14
382 YR. R. I. POCOCK ON 29. The Affinities of the Antarctic Wolf (Cmis anfawticrts). Bp 12. I. I’OCOCK, l”.I<.s., F.L.S., l?.z.s.> ChlKltGl. of nIammals. [Received aud Read April 8, 1913.1 (Text-figures 70-74.) IXDEX. Page _._ 383 384 391 1. Published ricas on the afinities of C. nntarcticzhs 2. Skull clraracters of C. antarcticus aid C. latrans ... .,, ._. .. ... 3. Esteriial oliaracters of C. antarcticus and C. latrans ... ..,... The story of Cui& ui%turcticus has heen told by Darwin *, by Hamilton Smith j., and more recently by Mr. Bupert Vallentin from first-hand information, and by several authors indebted either to Darwin’s or Hamilton Smith’s account, or to the accounts of tmvellers who visited the Ealklanrls before Darwin’s time. Iteferences to the literature clown to 1890 may be found in Mivnrt’s Monograph of tlie Canitla?,’ published in that year. According to Mr. Ballentin, d%7& ct?~tcwctiez~s becxnie extinct in 1870, without leaving a trace of its former existence in the Fidkiantl Islands ; and since all the known material of the species appears to he preserved in London and Paris, I have atteniptetl to supply tlie want expressed by Allen 4 by figuring a skull of one of the specimens in tlie British Museim. I have not, however, given detailed ineasureinents of the skull, because these may be found in Mivart’s monograph and in the paper by Huxley mentioned below. Some six or seven yea.rs a.go, when trying to identify some South American dogs exhibited in the Zoological Gardens, I took tlie incideiital opportunity of looking n.t the skulls of a few of tIie species of Keotropical Cmida? contninecl in the Ihitish Museuin, to learn, if possible, something of their ;tfiiiitics to one nnotlier and to the better 1mow1~ species inhabiting North Anierica and tlie countries of the Old World. Amongst the species examined were Cc~,nZ’s untccrcticzis, the so-cdled Wolf uf the Falklaritl Islands, mil Cunis Zutrms, the Coyote or Prairie \volf, which ranges roughly from Cannda to Mexico. The exrniiriatiun was inade without any intention on my part of Rtlding to the literature of tlie subject, TT-ith which I was only acquainted in a very general \my ; aid after satisfying myself that C. a~~,tcwcticw was relat.ed to certaiii Neotropical forins, of which C. thous (= cnncriz*orzcs) may he t&en as an exainple, nncl that the afinities of C. Ictti-ccns lie with some of the so-called jackals and wolves of the Old World, I was contented to let the imtter rest. * In Watcrhonne’s Zonl. of 1I.M.S. Beagle,’ i\laninialia, p. 7, 1839. + Iu Jnrdine’s X&t. Libr., IZliilIiitialiii, is. 1’. 282. cliiioirs, sl\ iii. p. 45, 1904. Tlii.; ]xiper is qiintrrl liy smile nf tlir iiitwwtiiig and piixAiiig poiiit,s coiiiiected with iwmiril in “l‘lic Field,’ Oct. 1, 1904. Uiiir. Espcd. l’:itagoiii:i, iii. lit. 1, 1’. 153, 1905.

The Affinities of the Antarctic Wolf (Canis antarcticus)

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Page 1: The Affinities of the Antarctic Wolf (Canis antarcticus)

382 YR. R. I. POCOCK ON

29. The Affinities of the Antarctic Wolf ( C m i s anfawticrts). Bp 12. I. I’OCOCK, l”.I<.s., F.L.S., l?.z.s.> C h l K l t G l . of nIammals.

[Received aud Read April 8, 1913.1

(Text-figures 70-74.)

IXDEX. Page _._ 383

384 391

1. Published ricas on the afinities of C. nntarcticzhs 2. Skull clraracters of C. antarcticus a i d C. latrans . . . .,, ._ . .. ... 3. Esteriial oliaracters of C. antarcticus and C. latrans ... ..,...

