Ne 7 
1YVYd!IT LIBRARIES SMITHSONIAN _INSTITUTION  NOILNLILSNI_NVINOSHLINS 


INSTITUTION NOILNLILSNI 
Sa1UVUGIT LIBRARIES 
INSTITUTION NOILNLILSNI 


SAIYVeSsIILIBRARIE 


Saiuvudl) 
INSTITUTION 


S3aluvagit 


IYVYPIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3JIYVYSIT LIBRARIES 


,— NY \ aa < . NAS = — — eS SS \ 
PY Sp Cz ASN a KS of z al “Zz SN 
Mf 2S 3 aN 5 UL“ 3 Z aN 
lip 2 EN 2 @& : So 
= oe es s > @ = > = \ 
n ee a oe 7) 2 “a * 
PTITUTION NOILNLILSNI NVINOSHLINS SJIYVYSIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI 
— Ww — — 
: ce = AS cc « Wt YY ye cc 
— Po NK 2 / ~ 
= 2 BWC <= C cf Mf 2 a 
is =a FS = = Gy = 
[ea] — oO ar a Sod a 
= oO =a [@) oii (@) on 
aad — eT <= a4 x eal 
IYVYRGIT LIBRARIES SMITHSONIAN INSTITUTION | NOLLNLILSNI_NVINOSHLINS S31 YVHIT LIBRARIES 
= z Fe 
wo Asses "2 y, a LEXI 2) KE SO aN i KGLO) 
a VOR ON —- QGZy xa fk SE Kam 2 Qe £E£ \ | Kea 


S3!yvugil LIBRARIES 


BTITUTION NOILNLILSNI NVINOSHLINS SSIYVYEIT_ LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI 


22) = wn 2 Zz 2) = wo 
a = Vij = = < < 
Xs 2 &? 2 Uy. & We - = 
LK. SRA. WE, tf D n f Uj? ” Ne NA. a 7) a 
& FAYE 0 YY fog COTE: oO Se ® SV} re) x Oo 
AS ee Of te raat or = RAN ASE = ae 
aN =e = S = » NS = = = 
1YVYEI1 LIBRARIES SMITHSONIAN INSTITUTION, NOtEME ES Mies Mv INOSH DiS LIBRARIES 
Zz Zz Zz 2 
lJ Vi WW w uJ 
: =e ¥%,* : : &\ : : 
= < Yt 2 < 2 NY 2 : 
A Ss Rea Y= i a X SS = = 
fe) j oO = re) “SN = oS 
TITUTION NOILNLILSNI NVINOSHLINS S3IYVuYdIT LIBRARI ES NSO AN eat NOILNLILSNI 
z= Ge za ig Jz z 
e a S @ 2 o S) 
Ls i= 25 = x9 = 
> cea 5 2. = 
KF > Ee > & 3 > Ke 
= a = = = = 
zZ i sie i Z i Z 


= GG, 2 = 2 = So : = 
a ppp =| 
2 f¥) = z2 = z E QV" Zz S 
> = = = > = ‘ >" = >| 
Z 7) 2 7) 2 ” ‘4p Fa a | 
_LIBRARIES SMITHSONIAN INSTITUTION _NVINOSHLINS S3IYVYGIT LIBRARIES SMITHSONIAN i 
5 2 5 i a us a mA | 
A = aa as = ns ~ = 
a < io < = = a e 
= oc 3 cc = ra S ce 
= oO. 5 fa) rt a 5 fool 
= er = ma z ad = o 
STITUTION NOILALILSNI NVINOSHLINS S3I1YVYEIT LIBRARIES Bes MES Shel Uae) Shy NOILNLILSNI NVINOSHLIWS 
Zz is Ze ie z c 
ro) = ( Oo = oO Kors ro) = 
= a = a oe ced =) a 
= can res Ba Ea > = lh 
= Pa) ra a = * za rE a 
: ; z en eee ee 2 B 
= wn S = n” — wn = wn ie 
1YVvVuad Woe B RAR | ES SMITESONIAN SINS Tene NOILALILSNIT NVINOSHLINS S34 1uyvuad Mott BRARI ES ,SMITHSONIA 
= WW = = eS < = = Ws & 
2 RE 8 : SY ff 2 B 2 RK 2 
— = NS 2 E 2 “yf = = = ad 
a » 2 F ees: a Tas | a 2 
*. Ww . - 
BTITUTION NOILALILSNI NVINOSHLINS S3iu¥vVudiT LIBRARI ES SMITHSONIAN INSTITUTION NOILNLILSNI 
— Ka n %2 . — 
NS Si ty Yy 
= WX 5 Pl S 
WN S Y 
oc \ Wi foe 
S Y = Gi, fsa) 
BS, e) = 
=! Zz = 


a 
SMITHSONIAN 


INSTITUTION NOILNLILSNI 


NVINOSHLIWS 
SMITHSONIAN 


NVINOSHLIWS 


NVINOSHLINS S3IYVUYEIT LIBRARIES SMITHSONIAN 


NVINOSHLIWS- 

as 
Xx 

SMITHSONIAN 


JTION NOILNLILSNI 


yy = = < = = < = « 
g — — = — _ 
ly, % 2 2 = = a = 
i 2 8 : g 2 SSNS 
= ce z a E Wy 2 : 
= S > = > > = BN: > : 
n = ” — Ze 7) 2 z ¢Y 
TION NOILNLILSNI NVINOSHLIWS INSTITUTION NOILNLILSNI_ NVINOSHLINS 
” a wa = 7) a v 
uw wo us a uw wn = 
Te 4 oy = — = = 
x =F a _— a = e 
< GS S q < = < c . SN - < 
rs = WN" = 5 2 NY": 
a Ohi ie o = Ss) Yo 
= = a) zs J 2 : 


4171 LIBRARIES SMITHSONIAN INSTITUTION NVINOSHLINS S3!IYVYSE!IT LIBRARIES SMITHSONIAN 


INSTITUTION NOILNLILSNI 


Sa1uVvyalt 


INSTITUTION 
[ lien Tl | mre rie | 


Saluvugit 
INSTITUTION 
INSTITUTION 
Saiuvugit 


TION NOILNLILSNI S3IYVUEIT LIBRARIES NOILNLILSN!I NVINOSHLINS 


w ce oe ~ Cc 

g 2 = ae ee < Z = : 
3 = 2 = \\ = = = = 
x ce) x. BD VK fe) as fe) _ 
Z S z aN 2 z : 
g = = Ee . S Z = : 
z a 2 | a . a . 3 ; ai : 

LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31YVYaIT LIBRARIES SMITHSONIAN 

= ” 3 8 ey ie: 2 ri ; 
eh = ~ « nd a w = ds = oe euw 
= oe. ty Z = ited a es we. ¢3 Z z 
z 2fG 3 : z : : 2M 
= = Ui = = = e 2 5 Ue : 
7 2 = o _ fe) = S) Ze C 
a z —! 2 = 2 AL Z 
TION  NOILNLILSNI NVINOSHLINS S3i¥vVygII INSTITUTION  NOILNLILSNI NVINOSHLIWS _ 
3 SEC . S : Sa age 
— S = o = w = w : 
= Se 5 S E 3 5 > We ! 
= A Ss > = = a SE 
: NS" 5 ~ ah = - = SS"; 
= oe Ne oD z o 2 ee Bishi 


igi1_LIBRARIES 


~~ 


SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3!YVUSIT_ LIBRARIES SMITHSONIAN 


‘ 
» 


S3t4YVYaIT_ LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_ NVINOSHLIWS 


NVINOSHLINS S3iuvysIT_Li 


SX 
TION NOILALILSNI_ NVINOSHLIWS 


AN Wa 
SMITHSONIAN 


NVINOSHLIWS 
~» 
SMITHSONIAN 


NVINOSHLIWS 
NVINOSHLIWS 


Na 
SMITHSONIAN 


LIBRARIES SMITHSONIAN 
rIDBDADICC CpsiTUCANIAN 


ig a 2 i 2 io 2% 
4 4 ae + = = : NY 
< Be a = an < = \\ 
= eS cS sy SS 

x + + a a a 4 x NN 
ar 2 2 a3 z a 2 ‘ 

€!17 LIBRARIES SMITHSONIAN INSTITUTION NOJLNLILSNI NVINOSHLINS S3IYVYSIT LIBRARIES SMITHSONIAN 
i Ss a Ss ie e ce S Zz 
= 5 e F 2 = % = i 
> = > = a = > =e : 
re — ra re 2 - Z i 2 
Ys ine ae Z is Mae o 2 g 

TION NOILNLILSN! NVINOSHLINS S3I4YVYSBIT LIBRARIES SMITHSONIAN NOILNLILSNI NVINOSHLINS 
nw = nw = we: ” 2 ” =z y 
g = Z of) ae! = z & < : 
= z= =| “2 WS 3 z = a = 
z re) + D9 Mew = re ae re) x 
z 2 g 2 RL 8 2 2 2 g 
S aS z is . 2 = 2 5 2 
0 el fier g a Spe 5 . 2 5 z 

LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31XYVHUSIT LIBRARIES SMITHSONIAN 

2 2 zZ ih a, 3 ” 3 
4 ery i Yj “4 ow pe XY 3 4 aa) tN LBB ‘= 
5 OE a 7 es OS a a or i a 
a <x WY, YY a <x pe NN < a <1 4 c 
4 “ Wj: , 4 = = Ss oe = a yg = 
o aA ne re) S re) a re) val aos o 
a nd Pa al z ene z al za 

TION NOILALILSNI NVINOSHLINS S3J!IYVYEIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS 
5 a am: : Si a g aig 
= @ = o = ly = KS @ = 
= a or ee) = LM a os y vy 2 5 
= > > > Ud > raat 2) > z 
Ee é aie - sed 4, wy ER \- <]/ =a — 


THE PLEISTOCENE MAMMALIA. 


Paces 1—62; Puares I, II. 


| 
| GIANT DEER. 
| 
| 


@£  pPalxontographical Society, 1927. 


AD 
D32 Lz 
SC A MONOGRAPH 
2, 
fot. PAL 


SMITRSO Ayres 


e 
Ole litle JOAN TUE AUG 21 1689 


CIBRAY 
THE GIANT DEER. 


BY 


SIDNEY H. REYNOLDS, M.A., Sc.D., F.G.S., 


4Y IN THE UNIVERSITY OF BRISTOL 


Paars 1—62; Prates I—II. 


LONDON: 
PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 
Drcemper, 1929. 


Made and Printed in Great Britain. 


' 
Adlard & Son, Ltd., 21, Hart Street, W.C. ik 


MONOGRAPH 


Eke Sh MAM MALTA 


PIS ROC Ran te PER EOD: 


THE GIANT DEER. 


Order—UNGULATA. 
Suborder—ARTIODACTYLA. 
 -Famity—CERVIDA. 
Genus—CERVUS. 


Species—Cervus giganteus (Blumenbach), Lydekker. 
Cervus (Megaceros) hibermcus, Wilde, Ball. 
Cervus megaceros, Hart, Adams, Newton. 
Megaceros hibernicus, Owen, Carte. 


A COMPLETE list of synonyms 1s given by Woodward and Sherborn, ‘ British Fossil 
Vertebrata,’ pp. 332—3. In addition to the misnomer Irish Elk, the animal has been 
termed the Irish Giant Deer and the Giant Fallow Deer. In the following pages the 
terms Irish Giant Deer, the Giant Deer and the Megaceros are all employed. 


I. HISTORICAL INTRODUCTION. 


It is not to be wondered at that the remains of an animal so conspicuous as the 
Irish Giant Deer or Megaceros, and occurring in such large numbers, attracted attention 
in Ireland at a very early date, the first account being T. Molyneux’s description (1)! 
in 1697 of a fine skull with antlers found at Dardistown near Drogheda. Molyneux, 
in a paper very remarkable for the time at which it was written, showed clearly that 
the antlers were not in any way related to those of the Elk, but believing the American 
Moose, an animal with which he was imperfectly acquainted, to be distinct from 
the Elk, he proposed to refer the Irish bones to that species. He suggested an epidemic 
cistemper as a cause of the animal’s extinction. 

The earliest reference to the occurrence of the Irish Deer in England is T. 
Knowlton’s (1746) (5) description of a skull with antlers found at North Dreighton in 
Yorkshire. Pennant (1781) (9) pointed out differences between the antlers and those 
of the Elk and quoted measurements. The Klk is not the only living animal to which 


+ Numbers in parentheses refer to the bibliography. 


2 PLEISTOCENE MAMMALIA. 


other early writers have assigned Megaceros antlers, for both Pallas (7) and Buffon 
adopted the view that they were those of Reimdeer. 

