DET NORSKE VIDENSKAPS
-AKADEMI I OSLO
RE S·U LTATER
AV DE NORSKE STATSUNDERST0TTEDE SPITSBERGENEKSPEDITIONER
BIND I
Nr. 4
ALF WOLLEBÆK:
THE SPITSBERGEN REINDEER
(RANG/FER TARANDUS SPETSBERGENSIS)
UT GITT PÅ
DEN NO RSKE STATS BEKOSTNING VED SPITSBER GEN KOMITEEN
REDAKTØR: ADOLF HOEL
OSLO
I K O MM ISJON HOS JACOB DYBWAD 1926
Nr. 4
THE SPITSBERGEN REINDEER
(RA NGlFER TA RA NDUS SPE TSBERGENSlS) (ANDERSEN 1862)
BY
A LF W O LLEBÆK
WlTH 16 FIGURES IN TEXT, 6 PLATES AND I MAP
SPITSBERGEN
,,
'l=t----
'
---
Scale
~I :,.'(i:· j_;f;r'
25 30"
80
\
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Contents.
Page
Craniological Investigations .
. . . . . . . . . . . . 5Introduction, with Explanation of Measurements ... . . . 5
Remarks on the Material Examined... 13
Cranial Length . . . 15 Palatal Length... 16
Total Cranial Breadth, Breadth of Forehead, Length and Breadth of Nasal Section. . . .. 16
Condyli occipitales... . . 18
Breadth of Cranium at the External Auditory Meatus.. . . .. 18
Nasal Aperture . . . .. 18 Distance between Cheek-Teeth Rows . . . 22
NasaJia ...
.... ... ... .... ... . ... 24
Os supramaxiJlare accessorium. . . 30
Ossa parietalia . . . .. 32
Cheek-Teeth in MaxilIae
.. . . .. 33
Distance between the jugals, Processus styliformes tympani, and Processus paramastoidei. .
.. . .
.. . .
.. . .
. . . ..
. .. . .
. .. . .
... 36
MaxiJlae ..
... .... .... . ... ... . ... ... .... . . . ... . ... . . .. ... 40Os zygomaticum . . . .. ... 41
Distance between Premolars and Canines . . .
.. . .
.. . . .
.. .
. . . ..
. . .. .
. . . .42
Limbs ...
' " .....
.. .
.. .
... .
..
... ...
.. . .
. . ..
. . ..
..
. ..
. . . ... ...
.. 43 External Characters ... ...
.46 46 46 47 48 49 49
50 50
66 70 Size ... Fatness ... Colour and Moulting ... .
,.
. . ..
. ..
...
. . .. . . ....
. . ..
..
.....
. . .Antlers ...
.Habits, Occurrence, &c ... Food
. . . .. . . . .Rutting
Season...
. . . . .....
....
..
. ....
...
...
. . .Occurrence, &c. . ..
....
....
...
...
....
...
. .Tables of Measurements of Spitsbergen Reindeer ... .
List of
Papersreferred to in the Text ... . . .
.. .
..... .Illustrations.
Page
Map of S pitsberge n . . . .. . .. . . . ... . ... . . . ... . ... . ... . ... . . . 2
Plate l. Side V i e w of lower Part of Cran i u m , showing ossa pterygoidea. a. N or wegia n Wild Re indeer (Male) from Sandalsfjel l, H a l l ingd al, Se ptember 1907 b . Male R e i n deer from Spitsberge n . . . .. ... . . .. . . . .. . . At the end Il a-d o N orwegian R e i n d eer, N asalia &c. from above . . . . III a-c. Skul I s of Gre e n land Rei ndeer from above . . . .. . . . IV a . Skull of Female Rei ndeer from N ord-Varange r, N orway, from above b. Skull of Female S p i tsberge n R e i n d eer, from above . . . .. . .. . . . V a. Sku ll of Female R e i ndeer from N ord-Varanger, N orway, from b e l ow b . Sku l l of Female Spitsberge n R e i n deer, from b elow . . . VI. Typieal Reindeer Tracts i n Spitsberge n . a. Ren sdyrlandet from Mt. Skiild kul l a , 481 m . Looking EN E. b. U p per Part of C o nway Valley near Water shed i n the D irection of Lundstrøm Valley. Looking up the Valley from a Mounta i n 876 m . . . .. . . .. . . . Fig. I. Re indeer Sku l l from above ( D i agrammatic), w ith M easurement L i n e s (V. pp. 6 l!) .... . ... . ... . . ... . ... . . ..... .. .. .... . . . .... .. .... . 7
2. R e i n d eer Skull from below ( D iagramm aticl, with Measurem e nt L i n e s (V. pp. 6-l!) . . . .. . . .. . . P 3. l�e i n d eer (V. pp. 6 Skull, s i d e view ( D i agrammatie), with Measurement Lines 11) ... . 4. N asalia &e. of Spi tsb ergen R e i ndeer, from above, rJ . . . . 5. N asalia &e. of Spitsberge n R e i n deer, from above, rJ . . . . 11 21 23 6. Nasa l i a &c. of Spitsberge n R e i n deer, from above, � . . . . .. 2S 7. Nasalia &c. of Reindeer from Karesuando, from above , rJ, 31'2 years old 27 8 --11. N asalia &e. of Reind eer, from above . . . . 29
12. Nasali a &e. of C alf from Finmark, N orway, from above . . . . 31
13. Ossa parietalia of R e i ndeer of d i ffere n t age s . . . 35
14. O ssa pari etalia of R e i n d eer of d i fferent sexes and age s . . . 37 15. S p i ts bergen Rei ndeer, rJ, Sasse n Bay, August 1911. Zool. Museum, Oslo 44 ,,16. N orwegian w ild R e i n d eer, rJ, Veodale n , Vågå, Dec. 1881, Zool. Museum, Oslo 45
Craniological Investigations.
Int roduct ion , with Expl an at ion of Measuremen ts.
T
h e reindeer, g r e a t r a n g e o f v a r i a t i o n i n r e s p e c t of appearance, and by Rangijer tarandus L i n . , i s d i s t i n g u i s h e d b y i t s its great power o f adapta bility under varying conditions of life. I t varies in size, colour, shape o f antlers, etc. And not only are its outer characters very variable, but also the size and shape of, for instance, the d i fferent parts of the cranium. Thus R. tarandus is considerably more variable than Alces alces, C. elaphus, and other species belonging to the Cervidae.The m a rked variations in the reindeer h ave resulted in Linne's old species o f 1 758, Rangijer tarandus, having been divided by severaI subsequent authors into n u merous sub-species. The American Caribou i n particular, has been grouped into a considerable n u m ber of sub
species, or even into severaI species. As a rule, the different "races" and
"species" are established principally o n the basis of e x t e r n a I eharac
ters, sueh as eolour, shape of the antlers, a n d size, and in most cases o n the basis o f very sean ty m aterial . As the reindeer is one of those animals whieh easily varies, it will j ust for this reason be of very doubt
ful va lue to try, on the basis of scanty m aterial, to determine whieh charaeters are really fixed and of such value as to j ustify the establish
ment of speeifi c races and species, as opposed to sueh eharacters as are due to sex and age o r are even merely individual variations.