The story of Cui& ui%turcticus has heen told by Darwin *, by Hamilton Smith j., and more recently by Mr. Bupert Vallentin from first-hand information, and by several authors indebted either to Darwin’s or Hamilton Smith’s account, or to the accounts of tmvellers who visited the Ealklanrls before Darwin’s time. Iteferences to the literature clown to 1890 may be found in Mivnrt’s ‘ Monograph of tlie Canitla?,’ published in that year. According t o Mr. Ballentin, d%7& ct?~tcwct iez~s becxnie extinct in 1870, without leaving a trace of its former existence in the Fidkiantl Islands ; and since all the known material of the species appears to he preserved in London and Paris, I have atteniptetl to supply tlie want expressed by Allen 4 by figuring a skull of one of the specimens in tlie British Museim. I have not, however, given detailed ineasureinents of the skull, because these may be found in Mivart’s monograph and in the paper by Huxley mentioned below.

Some six or seven yea.rs a.go, when trying to identify some South American dogs exhibited in the Zoological Gardens, I took tlie incideiital opportunity of looking n.t the skulls of a few of tIie species of Keotropical Cmida? contninecl in the Ihitish Museuin, to learn, if possible, something of their ;tfiiiitics to one nnotlier and to the better 1mow1~ species inhabiting North Anierica and tlie countries of the Old World. Amongst the species examined were Cc~,nZ’s untccrcticzis, the so-cdled Wolf uf the Falklaritl Islands, m i l Cunis Zutrms, the Coyote or Prairie \volf, which ranges roughly from Cannda to Mexico. The exrniiriatiun was inade without any intention on my part of Rtlding to the literature of tlie subject, TT-ith which I was only acquainted in a very general \my ; a i d after satisfying myself that C. a~~,tcwcticw was relat.ed to certaiii Neotropical forins, of which C. t h o u s (= cnncriz*orzcs) may he t&en as a n exainple, nncl that the afinities of C. Ictti-ccns lie with some of the so-called jackals and wolves of the Old World, I was contented to let the imtter rest.

* In Watcrhonne’s Zonl. of 1I.M.S. ‘ Beagle,’ i\laninialia, p. 7, 1839. + Iu Jnrdine’s X&t. Libr., IZliilIiitialiii, is. 1’. 282. cliiioirs, sl\ iii. p. 45, 1904. Tlii.; ]xiper is qiintrrl liy smile nf t l i r iiitwwtiiig and piixAiiig poiiit,s coiiiiected with iwmiril i n “l‘lic Field,’ Oct. 1 , 1904. Uiiir. Espcd. l’:itagoiii:i, iii. lit. 1, 1’. 153, 1905.

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TRE AKTARCTIC WOLF. 383

Rut in the summer of 1912, I received for review from the Editor of ‘ Nature ’ a copy of Dr. R. F. Scharffs volume, ‘ Distribution a n d Origin of Life in hinerica,’ 1911 ; and when I found it definitely st:tted therein that C. cciitai*cticics is closely related to G‘. Zatrans, and when I ‘saw the obvious difficulties in which Dr. Scharff was involved in his attempt to explain, on geo- graphical grounds, this singnlar :Itfilintioii, I ventured to reassine him by remarking, in effect, t1i:tt his belief W R S devoid of morpho- logical fonndation.

Now, an author who compiles a volume on zoology of the size arid scope of the ‘ Distribution and Origin of Life in America ’ cannot be expected to verify :dl the staternents of earlier and con- temporary writers. Nor in the present instance could Dr. Schnrff be justly criticisetl for not travelling to London to examine for liiaiself the preserved material of 6‘. antamticus, of which, i take it, there is no specimen in Dublin. Very naturally, therefore. he trusted to the verdict of others, and promptly replied to my remark with a request for my 1-easons for making it. Rut since J. coiild not ask the Editor of ‘ Sittnre ’ to give ine the necessary space for justifying the stn.ternent I had made, I pletlged myself t o do this elsewhere, and the matter that follows is an attempt to redeem that promise.

The acknowledged source of Dr. Scharff’s opinion about the niutunl affinities of C. cmtarcticm and C. latrcins was the followin= passage in Mr. Lydekker’s ‘ Geographicnl History of Mammal; 1896 :-“ Of the two indigenous mammals, the most remarkable is the Falklancl Island Wolf (C‘anis untnrcticz6s), wliich diff’ers ~narkedly from all the Canidze of tlre nrainland and is apparently closely allied to the North American Coyote (C. latrans) ” (p. 140). I therefore wrote and asked MY. Lydekker if he wonld kindly tell me his reasons for this conclusion, and he informed me that he took i t from Prof. Huxleg’s classic paper upon the critiiial arid dental chariicters of the C:initl=, published in the ‘ Proceedings’ of this Society, 1880, pp. 238-288. Upon looking rip this pn.per I find tlie following passages referring to tlre two species under discussion and bearing upon tlie question at issue :-

(1) . . . . . Rut sometimes there is a well-defiried though coin- pm.atively narrow s:tgittal area,, from t>lie centre of whicli a. lorn s:igitt:tl crc~st~ rises. This is well seen in some J:wkals, arid especially in C. rcibtc~rcticzcs (fi. 250).