The earliest Continental record 1s that of L. D. Hermann (1731) (4), who, as Cuvier 
reports, gives poor figures of remains from Massel in Silesia. Camper (1784) (10) 
was the first to pomt out differences not only between the antlers, but the skeleton 
of Irish Deer and Elk. Other early continental writers on the subject are von Rochow 
(1781) (8) and de Razoumowsky (1789) (11). 

Parkinson (1811) (12) mentioned a number of localities where remains of the Giant 
Deer were found, and alluded to the antlers from co. Clare presented to Charles I 
and now at Hampton Court. 

Goldfuss (1821) (13) issued a full description of the skull with measurements, and 
again compared it with that of the Elk ; Cuvier (1826) (15) also gave a general deserip- 
tion of the skull, his account being based on an example found in the Isle of Man 
and now in the Edinburgh Museum. He further showed clearly that the Giant Deer 
was widely distributed on the Kuropean continent. Desmarest (1822) (14) also 
contrasted the skull and antlers with those of the Elk. The remains of the Imsh 
Giant Deer were described by Oswald (17) and Hibbert (18) from the Isle of Man 
as early as 1825. 

The active study of the remains in Ireland itself was inaugurated by J. Hart 
(1825) (16), who set up in the Museum of the Royal Dublin Society’ a skeleton from 
Rathcannon, co. Limerick, comparing it in great detail with that of the Elk. He 
first used the name Cervus megaceros. A second paper published in 1838 dealt with 
the method of occurrence of the Giant Deer, but was chiefly concerned with describing 
the anatomical characters. Weaver (1825) (19) recorded the finding at Dundrum, 
co. Down, of Megaceros remains associated with recent shells, whence he drew con- 
clusions as to the modern character of the deposit. Several general questions relative 
to the Irish Giant Deer, such as the growth of the antlers, the question as to its 
contemporaneity with man and the conditions under which skeletons were found, 
were now attracting attention, and some of these subjects are dealt with in a paper 
by Scouler (1838) (24). 

The fact that the nearest relative of the Giant Deer is the Fallow Deer was first 
recognized by Hamilton Smith (1827) (20), who in his classification of Deer grouped 
the two together in the subgenus Dama. 

Hitherto such authors as had considered the subject were inclined to believe 
that the female Giant Deer, like the female Reindeer, was provided with antlers. 
Phillips (1836) (22), however, from an examination of a collection of bones found 
near Waterford, was able to show that the female was without antlers. 

The contemporaneity of Megaceros with Man was maintained by H. D. Richardson 
(1846) (31), who stressed the incorrectness of styling the Megaceros the Irish Elk. 


| The collections of the Royal Dublin Society and of the Royal Irish Academy have been 


incorporated in the National Museum of Ireland. 


Co 


THE GIANT DEER. 


Mantell, writing in 1851 (35), had no doubt as to its contemporaneity with Man, and 
even suggested that 1t was exterminated by the Celtic tribes. 

Several important contributions by Owen belong to this period. In his ‘ Report 
on the British Fossil Mammals’ (26), presented at the Cork meeting of the British 
Association in 1843, a comparison is made of the antlers of the Irish Giant Deer, 
Fallow Deer and Elk. In 1846, in his ‘ British Fossil Mammals and Birds’ (30), 
Owen gave by far the fullest account of the Giant Deer which had hitherto been 
published, and in 1848, in a paper (33) describe remains from Essex, he produced 
evidence of its contemporaneity with the extinct Pleistocene mammals—a fact which 
had been called in question by Richardson. The output of papers by Irish writers 
continued, Oldham (1849) (34) describing bones from Kiltiernan, and W. R. Wilde 
(1859) (42) mentioning the existence of sixteen crania in the Academy collection at 
Dublin. 

In 1859 appeared H. Denny’s important paper (40) entitled “On the Claims of 
the Gigantic Irish Deer to be considered as Contemporary with Man.” In this paper 
many interesting points are considered, including the distribution of the Irish Deer 
in the British Isles and elsewhere, the reason for the remarkable assemblages of 
individuals within certain limited areas, and the causes of its extinction. 

A number of British records belong to the next few years. Thus, in 1859, J. 
McEnery (41) recorded Megaceros from Kent’s Cavern, Torquay, and in 1861 R. 
Howse (43) described an imperfect skeleton from South Shields and a pair of antlers 
found after a storm at the mouth of the Tees. N. Moore (1867) (50) recorded Megaceros 
from the Cambridgeshire Fens. Dawkins, in his classical paper “‘ On the Distribution 
of British Post-Glacial Mammals ” (1869) (52), recorded it from nine caves and eight 
river deposits in Kneland and Wales. G. Busk (1875) (58) described the remains of 
several individuals from a bone-cave at Creswell Crags, Derbyshire. 

The question as to the contemporaneity of Megaceros with Man is alluded to by 
J. B. Jukes (1864) (46) in a paper “ On Some Indentations on Bones of a Cervus 
megaceros found . . . near Legan, co. Longford.” 

Papers by R. J. Moss (1876) (61) and G. Porte (1877) (62) refer to the remarkable 
discovery of Megaceros remains at Ballybetagh, co. Dublin, where the bones of a 
hundred animals were found. Many of these are now in the National Museum of 
Ireland. Porte lays stress on the erroneous character of the idea still prevalent, that 
the skeletons occur in bogs, and brings forward some fresh suggestions as to the cause 
of the destruction of so many of these animals, and of the great preponderance of the 
male. 

In 1878 appeared the first of a series of papers by Leith Adams referring to the 
Giant Deer. Many of the points on which he lays stress are discussed in the sequel. 
General accounts of the known facts relating to the Irish Giant Deer are given by 
G. H. Kinahan (1883) (79) and VY. Ball (1885) (81), and both allude to the finding of 
bones in association with flint implements at Ballynamintra, co. Waterford. Ball 


4 PLEISTOCENE MAMMALIA. 


gives a list of some of the other chief localities where bones have been found in large 
numbers. 

W. Williams (1878) (65) and (1881) (77) contributed important papers dealing, 
among other matters, with the cause of death of the Megaceros when found in large 
numbers, with the possible cause of its extinction, and of the frequent occurrence of 
the skull without the rest of the skeleton. 

Although A. Savin (1878) (64) had recorded Megaceros from the Forest Bed of 
Cromer, K. T. Newton (1880) (70) and (1882) (78) hesitated to admit it to the list of 
Forest Bed animals. Osborn (1910) (122), in his account of the Forest Bed fauna, 
groups C. dawkinsi of Newton and C. verticornis of Dawkins as members of the 
Giant Fallow Deer phylum. 

