a separate famly. The sngle lacrmal orfce rate famly (Antlocaprdae) to a subfamly (Antlocaprnae) nsde the orbt of Moschus separates the muskdeer from the Cervdae and corroborates wth ths of the Bovdae. The authors remarked: "Hstologcally horn formaton resembled that suggeston. However, the metatarsus of Moschus shows a closed gully at ts anteror sde (Flerov, of skn" and "the horn-formng process n pronghorns nvolves formaton of a keratn smlar to 9 Plate V) just as t does n the cervds. that of nals and claws". Combnng these state- Hentz (96) and Lenders (979) Emphaszed ments wth the fact that horn formaton s com- the value of the morphology of the gully at the mon n mammals, one mght wonder f horn form- anteror sde of the metatarsus as a dstnctve aton s a sgnfcant character for classfcaton. character between Bovdae and Cervdae. n bovds n addton to the smlarty n horn formaton, the gully s open over the full length O'Gara and Matson (975 : 840) lsted about of the metatarsus, whereas n cervds t closes above the dstal end of the shaft. n Moschus these two famlal characterstcs are contradctary (cervd-type meta- twenty features n whch the antlocaprds resemble the Bovdae. However, most of these are also found n cervds (e.g. cannon bone, loss of tarsus and bovd-type lacrmal orfces). As men- upper ncsors) or, they are varable n bovds toned earler n ths chapter, the lacrmal duct of cervds opens nto two orfces on the rm of the and/or cervds (e.g. orbts large to medum, contact between lacrmal and nasal bones). Other orbt snce the Mddle Mocene. Also the dfference n morphology of the gully on the metatarsus between bovds and cervds was already present at that tme (Hentz, T96). Ths ndcates that Moschus ancestral lne branched off from the Cer- smlartes lsted are: a) presence of a gall-bladder, b) one orfce to the lacrmal duct, c) presence of unbranched bony cores covered wth keratn, d) hypsodont dentton. Ad a) the of presence a gall-bladder n the 58

5 pronghorn ndeed makes t approach the bovds. But the presence of a gall-bladder n Moschus as the Glrdae, Murdae, Crcetdae, Equdae, Sudae and Cameldae. Dfferences n hypsodonty (Flower, 875) may ndcate that n prmtve cer- may be usuable for genus or speces classfcaton vods ths bladder was stll present and ts occur- wthn phylogenetc lneages, but certanly not on the level of famles or superfamles. Accordng to Frechkop (955) the pens ence n Antlocapra can be explaned as the retenton of an orgnal character. Flower (875) reports also that the presence of a gall-bladder of Antlocapra resembles that of cervds by the absence "seems to be a somewhat varable character even wthn the same speces". of a processus urethrae and Cowper's gland (Glandulae bulbourethrales). Ad b) accordng to O'Gara & Matson Antlo- The Tagelaphn and the Bovn are the only capra has only one lacrmal orfce nsde the orbt pecorans wth a varable confguraton of the or- as n bovds. Accordng to our observatons and fces to the lacrmal duct. Ths smlarty fts n the lterature (Flower, 875; Frechkop, 955) the wth the opnon (e.g. Thenus, 969) that these pronghorn has two orfces to the lacrmal duct. two trbes are closely related. The thrd and most Frechkop (955 : 66), however, reports that prmtve trbe of the bovnae, the Boselaphn, sometmes the two lacrmal orfces are fused nto have mantanedthe orgnal one-orfce-nsde-theorbt confguraton. one, whch could explan the contradctng observatons. Ad c) the horns of Antlocapra dffer from ACKOWLEDGEMETS those of the bovds by the forked form and by the annual sheddng of the horny cover. These dfferences were the man reasons to place the pronghorn n a separate famly. O'Gara & Matson ponted also occurs out that exfolaton of the horn covers n many bovds and from ths they concluded that the sheddng reason for separatng the pronghorn from the Bovdae. One can dspute n Antlocapra s no wheter these dfferences are sgnfcant for classfcaton on famly level or not, but the sheddng n bovds dffers largely from that n Antlocapra. Tn the pronghorn the sheddng occurs annually (n ovember) and s related to the rut. t takes about ten months to grow a new horn-cover (O'Gara & Matson, 975: fg- )' Ths parallels very much the growth and The authors are ndebted to C. W. Drooger, A. J. van der Meulen, P. Y. Sondaar and J. de Vos for readng the manuscrpt and dscussng the subject; to P. J. H. van Bree (Taxonomsch nsttuut Unverstet van Amsterdam) for readng the manuscrpt and permsson to museum study materal; to F. Popln and G. Petter (Museum at. d'hstore aturell Pars), C. Smeenk (Rjksmusum voor atuurljke Hstore Leden), D. van der Tooren (Taxonomsch nsttuut Rjksunverstet Utrecht) for permsson to study museum materal and to J. Luteyn for preparng the plates. The research of the frst author s fnancally supported by the etherlands Organzaton for the Advancement of Pure Research (Z.W.O.). castng of antlers n cervds and probably has the same functon n relaton to the rut. On the other hand the reported exfolaton of the horn-covers n bovds s rregular and ncdental and seems to REFERECES BROOKE, V., 878. On the classfcaton of the Cervdae, wth a synopss of the exstng speces. Proc. zool. Soc. London, 878: be related to the growth and/or renewng of the horn-covers. Ad d) The hypsodont dentton of the antlocaprds s frequently used as an argument to nclude ths famly n the Bovodea (Frechkop, 955; FLEROV, C., On the 90. classfcaton and the geographcal dstrbuton of the genus Moschus (Mammala, Cervdae). Ann. Mus. zool. Acad. Sc. URSS, : -0. FLOWER, W. H., 875. On the structure and affntes of the Musk-Deer ( Moschus moschferus Lnn.). Proc. O'Gara & Matson, 975). As can be concluded zool. Soc. London, 875 : FORBES, W. A., 88. Supplementary notes on the anatomy from the fossl record, hypsodonty s an adaptve attanment n mammals, whch occurs together wth brachydonty n many unrelated famles, such of the Chnese Water-Deer (Hydropotes nerms). Proc. zool. Soc. London, 88: FRECHKOP, S., 955. Rumnanta. Trate de Zoologe,