With the exeeption of the S p i t s b e r g e n r e i n d e e r, which already
in 1862 was given specific ran k by ANDERSEN [2], the reindeer of Europe
were considered as ane race, until E. LONNBERG [5] i n 1909 divided them i n to Rangijer tarandus typicus and Rangijer tarandus jennicus.
I n his treatise of 1862 ANDERSEN not only emphasizes special characters i n the cranium of the Spitsbergen reindeer but also certai n c h a racters i n other parts of the skeIeton , which should strengthen the correctness of the opinion that the Spitsbergen reindeer is a distinet race. Unfortunately, AND ERSEN'S treatise includes practically n o actual measurements, and only a few ratios. Further, the author does not m ention how large a n umber o f specimens of S pitsbergen, Lapland , a n d Greenland reindeer h e h a s h a d at h i s dispasal for purposes of comparison, nor does he state the sex of the speci mens exa mined
6 ALF WOLLEBÆK.
As regards the d i fferences between certain cranial or other parts of the skeleton of the compared speci mens, he merely states that the respec
tive parts are larger or smaller, narrower or broader, shorter o r longer, and so on, in the one than i n the other.
On comparing some skulIs o f male rein deer from Spitsbergen with severaI tame reindeer skulIs from Lapland, LILLJEBORG [ 1 2] ( 1 874) came to the conclusion that some of the d i fferences, asserted by AN DERSEN , between Lapland and Spitsbergen reindeer were not constant. A fter all.
the cran ium of the Spitsbergen reindeer did not d i ffer so much from that o f the continental reindeer as should be the case according to ANDERSEN'S statement. According to LILLJEBORG'S investigations there did not exist any other remarkable d i fferences i n the osteol ogical construction of the head than the somewhat smaller size and the peculiar shape of the points of the n asal bones. The other d i fferences pointed out by ANDERSEN between the S pitsbergen, Greenland, a n d Lapland reindeer, apart from the cranium , have not been subj ected to any' further i nvestigation by LILLJEBORG. But, i n spite of the fact that LILLJEBORG does not find the racial d i fferences mentioned by ANDERSEN in the construction of the cranium of the Spitsbergen a n d that of the Lapland reindeer to be so constant as might have been expected according to ANDERSEN, yet he maintains that the Spitsbergen reindeer is distinctly characterised as a separate race.
For furth er e n l ightenment on this question I have examined a num ber of reindeer skulIs collected i n Spitsbergen by N o rwegia n explorers (especially by Captain ISACHSEN a n d A. HOEL), a n d I have compared them with the rich m aterial of reindeer skulIs from oth er l ocalities stored at the Oslo Zoological Museum. In all, I have examined over 200 rein deer skulIs. This comparative i n vestigation
h a s proved that severaI of the d i fferences stated by ANDERSEN between the Spitsbergen and the Continental reindeer d o not exist, but, on the other hand, I find other characteristics, not previously pointed out, whereby the reindeer skulIs from Spitsbergen conspicuously d i ffer from all the others in m y m aterial. Before giving a fu rther account of the peculiar features found in Spitsbergen reindeer skulIs, I will make some re marks on the m easurements taken and the mate
rial at my disposal.
I have personally made all the measurements, and each skull has been measured i n exactly the same m anner. Besides the measure
ments o f the distances wh ich LONNBERG has made use of in his com parative studies o f reindeer races, I have also taken m easure
ments of some other distances used by other i nvestigators i n the study of racial m a rks. As, however, one and the same distance may be m easured and ind icated d i fferently by d i fferent i nvestigators, I find it
THE SPITSBERGEN REINDEER. 7
--- ---�
'�---+_---�B
Fig.
1 .Re � dee � SkUl � rom abo � e �� grammatiC), with Measurement Lines
(V. pp.6-
I I).8 ALF WOLLEBÆK.
necessary to define the nomenclature and to indicate in detail the points between which the m easurements h ave been taken .
All the measurements i n the tables give the shortest distance - t h e straight line - between t w o points.
l. Maximal Length oj Cranium
(Fig.
1, T -T)
measured from the anterior point of praemaxilla to the posterior point o f crista occipitalis. Even in calves a few days old this distance will be longer than the condylo-basal lengt h .
2. Condylo-Basal Length,
(Fig. 2, C)
the distance from the anterior point of praemaxilla to the posterior edge of the condyli occipitales (cf. O. THOMAS, [28], 1905).
3. Basal Length,
(Fig. 2, Bs.)
the distance fro m the anterior point of praemaxilla to the l owest anterior edge of jara men occipitale (cf. O . THOMAS).
For this distance LONNBERG (1906 & 1909) uses the designation Basicranial Length. In his work on red deer i n N o rway COLLETT [6], on the contrary, designates by basi-cranial length the distance from the posterior edge o f the condyli occipitalis to the point o f praemaxilla (above designated as the condylo-basal length). N either COLLETT (1909) n o r LONNBERG (1906) i ndicates i n his treatise on red deer what h e understands by basi-cranial length. W h a t LONNBERG m e a n s by his basi
cranial length is to be seen from his work "Om Renarne etc . " (1909), in
which h e says that the basi-cranial length i s the d istance from t h e
anterior edge of the joramen occipitale to the point of the praemaxilla.
What COLLETT has meant by the basi-cranial length appears from d i rect measurements o f the red deer skulIs i n the Oslo Zoological Museum , used by COLLETT. The basi-cranial lengths stated by COLLETT and LONNBERG in their treatises o n red deer are thus not di rectly com parable.
4. Palatal Length,
(Fig. 2, P).
distance from the point of praemaxilla to the posterior edge of ossa palatina at the end of the suture. (cf. O . THOMAS, 1905).
5. Maximal Breadth oj Cranium
(Fig. B- B)
measured between the outer edges o f the orbits.
THE SPITSBERGEN REINDEER. 9
� ____________ Øbr ____________ �
u
Fig.
2.
Reindeer Skull from below (Diagrammatie), with Measurement Lines (V. pp. 6�I l).10 ALF WOLLEBÆK.