T. Rupert Jones in 1881 (75) reported the finding forty years previously of 
Megaceros in the peat of Aldermaston, Berks. This record of its occurrence in 
peat is interesting, but in view of the long lapse of time between the find and its 
publication the record should be received with caution. 

The section of Lydekker’s ‘Catalogue of the Fossil Mammals of the British 
Museum,’ which contains the Irish Giant Deer, was published im 1885, and his 
important account of the various continental races included in ‘ Deer of all Lands’ 
in 1898. 

Measurements by W. H. Potter (84) of the chief skeletons in Irish and English 
Museums were printed in the ‘ Field’ in 1888, and in 1890 the publication of Wood- 
ward and Sherborn’s ‘ Catalogue of British Fossil Vertebrata’ rendered available a 
most useful series of references to the literature. 

Although papers were published by Rathke (1842) (25) and von Hichwald (1845) 
(27), comparatively little attention was paid to the Giant Deer on the continent from 
the time of Cuvier and Goldfuss till early in the second half of the nineteenth century, 
when Cornalia (1851-71) (38) and Gastaldi (1861) (44) described examples from various 
parts of Northern Italy. Cornalia showed that the Giant Deer was a contemporary 
of Man in the valley of the Po, which is probably second ‘only to Ireland as 
regards the abundance of its remains. Riitimeyer (1883) (80), in a comprehensive 
paper dealing with the mutual relations of deer, has a section on Megaceros. 

During the present century there have been numerous continental records such as 
those of Hagmann (1908) (119) from Alsace, Martelli (1908) (117) from Italy, Dietrich 
(1909) (121) from Swabia, Schweder (1913) (128) and Grevé (1913) (129) from Russia, 
and Chomenko (1915) (133) from the Caucasus. Other references will be found in 
the bibliography. 

The most interesting point connected with these continental records has, however, 
been the recognition by Pohlig and Nehring of certain distinct races. Pohlig (89), in 
his comprehensive paper (1892) on the Deer of the Thuringian “ diluvial ” travertine, 
recognizes two chief continental varieties, Cervus (Huryceros) germanie and Cervus 
(Luryceros) italie, which are considered in the sequel (pp. 15—17). 


THE GIANT DEER. 5 


Other interesting antlers are described by Gastaldi (1875) (60), Nehring (1891) (87), 
and Pohlig (1894) (94). A paper by Pocock (1912) (127) on antler growth in the 
Cervide bears on Megaceros, and will be referred to in the section dealing with antlers. 
. Among the more recent British records are those by Kermode and Herdman 
(1904) (109) from the Isle of Man, and by Bemrose and Newton (1905) (110) from 
Hoe Grange Cave, Longcliffe, Derbyshire. A very complete account of the occurrence 
of the Irish Giant Deer in the Isle of Man is given by Lamplugh (1903) (108). 

During the present century important papers have been published on the continent 
describing deer ancestral to or related to the Irish Giant Deer. Papers by Stehlin 
(1912) (125), Joleaud (1912) (126), Hescheler (1909) (120), Soergel (1927) (139), and 
Frentzen and Speyer (1928) (140) fall within this category. They are referred to in 
the sequel. Hescheler’s paper includes a very full general account of the Giant 
Deer, while in Soergel’s there are some novel conclusions regarding the homology of 
the tines. 

While it might have been supposed that the relatively close relationship between 
the Irish Giant Deer and the Fallow Deer was beyond dispute, in 1906 Lonnberg (112) 
maintained the almost abandoned opinion that the Giant Deer is an ally of the 
Reindeer, considering it no more related to the Fallow Deer than to the Elk, such 
resemblances as exist beg due to specialization rather than to affinity. Lonnberg 
based his opinion in part on the character of the vomer, which he maintained to be 
fully ossified and firmly united to the palatine, and, as in the Reindeer, completely 
dividing the nares. Loénnberg’s paper was replied to by Lydekker (1906) (115) and 
Scharff (1907) (116), who considered that the retention by the Irish Giant Deer of part 
of the proximal ends of digits 1 and rv, as well as the distal ends, while the Reindeer 
retains only the distal ends, makes any theory of the close relationship between the 
Trish Giant Deer and Reindeer untenable. As regards the vomer, Lydekker points out 
that its ossification may be correlated with the weight of the antlers, while Scharff 
states that in several skulls examined by him the vomer, though thickened, is free 
from the palatines. 

Sir H. H. Johnston’s ‘ British Mammals ’ (1903) (107) contains a full and admirable 
account of the Irish Giant Deer, as does Millais’ * British Deer and their Horns’ 
(1897) (99), while shorter accounts are given in the latter author’s “Mammals of 
Great Britain and Ireland ’ (1906) (113), and in a paper by Scharff (1924) (137). 


II. DISTRIBUTION OF THE IRISH GIANT DEER IN THE 
BRITISH ISLES. 


The following lists of localities in Great Britain and Ireland in which Megaceros 
remains have been found are based on those given in Woodward and Sherborn’s 
‘Catalogue of British Fossil Vertebrata.’ It would probably be impossible, even if 
desirable, to record all the Irish localities where Megaceros remains have been met 


6 PLEISTOCENE MAMMALIA. 


with. Leith Adams states that it is doubtful if there 1s a county where they have 
not been found. The Deer probably reached Ireland from Scotland through the Isle 


Fia. 1.—Half front view of a skeleton from Limerick now in the Ziirich Polytechnic. Drawn from a photograph 
taken by Dr. Arneld Heim and reproduced by permission of Dr. K. Hescheler. 


of Man. It does not seem to have been plentiful in Scotland, and has been found 
chiefly in the south-west. It has been met with in Denmark, France, Germany, 


THE GIANT DEER. 7 


Austria, Hungary, Northern Italy, and as far eastward as the caves of Altai in Siberia 
and as far south as the Caucasus. 

No specimens have been found in England comparable in perfection with those 
from Ireland, and no place can compare with Dublin for the number and excellence 
of the Megaceros remains. A list of those in the Science and Art Museum (now 
National Museum of Ireland) published in 1881 enumerated three complete skeletons, 
72 skulls in the great majority of cases with the whole or part of the antlers, and 
numerous shed antlers. Many specimens have been added since the publication of 
this ist. Other skeletons in public institutions in Dublin are those of Trinity College 
and the College of Science. 

For a general list of skeletons and antlers, see pp. 48—S0. 


RECORDS. 


Great Britain. 