(2) In the large size of the iipper niolars . . . . . C. anturctic.rcs presents the closest approximation to some specimens of C. Intraits (p. 266).

( 3 ) From the range of wriation of (7. crmcrizorzcs it can hardly be doubted that the examination of more extensive materials d l prore the existence of a.ti uninterrupted

: series of gi.ad:htions froin Cr. w t d w to C. ni~tarcticzcs and C. jubrctits (p. 266).

(4) Seven crania of C. I c i t ~ c c m , mhen measured, exhibit a con- siderable range of rariation, tlrough probably less than a larger series mould show. Rut, as they are, I mnst collfess iiiyself iuia1)lc t80 f i i i t l :nil inilmrbaiit I)reak in the

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384 MR. R. I. POCOCK ON

series of grwlntions of crmial and dental structure between C'aiiis l a t r m s ant1 C. aiatarctiocs on the one hand, and C. latrans and C'. occideiitnlis on the other. . . . . . I may further reiiiark that I can discern no difference of the slightest importance betuleen skulls of C'. b t m m and those of some of ow- domestic dogs (pp. 272-273).

(5) In the genus C'ccnis we have . . . . . as a lowest section the species of the G. cancr iaoi~~s and G. vetulus type (an- swering pretty much to the Agiia,i*m dogs of IIamiltoii Smith), the Hacaline section (C. uzcreics, C. a?ztJLus, C'. ~iaesontelas, C. antarcticits, C'. latrans), and the Lupine section (C. Zicpzcs and all its varieties) (p. 286).

Whether these pxigraphs justify Mi-. Lydekker's statement * that C. u)Lta.rctic?ts tliffers markedly froin all the Cnnic-lix of the inainlantl of South America, and is xpptrently closely allied to C'. Zati-ans, :tnd Dr. Schnrffs extension of this to the effect tliat C. antctrcticzcs is certainly closely related to C. latrcvns, must be left to intlividnii.1 judgment.

Pari\gr:*pll 1 merely points out one resemblance between C. antarcticus and some jaclmls. Pnragraph 2 similarly points out one resemblance between the two species, but conta,ins no suggestion of affinity between them. P:tragrnph 3 may be interpreted as suggesting affiliation between the extreme forms of South American Uanidze represented by C. eettc1u.s and G. jtcbatus, with C. ccntamticus lying midway between them. Paragraph 4 is more precise and stxtes that there is no im- portant strnctural break between C. avrtnrcticits and C. lntmns, and that the latter similarly intergrades witli 6'. occidantalis and C. fanzilictris. Parxgraph 5 , on the contrary, definitely associates G'. antctrctieus and 17. lutrcms, mcl ah the saiiie time severs the former from the group typified by C. .c;et.ccZits and the latter from the group typified by C. licpics 01- occitleiitalis, an arrangement not easy to recoiicile witli the views expressed by p:w:~gr:iphs 3 and 4.

Afte r reading Prof. Huxley's paper mther carefully for eriligliteninerit on this subject, I niust confess that I cmnot form any clear idea as t o his views of tlie d in i t ies of the species lie discussed, except in n broad sense.

If tlie srtbst;ince of paragrapl:s 4 ancl 5 aford some justifi- cation for Mi-. Lylelrker's tlecluation respecting t,he relationsliip between C. u~ztarcticl-ls ancl C. lutrnizs, it riinst, he admitted that p:iragraph 3 does not support the contention that C'. aiztnrcticus is quite iinrelntetl to the species of Ca,iiitl~ inhabiting the Sonth Anierican mainland. IIowerer that may be, the conclusions forced upon me by the exaiiiinn.tion of five craiiia of C. cmt- ccrcticics and twelve of (7. latrams 7 in the British Mnseum and

*; In the article iii 'Tlw Field ' (Oct. 1, 190&), nborc rrfrrrrd to, MY. Lydehker evinces lesa ahsuraiicc 011 these poiiits; Init lie c\.i(leiitlg corlld not bring liiinsclt to

t the anthority of Husl I use this term in i ts (11

Page 4: The Affinities of the Antarctic Wolf (Canis antarcticus)

THE AXTARCTIC WOLF.