6. Breadth of Forehead .
(Fig. l, P -P).
The measurements are taken between the outer ends of the suture between frontal and lachrymal.
Between these points, h owever, the frontalia have n o t their maximal breadth . In order to obtain a n expression for the b readth of the fore
head, I have, h owever, chosen to measure the distance between the above mentioned fixed points, which are always d istinetly marked, in
ste ad of the d istance between the u n fixed points between the middle of the orbits as used by some authors.
7. Breadth of Cranium at the External A uditory Meatus,
(Fig. 2, Obr.)
measured between the points of the p rotruding lateral parts of pars squamosa straight out from th e auditory meatus.
8. Breadth of Cranium behind the Canines,
(Fig.
1, h-h)
is measured between the outer sides of maxillae directly below the posterior upper points of praemaxillae.
9. The Breadth of Condyli occipitalis.
(Fig. 2, cd).
The distance between the outer edges at the base of condyli occipitales.
lO. Distance between jugalia
(Fig. 2, tp-tp).
measured between the o uter edges of the jugals at the posterior points of ossa zygomatica.
1 1. Distance between the Processus paramastoidei.
(Fig. 2,
pm-pm.)
12. Distance between Processus styliformes tympani
(Proc. styloidei, Fig.
2,
s-s).These short processus, to which ossa hyoidea are connected, are sometimes lying deeply concealed between bulla ossea and processus paramastoidei.
JtT"\... lr"\...
ø._ �� Fig.
3. Reindee r Sku ll, side view (Dia grammatie ), with Me asur em ent Line s
(V. pp. 6-11).-l ::r: (Tl (Jl '"ti ::j (Jl tXl (Tl ;:o Cl (Tl z ;:o (Tl z O (Tl (Tl ?Il
12 ALF WOL L E B Æ K .
1 3. Distance between Processus pterygoidei
(Fig.
2, pt�pt)measured between the outer sides o f processus pterygoidei (maxi mal distance) .
1 4. Distance from the Anterior Edge o f Orbita t o the Point of Praemaxilla
(Fig.
3,i)
measured from the superior edge of os zygomaticum in the a nterior edge of the orbit to the anterior poin t of praemaxilla.
1 5. Distance from Foramen infraorbitale to the Point of Praemaxilla
(Fig. 3, fl.)
Foramen infraorbitale may now lie j ust above the anterior edge, now over the middle, and someti m es so fa r back as a bove the posterior edge o f first premolar.
1 6. M axilla.
The l ength of this bone is measured from the anterior point i n front of t h e canines to t h e posterior point at the inferior edge of os zygomaticum (fig. 3, M ) .
The breadth is measured perpendicularly on the longitudinal axis.
The m axi mal breadth (height) of maxilIa is as a rule over or i m mediately in front of the secon d premolar ( Fig. 3, M h).
1 7. Os zygomaticum.
The length is measured from the point of the posteri or processus
to the anterior edge o f the bone at the suture a g a i n s t maxilla (almost
at the end of the sutu re against lachrymale ( Fig. 3, Z-Z).
The breadth is m easured perpendicularly on the longitudinal axis from the superior poi n t o f os zygomaticum in the anterior edge of the orbit ( Fig. 3 , z-z) .
18. Distance between the Superior Edges of Lacrymalia
(Fig. 1,
t-t)is measured between the anterior upper edges o f lacrymalia at the suture against maxilla.
19. Praemaxillae and the External Nasal Opening.
The d istance between the posterio r upper points are measured between the insides of the j aw branches at the anterior point o f the os supra maxillare accessorium ( Fig. I, i-i ) .
THE SPITS B E RGEN R EIND E E R. 13 For the distance at the middle of the nasal opening, see Fig. I, m-m . The length o f Praemaxilla Fig. l, i mo
T h e distance from the anterior edge of nasalia to the a nterior point of praemaxilla see Fig. l, ni).
20. Nasalia.
The posterior breadth is measured at the end of the suture between nasalia and m axillae (Fig. IN. br.). The smallest breadth o f nasalia usualy lies i n the anterior third-part of nasalia at the posterio r edge of os supra maxillare accessorium, more rarely farth e r back. Occasionally the breadth is smallest at the anterior edge of nasa lia ( Fig. I, N. br. ) .
2 1 . Breadth o f Os palatinum,
i s measured between the fi rst pair of premolars (Fig. 2, l p), the fi rst pair of m olars (Fig. 2, I ml, and the third pair of molars (Fig. 2, 3 ml.
22. Distance between Canines and Praemolares, see Fig. 3, h pr.
23. Length of the Cheek- Teeth Row, see Fig. 2, T .
24. Single Teeth.
Measurements are always maximal breadth a n d length.
Rem arks on t he Material Exam in ed.
O f the 2 1 Spitsbergen reindeer skulIs examined, the fol lowing 7 were brought h o m e by Captain ISACHSEN from his expeditions to the northem parts of Spitsbergen in 1909- 1 0 :
Z. M . No. 6615 0'. Total length o f cranium 300 m m .
- 6614 - 295 -
6619 - 6613 S".
6616 - - 6618 - - 6617 -
272 - 285 - 267 - 250 - 248 -
M a l e s . I n the speeimens with a total length of cranium of 300 and 295 m m . the milk-teeth are shed , the new premolars fully developed, somewhat wom i n the case of the former, not wom i n the case of the latter; this applies also to the molars.
14 A LF W O L L E BÆ K .
The third m a l e with a length of cranium 2 7 2 m m . is still a calf.
The thi rd molar has not broken th rough over the jaw-edge, the milk
teeth have not been shed. The antlers consist i n this speeimen o f only a single straight branch , 40 c m . long, without snags, and of a brow-tine, also without snags, 1 1 c m . long.
F e m a l e s . Only the specimen with a total length o f cra n i u m of 285 mm. has finished the shedding o f teeth. The cranium belongs to an old animal with premolars as well as molars m uch worn .
Specimen N o . 66 1 6 (Total length of cranium 267 m m . ) is i n the teeth-shedding stage. In the right maxilla the m il k-teeth a re a bsent.
O f the new premolars (permanent) the fi rst has j ust broken th rough.
the second enti rely and the third half broken th rough . I n the left m axilla the fi rst premolar (milk-tooth) is n ot yet shed ; the new premolar is to be seen above ; the bone at the front of the tooth has partly been broken through . T h e second and third premolars (permanent) have alm ost entirely broken through ; the third m olar, on the contrary, only about half broken through in b oth maxillae (first and second molars com pletely developed). In the left mandibula, all the milk-teeth are shed , the new pre m olars are not com pl etely full-grown . In the right mandibula, the third premolar (milk-tooth) is still left ; it has a very worn crown a n d very small short roots left. Below this and considerably above the jaw-edge the distal part o f the new pre m olar is to be seen . The fi rst and second premolars are somewhat more developed than in the left mandibula.