Caves. | Dogger Bank. 
Bleadon, Somerset. Erith, Kent. 
Brixham, Devon. Folkestone. 
Caldy Island, South Wales. Fulbrook, Warwick. 
Cheddar, Gough’s Cave. | Grantchester, near Cambridge. 
Creswell Crags, Derby. | Grays, Essex. 
Hoe Grange, Derby. |. Happisburgh, Norfolk. 
Hutton, Somerset. | Hilgay, Norfolk. 
Kent's Cavern, Torquay. | Ilford, Essex. 
Kirkdale, Yorks. | Ipswich. 
Long Hole, Gower. Jarrow Dock. 
Spritsail Tor, Gower. | Kmighton-on-Teme, Worcester. 
Ravensbarrow Hole, near Holker Old Park, London. 
Lanes. | Loxbrook. 
Vale of Clwyd, North Wales. | Lulham, near Peterborough. 
Wookey, Somerset. | Maybole, Ayr. 
Newmarket. 


North Sea off Lowestoft. 


River Deposits, etc. 


Oxford. 
Aldermaston, Berks. | River Rye, a tributary of the Derwent. 
Audley End, Essex. | Seaton Snook, Durham. 
Aylesford, Kent. Skipsea near Hornsea, Yorks. 
Barnwell, near Cambridge. | South Shields. 
Barrington, near Cambridge. | Stanway, Glos. 
Bedford. Twerton, near Bath. 
Brentford. | Ufton, Berks. 
Burwell Fen, near Cambridge. _ Walton, Essex. 
Chesterton, near Cambridge. Wastwater (stated to have been found near). 
Clacton, Essex. | Westbury, Wilts. 
Cowthorpe, Yorks. Whittlesey Mere, Cambridge. 


Coldingham, Berwick. Wigtown Bay. 


8 PLEISTOCENE MAMMALIA. 


Caves. 


Ballynamintra, Waterford. 
Cappagh, Waterford. 
Doneraile, Cork. 
Edenvale, Clare. 
Kilgreany, Waterford. 
Kallowen, Wexford. 
Shandon, Waterford. 


River Deposits, ete. 


Ask Bog, N.E. of Gorey, Wexford. 


Ballybetagh, Dublin. 
Ballinderg, near Athlone. 
Ballymackward, Fermanagh. 
Ballynamore, Londonderry. 
Belfast. 

Buttevant, Cork. 

Cappagh, Waterford. 
Cappoquin, Waterford. 
Castlebellingham, Louth. 
Chapelizod, Dublin. 
Cloggy Bridge, Cavan. 
Cloone, Leitrim. 

Dangan River, Meath. 

Dee River, Louth. 
Drogheda, Louth. 
Drumsna Bridge, Leitrim. 
Dundrum, Dublin. 


Treland. 


Dundrum, Down. 
Dungarvan, Waterford. 
Dunmow, Galway. 
Dunslaughlin Meath. 
Enniscorthy, Wexford. 
Enniskerry, Wicklow. 
Howth, Dublin. 
Johnstown Castle, Wexford. 
Kallowen, Wexford. 
Kilskeer, Meath. 
Kiltoghert, Leitrim. 
Knocklong, Limerick. 
Legan Bog, Longford. 
Leighlin, Carlow. 


Lough Derg, near Mountshannon, Galway. 


Lough Gur, Limerick. 


Lough Naglack, near Carrickmacross, Monaghan. 


Mobarnan, near Fethard, Tipperary. 


| Mullingar, West Meath. 


Naul Bog, Dublin. 
Newton Stewart, Tyrone. 


| Portumna, Galway. 


Rathcannon Bog, Limerick. 
Schiale, Limerick. 


Strokestown, Roscommon. 


_ Tuam, Galway. 


Turloughmore, Galway. 
Turvey, near Dublin. 
Waterford. 


The list of Irish localities has been kindly revised by Dr. R. F. Scharff. 


Balla Lheaney, Andreas. 
Ballaterson, Ballaugh. 


Close-y-garey, near St. John’s. 


Isle of Man. 


Strandhall. 
Kentraugh. 


III. OCCURRENCE OF THE IRISH GIANT DEER. 
A. CONDITIONS OF ITS OCCURRENCE. 


Although Hart (16), writing as early as 1825, showed clearly that the JMJegaceros 
skeletons do not occur in bogs or peat, but in the marl which is so commonly found 
underlying the peat, the error as to its occurrence in peat-bogs was hard to get rid 


of, and was attributed by Porte (62), writing in 1877, to the custom of giving the 


THE GIANT DEER. 9 


general name of “‘ bog” to every place where peat is cut for fuel. But while Porte 
stated that he was not able to learn of a single case of the Irish Deer being found ina 
real bog, Kinahan (79), as a result of his inquiry into the question, was convinced 
that the remains are sometimes found in peat.'! It is pointed out by Adams (1878) (63) 
that it is in the marly deposits or blue clay found at the bottom of the numerous lakes 
now silted up, which existed in Iveland prior to the period of peat formation, that 
the Megaceros remains chiefly occur. 

Kinahan refers to the occurrence of Megaceros in esker gravels in cos. Meath and 
Galway. Porte describes remains found beneath lacustrine deposits embedded 
among water-worn boulders. 


B. PossIBLE EXPLANATIONS OF THE CO-OCCURRENCE OF NUMBERS OF SKELETONS. 


The co-occurrence of a number of skeletons has sometimes been attributed to 
the animals having entered the water to wallow, or to escape the plague of flies at 
certain seasons of the year, and bemg unable to get out; or to their being drowned 
while attempting to cross a frozen lake; or to their having taken refuge in a snow- 
drift on a frozen lake when the weight of the herd broke the ice. It is suggested by 
Denny (40) as an explanation of the remarkable numbers of AJegaceros found in 
certain localities, that the animals formerly ranged over a wider area but were forced 
to congregate by the gradual submergence of the land. A somewhat similar suggestion 
had previously been made by Cumming (29) with regard to Megaceros in the Isle of 
Man. 

More often the association of many skeletons has been thought to result from the 
animals becoming mired, either when drinking or when fleemg from predaceous 
enemies, such as man or wolf, as is suggested by Scouler (1838) (24). This theory, 
as Porte (1877) (62) remarks, may be sometimes correct, but is difficult to reconcile 
with the occasionally wide dispersal of the bones, or with the frequent finding of the 
skulls and antlers without the rest of the skeleton, or, as is noted by Adams, with 
the fact that none of the bones show tooth-marks. Porte considered that the Bally- 
betagh herd could not have been mired. Had this happened, the remains would 
have been found in the lacustrine strata, and not, as was the case, entirely below them. 
Also, on this occasion, when the remains of 110 stags were found, several skulls occurred 
together without any other bones. Such accumulations may be due to driftage. 