6 O'GARA, B. W. & G. MATSO, 975. Growth and Castng Prodremotherum Flhol, 877 (rumnant prmtf de of horns by pronghorns and exfolaton of horns by bovds. J. of Mammalogy, 56 (4) : GARROD, A. H., 877. otes the on of the Chnese anatomy Water-Deer ( Hydropotes nerms). Proc. zool. Soc. London, 877 : GSBURG, L. & F. CROUZEL, 976. Contrbuton a la connassance d'ḣeteroprox lartet (Flhol) Cervde du Mocene Bull. Mus. naton. Hst. europeen. at., Pars, Sc. Terre, 58 (99) : GSBURG, L. & E. HETZ, 966. Sur les affntes du genre Paleomeryx (rumnant du Mocene europeen). C. R. Acad. Sc. Pars, 6: Les HETZ, E., 96. caracteres dstnctfs entre metatarses de Cervdae et Bovdae actuels et fossles. Mammala, 7 () : Les cervdes, 970. vllafranchens de France et d'espagne. Mem. Mus. naton. Hst. at., Pars, (. S.), (C), : -0. JEHEE, Y., 977. Descrpton du premer crane du genre l'olgocene eurosatque). Geobos, Mem. specal : -7- KAMPE VA, P.., 905. De Tympanalgegend des Saugeterschadels. Morpholog. Jahrbuch, Lepzg, 4: LEDERS, J. J. M., 979. On the osteology and functon of the dgts of rumnants some and ther bearng on Z. Saugeterk., 44 (5) : taxonomy. SGOGEAU, D., 968. Le genre Dremotherum (Cervodea), anatome du crane, denture et moulage endocranen. Ann. Paleont. (vertebres), 54 () : -64. SMPSO, G. G., 945. The prncples of classfcaton and a classfcaton of mammals. Bull. Amer. Mus. nat. Hstory, ew-york, 85 : -50. THEUS, E., 969. Phylogene der Mammala: -7 (Walter Gruyter and Co., Berln). Receved: February, 980. nsttute of Taxonomc Zoology (Zoologcal Museum), Unversty of Amsterdam, P.O. Box 05, 000 HC Amsterdam, the etherlands 60

7 Plate Fg.. Tragulus napu R.M..H. Leden, a lacrmal orfce. Fg. Moschus moschferus R.M..H. Leden. Fg. =.. Hydropotes nerms no R.M..H. Leden. 6/

8 Plate Fg.. Antlocapra amercana no. 90 Z.M.A. Amsterdam. Fg.. Gazella ruffrons no M..H.. Pars. Fg.. Tragelaphusspek no M..H.. Pars. 6/