Text-fig. 70.

385

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386 MR. R. I. POCOCK 011

the Museum of the College of Surgeons are:-(l) t,hnt C. ant- cwcticcss and C'. Iatmias are not closely allied ; (2) that 6'. alstu~-cttc~cs is niore nearly reLited to the C. thozrs (= cuncrivorus) p-onp of South American Ca,nid;e than to G. latrans; (3) that C'. lativ,iis niiist be affi1i:rted with such Old World species as C. pal l ips , C'. hpuster ancl C, unthzcs, and not with C. unta~cticus, Tlie first and third of these conclusions are borne out 11y the external characters of tlie two species concerned. My reasolis for these conclusions are as follows :-

l 'he suqittal urea uiad sayittnl cmst.-As Huxley and I\iTir?rt have shown, the skull of C. c~7atnrcticzcs has a well-iiiarked lyiifoi in sagittal area which, accortling to the evidence of available clanin, peisisted throiigliout life, :Llthougli in one of the three bpeciriiens in the British Museinn it is decidetlly narrower tlian in tlie t n o others. In the skulls of C. latrims that I have seen there is no tlihtinct lyriforin sagittal area, but iri adult skulls tlieie is a nietlian cariniforrn sagittal crest varying in Iieight with age. Even in two J oung slinlls, in both of which the slhenoitlal ant1 occipital sutures are open, while oiie still retains a, inilk canine behind the permanent canine, there is no lyriforin siiqittal wren. The significance of this depends upon the fact that the young of many species of Canirh of corresponding age or older show a stronger or weaker Iyrifoim aiea coriesponding Fvith the sinuosity of the upspreading temporal inuscle on each side, althongh in the young of no species of dog in which the adult possesses a caiir~i- form sagittal crest does the lyriform sagittal area &ow, I believe, the development arid definition it exhibits in the adult of G. atatn~cttcus. However that may be, if C. ccntcoctictrs ant1 C. Zatruias were closely related, we should nt least expect t o see a well-defined lyriforni sagittal area in the skulls of suhadult intlivitluals of C'. Zatrairs killed before the tempoial muscles had reached the siinmit of the claninin. But, as has been mid, this aiea is remarkable for its indistinctness in immature skulls of thnt species.

Thr occipitccl cre$t,- In C. cciitarcticics the occipital creqt, when viewed from above, is trans! ersely truncated and not angular ; when viewed fioni the side it only oveiha,ngs tlie vertical portion of the supi:wccipitcal to a small extent ; a n d uhen viewed fro111

Irehind it foriiis a tiiincatetl angle. In C. l a t ~ c w s this crest is angirlarly producecl h c k u ards in the middle line, or erhnngs tlte occipital area t o a much gi.eater extent, and is mole acutely angled from behind. It ~ a i i e b in shape and clepeloprnent in this species, hut neber, so f a r ac; I have seen, resembles that of

T%e malrcr bone.- In Cciicis roitccrct~ciis the anterior poi tion of the malar bone ih ni:iiked by a. stroug inasseteiic ridge trarerqrlg approximately the ni~dtlle of its outer suiface; the infelior edge of the bone close t o the maxilla is expanded corivexly t o aff;11(1 additional suppoi t t o the mnsseter inuscle ; its upper edqe close to tile ixaxilin is somewhat out-turnetl, foiniing R very :ipprec~ab]e lollo on on tlic snl)j:rcent poi tion of the niadlti :&o\ e the first

C. rr?Lt iWCtiCUS (text-fip. 70 & 71).

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s& AXYAWTIC WOLF. 387

molar tooth. I n C'. Zntrmzs the inxsseteric crest of t h e malar is low down on its external snrfxce, the inferior edge of t h e bone is scarcely at all exptnded, so that t h e area for the sttachmerit of

Trxt-fig. 71.

~ s. a

A. Occipital r q i o n of skull of Conis nnfavcticits. 13. Occipital region of skiill of C. latram.

s.a., sagittal area ; s.c., sagittal crest.

t h e masseter is niucli narrower than in C. antarcticzrs, and t h e upper edge of t h e malar is not noticeably out-turned, so tha t t h e hollow on t h e maxilla beneath i t is less pronounced (text-figs. 72 I% 73, pp. 388-9).