The two remaining calves (of 250 and 248 m m .'s total length of cranium) both still have all the milk-teeth lert. The second molar is not yet quite grown in either of them . I n the j aw-bone above the third molar only a n inconsiderable opening is to be seen.
The following 4 sku lIs were broug h t home from Spitsbergen in 1 898 ; they h ave been lent to me by the Trondhj em Zoological M useum .
Thj. Mus. No. 1 1 �. Total length 1 2 calf
1 3 1 4 -
of cranium 285 m m . 200
1 90 1 78
The largest specimen belongs to an old animal (�) i n which the premolars as well as the m olars are considerably worn. I n the th ree others (calves) none o f the molars are yet visible above the jaw- edge.
From the Tromsø Museu m I have obtained the following 1 0 craniums fro m Spitsbergen:
THE SPITSBE RGEN REINDEER.
Tr. Mus. No. 2 rf'.
4
-
3
�.
.- 10 - 7 - 9 -
8 calf
5 -
6
Total lengths of cranium 335 m m .
321 305 285 273 272 266
220 217 207
15
full
)
grownI n the three males the teeth are rather much wom, most i n the speeimen with a total length of cranium of 321 mm. The specimen with tot. er. length of 335 mm. has large st rong teeth, the molars especially being extraordinarily broad.
The same speeimen has the most developed antlers , the brow-tine, with 7 snags, shovel-formed.
The largest female cranium has teeth n ot much wom ; a l l (including the posterior molars) are fully developed.
The speeimens with a total length of cranium of 273 and 272 m m . o n t h e other hand, h ave both, m uch wom teeth. I n t h e smalJest full
grown female (er. tot. 266 m m . ) the teeth are yet only slightly wom.
I n the calf craniums the point of the 1 st molar is only j ust visible a bove the j aw-edge (the tip projects at most on ly one millimetre). All three specim ens have milk-teeth.
For com parison with these 21 skuIIs from Spitsbergen I have h a d an opportunity to examine about 200 skuIIs of wild a n d tame reindeer, ful l-grown males and females as well as calves, fro m the Continent, besides 22 Greenland speeim ens. In all of them the measurements were made from the same parts of the cranium as in the Spitsbergen skulJs.
Cranial Len gth.
The small skuIIs of th e Spitsbergen rei ndeer has been e m p hasi·
zed as one of the eharaeters on whieh the establishment of the Spits
bergen reindeer as a speeiRe raee is based . NITSCHE [22] (1893) has not however, from th,e m aterial he has examined, been able to point out any noticeable d i fference in size between the skuIIs of reindeer from Spitsbergen and from the Continent. NITSCHE has, o n the whole, com pared only 5 skulIs, a l l of young not ful l-grown animals. H e considers it h owever a faet, a s other authors (especially ANDERSEN) h ave previously held, that the Spitsbergen reindeer has a decidedly
16 ALF WOLLE BÆK.
s m aIIer l ength o f cranium than wild as weII as tame Russian and N orwegian reindeer. The m easurements m ade by m e further strengthen this. The sm aIIest o f aII the skuIIs o f fuII-grown males examined belong to Spitsbergen specimens, the largest belong to specimens from the forest tracts in Sunjel. The largest and probably the oldest males in my m aterial from Spitsberge n , with rather badly worn teeth , h ave a smaIIer length o f cranium (up to 335 m m . ) than a 3 1/2 year old male from Sunjel (Iength of cranium 340 m m .) i n which the shedding o f teeth was recently com pleted (the youngest and smaIIest o f the 17 ful l-grown m ales examined from this districtl. The smallest fulI-grown male from Spitsbergen has a length o f cranium o f only 295 mm. (basal l ength 274 mm.), th e largest Sunjel male o f 408 m m . Between these lim its Iie aII the rest o f the cranial lengths of ful I-grown males examined from various localities. Apart from the Spitsbergen specimens, the l i mits o f the cranial length s o f specimens from a I I the other localities i ntermerge. Specimens o f the same age from one locality may sometimes have larger and sometimes smaller cranial lengths thal1 specimens of corresponding ages fro m another l ocality, irrespective of whether one com pares reindeer from wood tracts with reindeer from mountain tracts, although a mongst these l atter there are man y lower measurements than a mongst specimens from wood tracts .
Palat al Lengt h.
I n spite of the great d i fference between the absolute measurements i n the m aterial o f the d i fferent ages, the ratio giving the relation between the total cranial length and the palatal length shows a remarkably slight variation. Between the material fro m different localities
there i s n o pronounced d i fference in th e ratio. Even i n th e sm all
Spitsbergen skulIs the ratio faIIs between nearly the same l i m its as i n the largest skuIIs from the Continent, and irrespective o f these latter belonging to rein deer from wood tracts or mountain tracts, the ratio is practicaIIy the sam e , viz. from 58 per cent. to 62 per cent.
and only in exceptional cases slightly higher or lower.
Tot al Cranial Breadt h, Breadt h of F orehead, Lengt h and Breadt h of Nasal Sect ion.
T h e ratio, giving th e relation between l ength and breadth of the cranium , varies individually rather considerably , which is essentiaIIy due to the m o re or less protruding orbits. Between the outer edges of the orbit the cranium has its maximal b readth .
THE SPITS BERGEN REINDEER. 17 I n the cranial breadth i n proportion to the length we have, accor
ding to ANDE RSEN, a racial m a rk whereby the Spitsbergen reindeer can be distinguished from the Greenland and Lapland a n i m als.
ANDERSEN says that the Spitsbergen reindeer has a broader and some
what more bullet-shaped head in proportion to its length than is fou n d i n the reindeer from Lapland and G reenland, a n d he further asserts that this character not only applies to ful l-grown animals, but also to calves.
In a Lapland calf crani u m , which was nearly 10 cm. longer than h i s longest specimen from Spitsbergen, he h a s found s m a l l er breadth a n d
·circu m ference than in the largest Spitsbergen cranium m entioned.
A mongst the skuIIs measured by me there is some, though inconsiderable, d i fference between the cranial breadth and length in Spitsbergen a n d G reenland reindeer. In males (ful l-grown) o f the former, the ratio varies between 46.2 and 49.5, in the latter between 41.4 and 45.3; in females between 48.1-51.3 and 42.3-47.9 respec
tively.