Adams (1880) (73), alluding to the commonly perfect condition of the antlers, 
suggested that the animals were sometimes bogged during the rutting season, when 
stags became much excited and took to the water in pursuit of a hind or a rival. 


' On the subject of the finding of Meyaceros in peat Dr. Scharff writes to me as follows: “I 
believe where the Giant Deer has been obtained in the turf (peat) it was either washed out from 
the marl deposit, or its original position was disturbed by bog movements such as occur in some 
localities.” 


2 


10 PLEISTOCENE MAMMALIA. 


The most satisfactory explanation of the occurrence of such a mass of skeletons as 
was found at Ballybetagh is that offered by Williams (77). The lake in which the 
skeletons occur occupied a hollow in the boulder clay with the sides sloping at an 
angle of sometimes as much as 45°. The lake formed a natural trap which the animals 
could easily enter, but from which escape was extremely difficult, partly owing to 
the steepness of the sides, partly to the depth and tenacity of the clay forming the 
floor of the lake. As the author points out, the large number found may be accounted 
for on the supposition that one or two lost their lives in this fashion every season. 
It is unnecessary to assume that the animals were destroyed as a herd. 

The suggestion by C. Reid (96) in the following paragraph also bears on this 
question : 

“Those familiar with the pools containing Chara will be well aware of the appear- 
ance of shallowness and of a solid floor, which is so deceptive. The Chara and Pota- 
mogeton may grow from a depth of several feet, but they often appear to form a 
carpet of bright green turf a few inches under the surface of the clear water. Any 
animal treading on this turf would immediately plunge head foremost into the water, 
and the wide-branching antlers of Cervus megaceros would become entangled amid the 
Chara stems, and still tougher Pondweeds, so that the animal would have scarcely a 
chance to escape.” 

Porte says that the condition of the antlers—full-grown, but showing in the burr 
and constriction round the base preparation for the fall—proves that the season of 
the year at which the animals perished was the end of April (supposing the habits of 
the Irish Giant Deer to be the same as those of Deer in Ireland at the present day). 


c. CAUSE OF ITs SPECIAL ABUNDANCE IN IRELAND. 


The immense numbers of this magnificent Deer in Ireland and the fact that it 
considerably exceeded its continental relatives in size, show clearly that the conditions 
in Ireland must have been exceptionally favourable to it, both as regards climate 
and in other respects. The scarcity of great Carnivora has always, as by Adams 
(1881) and Ball (1885), been considered a chief cause of its abundance, the Lion and 
Cave-bear not having been found in Ireland and the Hyena very rarely. Dr. Scharff 
writes: ‘‘ They probably lived in an open prairie country and were comparatively 
free from enemies. Hyzenas were not to be feared, for they probably became extinct 
(in Ireland) sooner than the Giant Deer. The tremendous sweep of the heavy antlers 
was a powerful defensive weapon against the Wolf, which was always abundant in 
Ireland. It would seem that Man camped mostly near the sea among the hills, and 
may only have arrived in Ireland when the Giant Deer was on the verge of extinction.” 

Among the localities where the Giant Deer has occurred in special abundance are 
the followmg: margin of Lough Derg, near Mountshannon, Galway; Killowen 
Wexford ; Ballybetagh Bog, near Kiltiernan, Dublin; Ask Bog, N.E. of Gorey, 
Wexford. 


THE GIANT DEER. Il 


D. POSSIBLE CAUSE OF THE FREQUENT OCCURRENCE OF THE SKULL WITHOUT THE 
REST OF THE SKELETON. 


A possible explanation is suggested by Williams, who, writing in 1881, remarked 
that in the previous thirty years nearly a hundred skulls had been found in Bally- 
betagh bog, but scarcely six skeletons. He stated that the atlas vertebra was in 
almost every case found with the skull, and suggested that when the carcase decayed, 
the head and antlers separated off with the atlas and sank, while the rest of the 
carcase was carried away by currents out of the lake. In one case the legs were found 
with the skull, the trunk having apparently floated away. 


E. CAUSE OF THE PREPONDERANCE OF THE SKULLS OF THE MALE. 


It has always been recognized that this is probably to be attributed to the great 
weight of the antlers causing the male to be more frequently bogged. According to 
Porte, a possible cause of the 36 skulls found at Ballybetagh being all male is that 
the animals perished through some sudden catastrophe when the stags and hinds 
herded apart. Dr. Scharff suggests in a letter that possibly the Deer went into 
Ballybetagh lake, which is one of the highest localities where they have been found, 
merely to escape the plague of flies at certain seasons of the year, and then got mired 
occasionally in the weeds. 

The preponderance of male skulls may sometimes be more apparent than real, for 
Dr. Scharff writes: ““ When a skull without antlers is found during the digging for 
peat, it is generally broken up and considered worthless, being often mistaken for a 
horse’s skull, whereas the stags’ skulls are known always to fetch a good price.” 


F. CONCERNING THE CONTEMPORANEITY OR OTHERWISE OF THE [RISH GIANT DEER 
with MAN IN THE BritisH ISLEs. 


The question as to the contemporaneity of the Irish Giant Deer with Man is one 
which has attracted naturalists from the earliest times, and although the clearness 
of the continental evidence for contemporaneity, particularly im France and Croatia, 
has caused the interest in the subject to be now mainly historical, it may still be well 
to state the opinions and arguments of the various writers on the subject. 