Q p e r cornassicd tooth.-In C. ccntccl'cticw t h e antem-external P R O C . ZOOL. SOC.-1913, NO. XXvII. 27

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388 MIL. I t . 1. POCOCK OY

cusp has the front border more rounded and the very fine crest that runs down it, is rlefiuerl on the inner side by a very indistinct groove. The nntero internal cusp is wider and rises further back and has no distinct little crest running inwards towards the

d h

bb w

il

0 * u1

.- c

E

antero-external cusp. 111 C. latimes the crest traversing the anterior edge of the antero-external cusp is more pronounced and is defined by a distinct groove, the two combining to make the edge of this cusp more cutting than in C. antarcticus. The

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‘rrm ~STAILCTIC WOLF. 389

antero-internal cusp is narrower and set distinctly more forwards than in C. ccnfarcticns, and there is a delicate crest running along its surface towards the base of the antero-external cusp (text- fig. 74, A, R, p. 390).

Lower cnrnassial tooth.-The main cusp is higher and more pointed in C. antarcticus than in 6. lcstrans, and the little crisp at its base on the inner side is inucti lower, so that it stands on a little higher level than the internal cusp of the talon. I n C. Zatraizs

M r-

this cusp is comparatively high up the main cusp of the tooth and is considerably above the inner cusp of the talon (text-fig. 74,

There are other minor differences both in the skull and teeth. The palatine bones, for instance, extend farther forwards with relation to the upper carnassials, and the margin of the posterior nares is also farther forwards with relation to the posterior molars in C. latrccns than in C. antarcticus : the incisor teeth are

c, n, p. 390).

37%

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3!)0 MK. n. I. POCOCK ON

Hmaller and the ci owns of the cheek-teeth are higher with relation t o their breadth in C. antarcticus than in C. lc&wns. But apart from these, the principal differences mentioned above are quite sufficient to disprove the claim that tlie two species are closely related. According to riiodern standai ds of classification they are subgenerically, if not generically, diitinct.

But tlie cliaracters above described tell us more than that. Taking C. lcctrairs first, it is obvious that in the cariniform sagittal crest, the anplar ly produced occipitnl crest, the position of the niasseteric ridge on the mdar bone, and in the points alluded to in connection with the upper and lower carna~sials, the species falls into line with the large wolves like C. occidentalis and ~ ? L ~ u s ,

Text-fig. 74.

C D A . Vertical view of upper caniassjnl of Canin Zutmnn. I$. Vertical riuw of upper cariiass1:11 of C. a7ctort.ticus. C. Iiitei~nal view of i o w n carnassinl of C. Zntmns. I). Iiiteriial view of lower earliassial of C. unturctictcs.

and 1vit.h C. pdlipes a.ntl C. lupaster, which, a.ccording to fancy, ma,y he cnlled large j:\cka.ls or small wolves. These resenihlances explain Mivnrt’s dismissal of the cranial and dental characters of C. dcctrnias with the remark, “ The skull possesses 110 distinctive clinrw.cters, nor have we been able 60 detect any in the shape of the teeth.”

On the other h m d , the skull of C. antccrcticus, with its lyriforni sa.gitta.1 a,rea and truncated occipital crest, agrees in tlie ma.in with the skulls of certain species or subspecies of Honth- American dogs in the British Museum labelled C. thozcs ( = cancrivortss), rztdis, sclateri (=nzicrotis), parvz‘dem, awostictzcs, gracilis, and fdv ipes . And in the skull of a dog, perhaps referable to C. gracilis, which

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THE ANTARCTIC WOLF. 31) 1

&me from Mar del Plata and died i n t h e Gardens, t h e above- (lascribed crests on tlie upper carnassial a re not better developed tllan i n C. antarcticLcs, and the masseteric ridge on t h e inalnr bone sllows a decided approximation to the condition seen in tha t species. This latter chemcter is st,ill better marked in t h e skull of another South-American dog, the exact locality of which is unknown, hiit which was a different species" from the Mar del Plata example, and the crests on tlie carnassinl exhibit the same feebleness of development. Biit it niay be noted that i n both these skulls tlie positions of the cusps on tlie upper and lower cama,ssials are more Zrctrcms-like than antarcticus-like, so tha t in this respect at. least they s e w e to bridge over the difference between t.liose two species ; a fact i n keeping with tlie idea tha t C. rc?~tarcticus is a specialised form of the group of South- Aniericm dogs above ;iI1ut~ed to, but specialised i n A direction away from tha t taken by C. Intrans ant1 its allies.