Also the corresponding ratio in the skuIIs from the Continent
i s of ten slightly lower than in the Spitsbergen skuIIs. Very rarely the breadth constitutes half of the length in the Continental skuIIs.
B y comparing the ratios from the d i fferent l ocalities o n the Continent, the variation l i m its a re fou n d to be fai rly homogeneous.
From none o f the localities is the m aterial distinguished by a ny icharacteristically large or small cranial breadth in proportion to length . Protruding orbits cannot be seen to characterise the skuIIs fro m any special locality. The rei ndeer of the wood tracts vary i n this respect just as much as those of the mountain tracts.
Com paring the breadth of the skuIIs behind the canines, (one 'of ',the measurements which define breadth of snoutsectio n)' the ratios prove to vary between somewhat narrower l i m its in the Spits
bergen skuIIs than in the skuIIs from the other l ocalities. The individual variations are very considerable. The measurements, h owever, which jll ustrate the length of the snout, the distance fro m the front of the
orbits to the fore m ost point of praemaxilla and the distance fro m the
Jara men inJraorbitale to the fore m ost point of praemaxilla, are fou nd
in the Spitsbergen skuIIs to be no d i fferent from the same measure
ments in the skuIIs fro m the other localities, if we compare the distances named with th e total length o f the skui\. The ratios for these distances vary between relatively narrow limits in all fullgrown specimens from the d i fferent localities. The distance from the fron t edge of the orbit to the foremost point of praemaxilla is only in a few cases under 57 per cent. and over 61 per cent. of the total length of the skull, the main p.art between 58 per cent. and 60 per cent. T h e distance from the
2
18 ALF WOL L E BÆK.
foramen infraorbitale to the fore most point of praem axilla i n propor tion to the total length of the cranium va ries i n about the same ratio as the fo regoing length proportion . I n a couple of ca ses the ratio is some tenths lower than 3 1 per cent, not i n a n y case h igher than 35 per cent. , a n d i n the main between 33 per cent. and 35 per cent.
Con dyl i occip itales .
As regards the dista n ce between the outer edges of condyli ocapL
tales, which LO N N B E R G considers as greater for woodland rei n deer than for mountain reindeer, I have i n my material found considerable variati on i n reindeer from Spitsbergen as well as from the Continent. I n animals o f the same age, sometimes speci mens from wood districts, sometimes from the mountains show the greatest distance betwee n the outer edge o f condyli occipitales, and I h ave o ften fou n d that even smaller mountain reindeer h ave larger breadths between condyli occi
pitales than rei ndeer of the sam e age from wood tracts. And as far as the Spitsbergen rei ndeer is concerned, o n e o f the smaller full-grown female skulls is seen to be supplied with condyli occipitales which i n breadth (57 mm. ) almost correspo nds to one of the la rgest males (58 mm . ) .
Br eadt h of Cr an ium at t he Ex te rn al A udit ory Meatu s.
The breadth varies individual ly rather considerably both i n males and females ; the males, h owever, have as a rule a n absolutely a n d relatively broa d e r occiput t h a n t h e females. The skulIs o f t h e Green
land a n d the Spitsbergen reindeer show n o evident divergence i n this respect fro m the skulIs fro m other localites.
Nas al Ap erture.
In the more tha n 200 skulls examined, the breadth at the middl e of t h e external n asal aperture, both i n t h e m ales and females, young and old animals, is nearly always greater than the breadth between posterior points o f praem axilla . In very rare cases these breadth measures are of equal size, a n d o n l y in a single case - i n one ol the skulIs from Spitsbergen - have I found the breadth of the extern al n asal apertu re largest i n the u pper part o f the aperture just below
T H E SPITSBE RGE N R EIN D E E R . 1 9 nasalia o r at the upper edge o f praemaxillae. Such a form o f external nasal aperture was found by LON N B E RG in the " Skogsren-race" ("Wood
lan d Reindeer Race") established by h i m ( i 909) , but besides the above-mentioned case from Spitsbergen I h ave seen on ly one exa m ple of such a fo rm, namely in a wild male from the Karelen ( Finland), whilst i n 50ther wild males fro m the sam e forest districts the nasal aperture was broadest in the middle.
AN DERSEN points out that in the Spitsbergen reindeer the posterio r nasal aperture is especiall y narrow ; not even the smallest Lapland or Greenland skulIs, says this author, h ave so small an aperture as the largest Spitsbergen skull . LILLJ E B O RG'S and later on NITSCH E ' S i nvestigations con firm A N D E RSEN ' S statements. LILLJEBORG compares the breadth over the ossa pterygoidea with the distance between the posterior protuberances o f the basal part of the occipital bone, and finds that the two d istances are about equal i n the Spitsbergen reindeer,
\vhereas i n the Lapland reindeer the former b readth is the greater.
Amongst my m aterials there are, h owever, n u merous exceptions to this rule. N ITSC H E , on the other hand, com pares the breadth of the p osterior nasal aperture with the distance between the j ugals, and finds that the latter d istance i n two Spitsbergen reindeer (o" 0") i s 3.3 1 --3.44 times as long as the breadth between the posterior points of ossa pterygoidea;
in 2 N orwegian and 1 Russian ta me reindeer, on the other hand, 2.86, 3.00 and 2.97 times as long. The absolute sizes i n N ISCH E ' S Spits
bergen reindeer for the distance between pterygoids of the points are 36 and 34 m m . ; for the distance between the j ugals, 1 1 9 and 1 1 7 m m . respectively.
In the largest full-grown male specimens i n m y Spitsbergen m aterial the posterior nasal aperture is defective, i n the other 4 males the con
ditions are as follows: Largest distance between the outer sides of oss a pterygoidea: 39, 33, 36, 36 ; distance between the outer sides
of the j ugals: 1 33, 1 20, 1 1 7, 1 1 5 . From this it is found that the dis
tance between the jugals are respectively 3.4, 3.6, 3.3 and 3.2 times as long as the distance over the pterygoid poi nts. T hese fi gures agree very well with those of NITSCHE. In all the other speeimens, th e G reenland , Scandinavian and Finland , the breadth over the posterior nasal aperture is greater compared with the distan ce betwee n the jugals. I f the distance between the jugals is indicated with a and the breadth over the ossa pterygoidea with b , then
i
will be i n fullgrown males fro m :
20 ALF WOLLEBÆK.
I
l-<lJ OJ) Ol l-<lJ bil >-Q l-I
o >-Ol -� ;:;"O '" bil '" <lJ ..I< "O l- <lJ
'"
., Ol '" Ol bL o '" 0"0
Ol ..I< l- l- Ol z'"
-a 'C o Ol <lJ Ol :E '00 '"
'" Ol '00 > .e: > '" ;:J ",.:ti
<lJ
Jl .5 u blJ I- m l- ...