Molyneux, the earliest describer of the Irish Giant Deer, says that it became extinct 
in Ireland so long ago that no tradition of its former existence survives. Owen (30), 
who discusses the subject, remarks that “the total silence of Caesar and Tacitus 
respecting such animals renders their existence and subsequent extirpation by the 
savage natives a matter of the highest improbability.” Later writers have held 
conflicting views. Thus, Richardson (1846) (31) maintained that it was contemporary 
with Man and hunted by him for food and clothing, and as evidence of its relatively 
recent extinction stated that he had often peeled the periosteum off the bone. 
Mantell (1851) (35) considered that it was probably exterminated by the Celtic tribes. 
The evidence available at the time of writing (1859) was assembled by Denny (40) 


12 PLEISTOCENE MAMMALIA. 


without adding any fresh arguments. Jukes (1864) (47) described some indentations 
on limb-bones, which he believed could only be produced by a cutting instrument ; 
but Carte. in the discussion on Jukes’s paper, attributed the indentations to pressure 
and friction, and in 1867 described bones from Lough Gur showing indentations 
which he thought due to the same cause. Ball (1885) and Ussher allude to the 
occurrence of long bones split as if for the extraction of marrow, in association with 
stone implements in Ballynamintra Cave, co. Waterford. Adams, in 1878, stated 
that none of the bones showed that man or beast preyed on the animal, and in 1880, 
in his paper on “ Recent and Extinct Irish Mammals” (73), expressed the belief that 
there was no valid evidence showing the contemporaneity of Megaceros with Man in 
Ireland. Later m the same year, however, he changed his opinion, and in a communi- 
cation consisting of preliminary notes by himself and Ussher, recorded the occurrence 
at Shandon of implements made from the undoubted bones of Irish Giant Deer. A 
year later (1881), in describing the bones from Ballynamintra Cave, co. Waterford, 
he recorded (76) the finding in association with hammerstones of bones of Megaceros 
split as if for the extraction of the marrow, and stated that it could scarcely be doubted 
that the regularity of fracture was due to their having been split for the purpose. 
An examination of these bones, which are now in the National Museum at Dublin, 
and comparison with others in the same collection, lead one to doubt whether the 
splitting may not be due merely to the natural processes of weathering and decay. 
Similar split bones have been found in the Clare caves. There is some evidence of 
the co-occurrence of Man and Megaceros in the Isle of Man, and in England Megaceros 
bones have been found associated with human remains at Kent's Cavern, Torquay, 
while in France it was constantly hunted by Man. 

Some of the more recent writers on the subject have believed in its survival to 
alate date. Thus, Sir H. H. Johnston (106) was of opinion that it probably lasted m 
Ireland to the end of the Neolithic period, J. G. Millais (112) that it may have 
become extinct only in Recent or Prehistoric times. He alludes to a skull in his 
possession in the forehead of which is a deep cut, evidently made by a hatchet. 


G. PosstpLE CAUSES OF THE EXTINCTION OF THE [IRISH GIANT DEER. 


A long succession of writers, ranging from Molyneux (1697) to Sir H. Johnston 
(1903), have discussed this question, and the following suggestions have been made : 
1. That dense herds of these big animals may have been attacked by some 
microbial disease. 
. That human hunters perhaps by pits, perhaps by huge drives, which sent 
herds over precipices or into bogs, were the chief cause. 


i) 


ww 


. That its disappearance is due to a change in climatic conditions. 


nS 


. That its extinction is due to over-specialization in structure, possibly 
leading to sterility. : 


THE GIANT DEER. 13 


The first of these causes was advocated by Molyneux (1697). He rejects the theory 
of the Noachian flood, suggesting that extinction may be due to “ certain ill-consti- 
tution of air in some of the past seasons long since the Flood, which may have 
occasioned an epidemic distemper.” He quotes examples of such distempers affecting 
Reindeer in Lapland. Any which survived the epidemic, he suggests, were killed by 
human hunters. Hichwald (1845) favoured the view that it was exterminated by Man. 

The third of these suggested causes was maintained by Williams, who considered 
that the deposit of clay overlying that containing the Megaceros remains is of glacial 
origin, and hence that cold was the cause of the animal’s extinction. In this conclusion 
he is supported by much modern opinion. 

From the analogy of many groups of animals in which over-specialization either in 
size or in some other respect is closely followed by extinction, it seems probable that 
here is to be found the most important cause of the extinction of the Irish Giant Deer. 


H. DISTRIBUTION OF THE GIANT DEER IN TIME. 


Although, as has already been mentioned, Pohlig derives the Giant Deer from 
Cervus dicranius of the Upper Tertiary, no discovery has as yet definitely carried 
back the race to an earlier date than the Upper Pliocene. Both in England and on 
the continent, especially in Italy, remains of this date are well known. In England 
ancestral Fallow Deer like Cervus verticornis are found in the Red Crag and Forest 
Bed of East Anglia, and forms like C. belgrandi, intermediate between early Fallow 
Deer and Giant Deer, occur in the Forest Bed. In Italy ancestral Fallow Deer are 
found in the valley of the Po. Numerous records of Giant Deer (Cervus (Huryceros) 
germane) from Northern and Western Germany show that these animals had a wide 
distribution in that part of Europe prior to the last great increase of cold in Magda- 
lenian times. There does not, however, appear to be any clear evidence that the 
Giant Deer survived in northern continental Europe into post-Magdalenian times. 

Numerous records show that the Giant Deer was widely distributed m England 
during Paleolithic times and spread thence to the south-west of Scotland and the Isle 
of Man, whence it probably reached Ireland. Though recorded from the peat in 
several localities! there is little evidence of its survival in Great Britain as late as 
Neolithic times. It is, nevertheless, commonly quoted as the sole extinct member of 
the Paleolithic fauna which survived, attaining its maximum in the Irish deposits 
of the period, and persisting, so it has been claimed, even to Prehistoric times. 

Whether it survived the latest period of refrigeration is not so clear as was once 
thought to be the case. To take first the Megaceros-bearing deposits of the Isle of 


| Owen (‘ Brit. Foss. Mamm..,’ p. 466) records it from the peat of Hilgay, Norfolk ; Seeley (‘ Quart. 
Journ. Geol. Soc.,’ xxii, 1864, p. 479, and ‘ Geol. Mag.,’ 1866, p. 499) alludes to its presence in the 
peat of the Fen country; N. Moore (‘ Ann. Mag. Nat. Hist.,’ xx, 1867, p. 77) to its presence in the 
Fens, 


14 PLEISTOCENE MAMMALIA. 


Man, which closely resemble those of Ireland. The nature and age of these deposits 
are exhaustively discussed by Lamplugh (108), and the conclusion is reached that 
Megaceros must be regarded as “ either a late glacial or early post-glacial mhabitant 
according to the limits, in any case arbitrary, which we may assign to the Glacial period.” 
With regard to the Irish deposits, Williams has suggested that the Megaceros marls 
were accumulated during a mild and genial interlude which followed the main melting 
of the ice, and was succeeded by a return of glacial conditions. J. Geikie was inclined 
to accept the view that the Megaceros beds are of interglacial age, and Lamplugh 
says: “It would be rash to deny the possibility of the Megaceros marl being the 
product of a genial interval between two cold periods.” It may be doubted whether 
even in Ireland it survived the increased cold of Magdalenian times.! 