The external chamctevs of G. cintrtrcfiicics and C. Intrans also a.fford no justification for the claim of close rel;r.tionship between them. In t h e first place the ears of C'. cc?~tctrcl%czcs :we very sin:i,ll, srnxller indeed comp,a.vatively, I believe, t,Iian in any wild species of tlie dog family, with t h e exception perhaps of C. sclakri, C. (LVyctereutes) procyonoides, and Vtdpes ( A lopex) lngoprcs. I n G. latrans, on t h e contrary, they a re as large as in most, at all events, of the species of Canis. An idea of their length i n t h e two species may be gathered from t h e measurenient of a specimen of each of approxiinately the wine size given by Mivai*t, tlie ear of G. an,tccrctic~i~s being 6.5 cni. (= about 2: inches) and tha t of C. Iatr(ms 14 cni. (=xhout 54 inches), or inore than twice as long. It ma,y be added tha.t the measurement of 2 inches 9 lines assigned by Weterhouse t o the ear of C. antarcticus confirms Mivnrt's statement.

As regards colour C. latrans varies from grey to greyish fawn, mixed with black >hove, and shows t h e chamct.eristic cloutled or pntc.l\y colora.tion caused by t h e running togrt,her of the bands of t h e individu:d long coarse lixirs of t h e back antl sides seen in so many of the so-c:tlled wolves and jackals. One of these long coa.rse hairs, pulled a t random from a skin, measured a.boiit 3 inches long, t h e bhck t ip being 1 inch ( I S rnni.) a n t 1 t h e wliitisli area. below it 1 inch (25 nnn.). The wliole of the ventral surface from the'chin t o t h e root of t h e tail is nsually white or wliitisli, antl always apparently nia.rkedly paler t1iw.n the ]pack atid sides, thoiigh sometimes the continnity of t h e light t in t is inter- rupted on the throat by an infusion of fa.wn. There is no dark p i tch above the hock on t,lie liintl leg, and the tail matches the hack approxinintely in colonr throughout, t h e t ip and the gland- spot 1)eing hlaclter than t h e rest,.

I n C. aiztcircticts the coat is thick and soft, and comparatively short, with none of the long coarse hair seen in C. Zutrans. One

* The Sontli-Amel.iaaii clogs of this gronp are i l l snch a systematic mnddle t h a t it is very ditticult to identify sprrimPns without rl coinplete revisiou of the whole series.

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392 OS THE ANTARCTIC WOLF.

of the longish hairs pulled at random from t h e back mensured only 13 inches (37 mm.) in length, t h e dark apical t i p being 4 inch (6 mm.) and t h e pale band below it inch (3 mm.) in length. The prevailing colour of the body is brown, relieved by t h e fine speckling due to t h e narrow pale band on t h e individuad hairs just described. The loaer side is white onIy on the posterior portion of t h e beliy and on t h e upper end of t h e throat,, t8he chin and lower jaw being white stained with a fuscoiis tint. Apart from t h e areas described t h e ventral surface is brownish. There is, moreover, as Miva.rt said, R fuscous patch a.bove t h e hock of t h e hind leg and t h e tail is particolonred, its b a d portion being like tlie back, its t ip white, and t h e intermerlia~te area blackish, t h e colour of this area gradually blending p r ~ ~ i n ~ i t l l y with the brownish basal portion, bu t being quite black distally and sliarply defined from the white terminal area..

Perhaps it may rightly be claimed t h a t these differences in the length of t h e ears mid in colour do not count for much in themselves. Tha.t may be so. Kevertheless, if C. asitcircticus anrl C. latrans were only known from their skins, it is quite certain that t h e latter woulrl be phcecl in the same a t e g o r y with such species as C. pallipes and C. lupaster, ancl that C. antarcticzts would be excluded therefrom. The latter would be difficult to classify ; but there is one significant colour-feature connected with t h e species. This is tlie p~ rice of the Clark patch above the hocks ; and the interest of tl ies in t h e circumshnce that it is a very common fen.ture i n va.rious species of t h e smaller South-American dogs and occurs in some of t h e species of Ydpes, like F7. chasm *.