., S m Ol <lJ <lJ
l- .e: ., -S 'Ol ...
lJ "5 > :.:: Ol .e: "O
o.. ... O- :.:: --
I
Cf] O Z O O:::: .,I
Cf] :::.� 0_2.9 2.8
-2.8 2.7 3.1 2.8 3.1 2.P
2.8 2.7 3.0 2.P 2.4 2.9 3.1
-2.8
3.03.0 2.8
2.62.9 3.0 3.1 2.9 2.7 3.0 2.0
2.9 2.7 2.8 2.5
2.8 2.8 3.0
3.1 2.9 2.P
3.1 2.6 2.8 2.7 2.8 2.7 2.7 2.6 2.8 2.9 3.2 2.9 2.6 2.7 2.5 3.0 2.7 2.8 2.7 2.8 2.9
--3.0 2.5 3.2 3.0 2.9 2.7 2.6 3.
23.2 2.9 3.1 2.9 2.8 3.0 3.0 2.8 2.8 2.9 2.8 3.1
2.6 3.02.9
-2.7 2.9
2.9
2.9 3.0
2 8 2.7 2.P
3.0 2.4 2.8
2.8 2.6
-2.9 2.9
From the above table it appears that
�
d oes not i n any of the specimens measured gi ve so high a flgure as the h ighest in the Spitsbergen reindeer.
On ly in a few cases a re they as h igh as the l owest in the Spits
bergen reindeer. Further, it will be seen that between speci m ens from th e various l ocalities shown in the table there is not any striking d i ffe r
ence in the conditions mentioned.
Besides the d i fference in breadth , the shape of the posterior nasal aperture is also somewhat different in the Spitsbergen rei ndeer fro m
the oth ers. A proflIe view o f the cranium (PI. I ) s h ows that t h e pterygoid
points d o not slope downwards so much in the Spitsbergen reindeer as in the G reenland reindeer as well as in wild reindeer fro m Southern N o rway, tame rei ndeer from Finmark, Sweden, and Finland. In all these the "slope" is very similar, irrespective o f whether they are
"woodland" or " m o u ntai n " reindeer. Also i n a large, fulI-grown spec
imen (male) from King William Land, Arctic America , both b readth and shape o f the posterior nasal aperture a re entirely in accord with the G reen l a n d and European specimens.
The vomer, which in the genus Rangifer, as opposed to the genus Cervus, is united even with the posterior part of crisfa canalis of palatinae, extends, in ful I-grown reindeer from the Continent, as far back as the ossa pterygoidea. In the cal fskulls the vomer is united with the crista can alis enti rely or alm ost entirely back to the posterior part of the latter. I n skulIs, both of calves and ful l-grown specimens from the Continent, the vom er stretches forward and practically right
THE SPITSBERGEN R EIND EER. 21
Fig. 4. Nasalia &c. of Spitsberge n R e i n d eer from above, cl.
22 ALF WOLLEBÆ K.
up to the basal part of os sphenoideum posterius, so that os sphenoi
deum anterius, in the ventrai view, is practically altogether hidden by the posterior and upper point o f the vom er.
I n Spitsbergen skulIs, on the contrary, the vomer seldom extends either so fa r back o r so far up with the poste ri or poi nts. I n all calf skulIs fro m Spitsbergen the posterior part o f os sphenoideum is therefo re for the greater part visible, and this is the case even i n full
grown ones, i ndeed even in o I d speeimens, as the ossi fication o f the poste ri or part o f the vomer is only seldom so far advanced as in the Con
tinental reindeer. (G reenland reindeer and the above-mentioned cranium fro m King William Land correspond in this respect to the m aterial from the European continent) l.
Dist ance bet ween Cheek-Teeth Rows.
I n all reindeer skulIs the d istance between the first pair o f pre
molars is smaller than between the fi rst pai r o f m olars. I n other words, the first half of the teeth ro\vs is i n all cases divergent backwards.
The d i fference between the two distances varies for males fro m the Sunjel forest d istricts between 7 and 1 6, from I nari 5-- 1 3 , fro m U tsjoki 5- 1 4, from Karesuando 9- 1 5, from N o rth Varanger 7- 1 5, froIll South Varanger 6--- 1 6, from Petschenga 7- 1 4, from Karasjok 6- IS.
froIll M agerøy 9- 1 5, froIll Greenland 7- 1 6, for the S p i t s b e r g e n
l It may be of i nterest i n th i s eon n eeti on to be rem i n ded of the phyloge netie and system atic importance of lJomer in ccrl'idae. ADOLF RORIG writes in his work: "Oas Wachstum des Schad e l s von Capre o l u s vulgaris, Cervus e l a p h u s u n d Dama vu lgaris" (Bi b l i otheeia M ed iea, Abt. A . Anatom i e , h erausgege b e n von Professor Dr. Wilh. I�oux, Stuttgart 1904) p. 3: "Der Bau des Pflugseharbeines, Vomer. hat
hervorragende Bed eutung u n d sowo h l phyloge n etisch e wie systematisehe Wiehtigkeit fUr die Glieder d e r Cervidenfam i l i e e rlangt. Bei e i n e r gewissen Gruppe dieser Fam i lie is d i e se nkreehte Platte des Vom e r weiter nach ri.iekwarts ausgede h n t und weiter n ach u nten verlangert als b e i anderen, so dass s i e e i n e kniieherne Seheide
wan d b i l d et, d i e das Rhinari um i zwe i viil lig v o n e i n a n d e r getre n nte Abteilungen scheidet. Ein solches Verhalten i st b ei samtlichen e n d ogenen ame rikani sehen C e rvi denarten (i nc\. d e s Renn tieres, Rangifer tarandus) festgeste lIt word e n . Ahn.
l ichen Bau des Rhi n ari ums zeigen aueh die Schad e l aller ge ologi seh a l t e r e n C e rvid enformen. Die amerika n i schen Cerviden haben daher i n d i e s e r H insieht e i n e n alterti.imlieh e n C harakter bewah rt. Be i al l e n i n der alten Welt (Euro pa u n d As i e nl heim i sch e n Cervide nformen (e i n seh l i e ssl ich d e s i n d er Post p l i oeanze i t ode r v i elleieht aueh seh on frliher naeh Ame rika ausgewanderten Wapiti (Cerl'us CClna
densis) ist d er h i ntere Raum d e s R h i nariums d ureh d i e senkreehte Platte d e s Vomer n i c h t in zwe i Abte i l u ngen geseh ieden. Und e i n e n gl eiehen anatomiseh e n B a u zeigen u n ter v orweltlichen C e r v i d e n Cervalces am erican u s u n d C e r v u s mega
ceros. Diese h i er erwah nte Versch i e d e n h e i t des Baues der senkreehten Platte des Pflugscharb e i n e s hat zu der Tre n n ung in ticho- Llnd holoth i n e Cervide ngruppe die Veranlassung gege ben".
T H E SPITSBE RGEN R EIND E E R . 23
Fig. 5. Nasalill &c. of Spitsbergen Re indeer from above, cf'.
24 ALF WOLLE B Æ K .
r e i n d e e r o n t h e c o n t r a r y b e t w e e n 1 2-2 0 m m . I n the Spits
bergen reindeer there is thus, to j udge from my m aterial, an indication of a stronger d ivergency i n a backwa rd d i rection o f the teeth-rows in their front half. For specimens fro m G reen land and the Continent, on the contrary, the distance changes between the fi rst pair of premolars and the first pair o f m olars with in fai rl y similar variation l i m its.
Nasal ia.
Am ongst the osteological characteristics in the rei ndeer crani um , which i n a prominent d egree h ave been the object o f attention in the study o f d i fferent reindeer races. the form and size o f nasa lia may be specially m entioned . In the fi rst pl ace, says LONN BERG, i n h i s charac
teristics of woodland and mountain reindeer, it is evident that the shape and appearance of the nasal bones are quite unlike i n both forms, and previous authors like AN D E RS E N , L I LLJ E BORG, N ITSC H E , CAM E R A N O, and others, assert i n a special degree the d i fferences in the shape o f nasalia, i n distinguishing Spitsbergen from Lapland and oth e r rein
d eer, and the d i fferences have been considered so strongly fliarked, so constan t, that they have been given value as racial marks. My m aterial of wild and tame reindeer both from Scandinavian m ountain districts and from Finland forest tracts, sh ows all transitions between flat nasal bones, w h ich according to LO N N B E R G are characteristic of m ountain reindeer, and h ighly arched, nasal bones which is said to be characteristic o f woodland reindeer.
My material fro m Spitsbergen, on th e contrary, sh ows considerably more conform ity with the characteristics o f the form and size o f the nasal bones as employed by previous i n vestigators.
A N D E RSEN maintains that the Spitsbergen reindeer, com pared with those o f Lapland and G reenland, have the narrowest nasal bones. T h e largest and broadest nasal bone, according to this author, is possessed by the Lapland animal ; even the G reenland reindeer has a smaller nasal bone than the Lapland type. As to the length o f the nasal bones, A N D E RSEN states that i n the Spitsbergen reindeer it can hardly be contained three times in the length o f the head, whilst in the Lapl a n d reindeer t h e l ength of nasalia is contained alm ost four t i m e s i n the length o f the head. Nasalia are thus said to be relatively longest in the Spitsbergen reindeer.
My m aterial show that i n the Spitsbergen reindeer there is gener
ally a greater d i fference between the breadth o f the nasal bones at their narrowest and their breadth in front, than i n both the Greenland and other reindeer. Thus, in the Spitsbergen reindeer the foremost half of the nasal bones is not. as a rule, so equal i n breadth as i n Lap-
T H E SPITSBE RGEN R E I N D E E R . 2 5
F ig. 6. Nasa/feI &c. o f Spits berge n Reind eer from above, Cf .
26 ALF WOLLEBÆK.
land, Greenland, and other rei ndeer ; the breadth i n front is relativel y greater in the Spitsbergen rei ndeer. From Fig. 6, a full-grown female from Spitsbergen, it will h owever be seen that th e front half o f the nasal bones m a y be a l m ost entirely equal i n breadth righ t to th e tip. I n genera l , h owever, the breadth right i n front o f the n asal bones i n the Spitsbergen reindeer even absolutel y is not very much smaller than that o f the rest of the reindeer i n m y materi a l , i n spite of the skulIs being a good deal smaller. The breadth o f the posterior part o f the nasal bones, on the other h a n d , is smaller, a n d not i n a n y o f the full-grown specimens i n m y material from Spitsbergen is the breadth of the nasal bones in front less th a n the half of their greatest breadth behind ; most o ften the b readth i n front is over 60 per cent. o f the breadth behind (in 3 out of Il ca ses it i s less than 60 per cent. ) .
W h e n N I TSCHE states in the c a s e o f the L a p l a n d ta me reindeer that the nasal bones h ave their n arrowest part "am Vorderende der N a senbeine", it would be a m i stake to understand this oth erwise than that the n arrowest part o f the nasal bones is, a s a rule , further forward in the Lapland than in the Spitsbergen rei n deer. Most of ten indeed the nasal bones a re somewhat widened i n the front part also in the Lapland and other reindeer.
Confi r m a tion o f A N D E RSEN'S opinion as to the l ength o f the nasal bones being contained scarcely th ree times i n the cra n i u m length or Spitsbergen reindeer, wh ilst it is conta i n ed nearly fou r times in the Lapland a n i m a l , I h ave not, h owever, been a b l e to fi n d in m y materi a l . T h e l ength of the nasal bones varies in t h e Spitsbergen reindeer aS well as in the Greenl a n d and Lapland types between l 3 and l � of the total length o f the skuI!'
The d i fference i n breadth a n d shape is, h owever, rather striking.
Besides the mentioned greater b readth of the nasal bones i n front, their a n terior end is incised otherwise than i n Greenl a n d , Lapland, a n d the other skulIs in my m a terial . LI L LJ E BO R G has already pointed out the characteristic shape of the a nterior part o f the n asal bones i n the Spitsbergen reindeer. L I L LJ E BORG says that the n asal bone from the inner side outwards is cut o b l i q uely, so that the front edges o f both nasal bones form a n acute inwa rd angl e, whilst in the Lapla n d rei ndeer the front ends are cut almost square and with a deep sinuosity. This d i fference i s pointed out by LILLJEBO RG to be the m ost essential one between Spits
bergen a n d Lapland rein deer, and, alth ough the characteristic quoted from LI LLJE BORG as to the shape of the nasal bones is not a l\vays an altogether a pt one, yet it is correct in the m a i n , a n d this is con fi rmed not only by the materia l exami ned by me, but also by other auth ors, e . g., N I TSCH E and especia l l y CAME R A N O l41 ( 190 1 ) , who h a ve had con
sidera b l y m ore material th a n L I L LJ E BORG. N ot on l y speci mens o f tame
T H E SPITSBE RGEN R EINDE E R . 27
Fig. 7. Nasalia &c. of Rei ndeer from Karesuando from above, c!'. 31 2 years o ld.