There is one other little point tha t may be referi.ed to. Darwin s a ~ s he was informed t h a t the cries of C. cc~~tarcticus resembled t,liose of t h e Sontli-American species C. C ~ Z U T ~ . I ha.ve never heard C'. azrrrce bark or bowl, hut t h e keeper in the Gardens informs me tha t examples of wild dogs from hlar del P la t s and Cordova, which m e closely allied to and perhaps only ra.cially distinct from C. m u m , bark after the manner of foxes. On t h e it-hole, however, they are silent dogs in captivity, ancl, like the foxes, never succumb t o the ternptiitioii of joining in the Iiowling concerts in which the dinpos, jackds, prailie wolves, and large wolves i n t,he Gardens indnlge, and which they seem unable to resist contributing to. Personally I heliere t h a t voice i n mammals is often a good guide t o affinity; and, in the present case, the voices of C. antamt icus and C. lcitwciis bear out my opinion of the relationship of these species t o otliers, sliown by structural characters.

Finally, if t h e conclusions a.hox-e put forwai,rl m e correct,, Hnxley's classificat,icn, expressed in p ~ r . 5 (p. 384), mnst be emended by transferring c. rk?itarcticm~s to the lowest section of

* The presence of this patch in snnie of the primitive Cunii lz is well worth more sttention thaii it has rwuived. I do not know what it may mean, any more thaii I kiiow wh:it the pile area hehind the shoulder, observable in ~iiaiiy Caiiida, biitli wild aud doniestic;itcd, may mean.

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ON’ AVIAF OSTEOLOOY. 393

dogs, containing C. thous (=canm*uorzcss) and vetulus, and by uniting his S a l i n e and Lupine sections. Possibly these sections contain groups worthy of subgeneric, if not of generic, recog- nition: C. antarcticus and C. thow, for example, may be sub- generically or generically separated by the structure of the mandible; but I do not see how C. latrmw is to be distinguished other than specifically from such forms as C. paU$es, anztihm, or even lupus.

30. On the Patella i n the Plielacrocorncidz. By Ur. R. W. SHUFELDT, C.M.Z.S.

[Received April 14, 1913: Read May 20, 1013.1

(Plate LXI.”) , While recently employed in preparing a detailed amount of the

skeleton of Harris’s flightless Cormorant (Nalznopterurn harrisi), in which the osteology of that species is compttred with that of a number of others of the family, I became interested in the morphology of the patella? of tliose birds.

Many years ago I published a number of papers t on the skeleton in the Cormorant, in some of which the pa.tella of the Pha,la- crocoracidE was referred to and figures given of it. But my inaterial, at the time to which I refer, was very limited-in fact I think there u-ere but tlie skeletons of one or two species of those birds a t hand, and this included the skeletons of two or three young ones.

I n t h e higber groups of birds the patella, when present, is usually small, and offers but little of value to tlie avian taxonomist. This, however, is not tbe case when we come to examine into the osteology of. some of tlie groups occupying lower positions in the system, and especially is this true of many of the Pygopdine forms and their allies near and remote.

Owen paid but scant attention to this bone of the skeleton in Aves, devoting less than half a paragraph to it, thus :-‘‘ The chief of the sesamoid bones in the hind limb is the patella : it is of unusual size in the Penguin, is ossified from two centres, and articulates with the procnemial process of the tibia : it coexists with the long i-otular process in the Loon, fig. 34, I ; it is large a.nd of an angu1a.r form in the Musk-cluck (Bizizcra): in the Merganser the pa,tella is largest and deeply notched j in the Coot i t is elongate. I n most aerial birds a patella is want,ing” 2. There is no reference made here either to a Grebe or n Cormorant, and

* For explanntioii of the Plat,e see p. 402. + Sliufeldt, R. W. “Osteology of the Cormorant,” Scieiice, Doc. 7, 1883, p. 739;

(‘ Concerning someof the forms asmmed 1,y tlie Putella in Birds,” Proc. U.S. Nat.

“ Osteolop of the Rtrgaiiopodes,” Mrm. (‘arnrgie Museum, Pitt Hates i i i i d marry trxt-figure.

Comp. Anat. and I’hys. of \‘ertel)rates, vol. ii. p. 83, Lo11do11

Frh. 8, 1884. vol. iii. No. 63, p. 143; ifiid. Apr. 18, 1884, h’o. 68, pp. 474, 475.

Mns. 1884. sii. pp. 324-331.

1903, vof. i. h’o. 3, Art. 3, pp. 16-70.

Nnineiocis text-figures.

Owen, Iticliard. iati6.