28 A L F WOLLEBÆ K .
rei ndeer fro m Finmark, Karesuando, and Russia, b u t also w i l d reindeer from South Norway, Greenlan d , N ovaya Zemlya (wild ?), and Arctic A merica are m ost often strikingly divergent fro m Spitsbergen reindeer in the shape o f the nasal bones. All have, as distinct from the Spits
bergen rein d eer, the nasal bone of ten cut square or m ost often cut aslant (generally somewhat incised ) from the outer side forward ( practically never backwards), with a deep sinuosity, so as to form an of ten rel a
ti vel y long forward-pointed tip on the inward side o f each nasal bon e . These t i p s on e a c h s i d e of the middle-l ine very frequently project farther forward than the sid e-parts o f the nasal bones bordering on the os supramaxillare accessorium . In the Spitsbergen reindeer the tips are either entirely absent or relatively sJightly prom inent. No other specimen o f the Spitsbergen skulIs has such long tips as the one from which Fig. 6 is taken . The d i fference between the shape o f the nasal bones i n Spitsbergen reindeer and the others wil l , on the whole, appear better from the appended figures than from a description . Fig. 4 shows a strongly n otched nasal bone of an old m a l e from Spitsbergen which is the nearest approach to L I L L.J E BORG'S descrip
tion. Projecting points on each side o f the m id-line are completely wanting.
Fig. 5 gives a sketch of another common form . With this figure N ITSCH E ' S sketches pp. 1 22, 1 23 show the closest agreement.
Fig. 6 represents the nasal bone in m y S pitsbergen material whose shape bears the greatest likeness to the Finmarken and other reindeer.
For com parison Fig. 7 of a 31 (2 years old uncastrated male from Karesuando is included , also photographs o f N o rwegian and Greenland skulIs (Pl. I l-I l ! ), and Pl. I V-V o f skulIs o f a S pitsbergen and a
Lapland reindeer.
The d i fference in shape h a s prov ed constant, n ot on l y a m ongst all
the fulI-grown specimens - females as well as males - but also in calves, and even i n the smallest calves only a few days old this d i ffer
ence is already strongly ex pressed . Sketch es are given o f nasal bones i n 2 o f the smallest calf skulIs i n m y material from Spitsberge n , of which the one ( Fig. 8) lacks every trace of pointedness on each side of the m id-line i n front, the other ( Fig. 9) is supplied with quite sh ort points.
The other calf skuIIs i n the Spitsbergen m aterial are sometimes like the latter, sometimes most like the former. For comparison I give figures of a calf skull from King William Land, ( Fig. 1 0 ) from N a m d al ( Fig. I l ) and Finmark (South Varanger, Fig. 1 2) . The d i ffer
ence is clearly seen from the figures, and further explanations are unnecessa ry.
LI L L.J E BORG holds (p. 839) that jrontalia often shoot a little forward between the nasal bones in the Spitsbergen reindeer. This condition is neither characteristic nor constant in this reindeer. I t varies
T H E S P I T S B E R G E N R E I N D E E R . 29
8 10
9 1 1
Fig.
8-1 1.
Nasalia &c. of Reindeer fro m above .Fig. 8-- 9 . C a l f from S p i tsberge n . Fig. 10. Cal f from K i n g W i l l iam La nd.
Fig. I l. Calf from Namdal ( N o rway).
30 A L F W O L L E B Æ K .
j ust as much i n Spitsbe rgen rei ndeer as in the other types o f whieh I h ave had m aterial fo r examination , a eondition to whieh also N I TSCH E has already ealled attention .
O s supr am ax ill are accesso r ium.
The form o f this small bone, whieh on eaeh side borders the fron t p a r t of t h e n a s a l bones, is stated by N I TSCH E ( 1 893) t o be d i fferent in the Spitsbergen and the Lapland reindeer.
N I TSCH E [ 221 , p. 1 23 , says : " Ferner ist das Os supramaxillare aeeessorium bei den Spitzbergen ren n e n vie I sehmaler und m e h r spindel
fOrmig als bei den Lapplandsrennen , bei denen es ungefii h r ein Del
toid , d. h. ein langgezogenes Viereek mit zwei riiekwarts gewendeten la ngen und zwei vorwarts gewendeten kurzen Sei ten bildet. Aueh grenzt aas Zwisehenkieferbein bei den Spitzbergenrennen nur mit einer gan z kleinen Stelle an d a s Vorderende d e s Os supra m a xillare aecessorium , wahrend bei den zahmen Rennen diese Berii h rung sieh auf die ganze aussere kurze Seite des Viereeks erstreekt" .
The great majority of the speei mens of S pitsbergen reindeer i n m y material eorrespond i n a l l essentials with N I T SC H E ' S d rawings a n d h i s a bove deseription of t h e bones m e n tioned . H owever, they a re as a rule not quite so n arrow and fusiform as N I TSC H E depiets them , and i n some older male skuIIs the departure from the form deseribed by N I TSCH E i s very eonsiderable ; instead o f fu siform i t i s sh ort and a l m ost sq uare, about as long as broad (see Fig. 4 ) . I n t h e Green l a n d speeimens a n d i n reindeer from t h e Continent t h e o s supramaxillare accessorium is very variable in size and shape. I n severai eases the branehes of praemaxillare projeet righ t up to the nasal bones, a n d nothing or very l i ttle o f the
o s sup. accessorium is then visi ble l , in other eases the o s sup. ace. i s slender, fusiform as is general in the Spitsbergen reindeer, in other eases l a rge , broad, and of a n approximately square shape. The Russian woodland reindeer, Finmarken mountain reindeer, G reenl a n d a n d South N o rway reindeer, a l l vary so eonsiderably as t o t h e s h a pe o f the o s sup. aee. , that it is not possible to c l a i m a n y speeial shape as prev
alent in the one more than i n the other. This ca n only to a eertain degree be done as far as the Spitsbergen reindeer is eoneerned , in whieh os sup. acc. usually has a slender fusi form shape.
l T h e cond i t i o n i s th e n a s i n t h e g e n u s Cervus, for w h i c h i s stated a s b e i n g c harac
teristic that prae m a xi l l a ri a reac h up to the nasal bones, i n contrast to the Rangifer genus, i n w h i c h praemaxi llare i s said to reac h only u p to the os s upra m a x i llare a c ccssori u m , n o t t o t h e n asal bones (cf. LI L LJ EBORG [ 1 2J , pp. 787 and 830 83 1 \ .
T H E S P I T S B E R G F N R E I l': D E E R . 3 1
Fig. 1 2. i\'asa lia &c. o f C a l f fl"om F i n mark, N orway, from above.