IV. ORIGIN, RACES AND AFFINITIES OF THE GIANT DEER. 


Though there is little evidence available, it is possible that the Giant Deer was not 
of Kuropean but of Siberian origin, and that it migrated westward under the influence 
of the increasing cold, of which the later Plocene beds afford proof. On the other 
hand, Pohlig (89) derives Megaceros from Cervus dicranius, a small Upper Tertiary 
form with a wide distribution in the Mediterranean region. 

The earliest fossils attributed to the race of the Giant Deer are the fragmentary 
antlers described by Owen (37) and by Dawkins from the Red Crag of Suffolk. It 
may be doubted whether the former specimen is sufficiently well preserved to justify 
any definite conclusions concerning it. Soergel agrees with Dawkins in attributing 
the latter specimen to Cervus verticornis. 

Various authors have described the several “‘ races’ into which it is possible to 
divide the Giant Deer, one of the most comprehensive accounts being that of Lydekker 
(100), who recognized five races, the German, Irish, Italian, French and Forest Bed 


b) 


races. 

The earliest race is that of the Forest Bed, to which such forms as Cervus vertt- 
corms are referable. The antlers are short, directed upwards and outwards, with 
palmation generally slight and a simple brow tine. Some of the early forms described 
from Italy, such as the simple antlers with short tines described by Alessandri from 
Monteferrato, seem to belong, as Soergel suggests (130), to an animal closely allied to 
C. verticorms. 

C. dawkinsi from the Forest Bed, which is commonly grouped with C. verticornis, 
has a more expanded palm, and is perhaps, as Soergel suggests, more closely related 
to Alces than to Megaceros. 


! Since writing this sentence I have been shown the as yet unpublished script of a paper by 
Dr. J. W. Jackson on the animal remains of the Kilgreany Cave, Waterford, in which he writes : 
‘With regard to the giant Irish deer, there appears to be good evidence that it did not long survive, 
if at all, the last cold phase, which is generally regarded as of early Magdalenian date.” 


THE GIANT DEER. 15 


Another early form is that described by Stehlin (125) as Cervus (Megaceros) dupuisi 
from the Pleistocene of Rosiéres, near St. Florent. This was a relatively small animal, 
about as large as a Red Deer, but it resembles Megaceros in the fact that a cross-section 
of the mandible is wide and rounded below, while that of the Red Deer is deeper, 
narrower and more pointed below. A related form, Cervus pachygenys (Pomel), is 
considered by Stehlin to be more primitive than C. dwpwisi in respect of the small size 
of the antlers, while as regards the depression of the antlers and the length of the 
molars it seems to be more highly specialized than C. dupwisr. Joleaud (126) has 
also described C. pachygenys, which he considers identical with Lydekker’s C. algericus, 
and agrees as to some of its characters being archaic and others highly specialized. 
He suggests that it and Megaceros may have had a common ancestor, and regards it 
as worthy of subgeneric distinction under the name of Megaceroides. 


Fic. 2.—Fragmentary skull and antlers of a young individual from co. Limerick (no. 17 Nat. Mus., Dublin). Both 
antlers are broken; the right one retains the brow tine which springs at the very base of the antler and already shows 
flattening. The left antler retains the 2nd tine. The most interesting features shown are the conspicuous pedicels 
from which the antlers spring. In old animals the antlers spring directly from the skull-wall. 


Many important occurrences of the Giant Deer have been recorded from the 
interglacial and other deposits of the Rhine Valley and from Northern and Central 
Germany, which show that Megaceros had a wide distribution prior to the last great 
advance of the ice. It occurs associated with Macherodus, Hippopotamus, Rhinoceros 
mercku and Elephas antaquus as a member of the warm Chellean fauna, and is found 
also in Mousterian and Aurignacian deposits. 

The German examples, which are seen in nearly all the Rhineland museums, 
have been described by Nehring, Pohlig, Soergel, Frentzen and Speyer and others, 
and there has been difference of opinion as to how they should be grouped and as to 
the systematic rank of the divisions adopted. Pohlig has grouped all the German 
examples together as Cervus (Huryceros) germane, which he considers worthy of 
specific distinction from C. hibermcus. The chief distinguishing features of C. ger- 
mane are: (1) the antlers are smaller, have a narrower palm, and are more upwardly 
directed than in C. habernicus, and consequently have a smaller span; (2) the tines 


16 PLEISTOCENE MAMMALIA. 


on the front border of the palm are much less developed than in C. hibernicus, and all 
those of the palm are characterized by a sharp inward bend. The brow tine is 
commonly well developed and sometimes bifurcated, and a wide gap separates it 
from the second tine, which commonly springs close to the palm. In some cases 
there is no back tine. 

A variety was described by Nehring (87) to which he applied the name Cervus ruff, 
regarding it as a distinct race. The antler to which this name was originally applied, 
and which was found near Kottbus, has a well-developed somewhat spoon-shaped 
brow tine and a long second tine, but only shght mdications of a back tine. The 


€ 
@ Fic. 3.—Stages in antler growth, after Owen, locality not stated. a, rose; b, palm; c, brow tine; d, 2nd tine 
sometimes regarded as a bez tine; e, back tine. The above lettering is retained in all the figures of antlers. Owen 
gives a full description which is quoted in the text (p. 24). 


palm is not very much expanded, and four well-developed times spring from its ter- 
mination rather than from either the anterior or posterior border. The probably 
young antlers shown in Text-fig. 4 bear a considerable resemblance to those of the 
ruffi type, and it seems likely, as Frentzen and Speyer suggest, that Nehring’s 
specimen may have belonged to a young individual. A second example attributed 
to Cervus ruffi by Nehring has a well-developed back tine. His figure is reproduced 
in Text-fig. 31, A, p. 60. Specimens described by Dietrich (120) from Canstadt and 
Jarotschin (Poland) are of essentially the same type. The fine antlers from Knielingen 
described by Frentzen and Speyer (139) are considered by these authors to represent 
an extreme development of the ruffi type connecting it with the dama type of antler. 
The resemblance to the antlers of C. dama lies particularly in the fact that a series 
of tines springs from the posterior border (Text-fig. 12, b). These are strongly bent 
backwards and upwards as in the typical germane variety. 


THE GIANT DEER. ily) 


The Italian race Cervus (Huryceros) ttalie of Pohlig (‘Text-fig. 12, a) does not differ 
very markedly from the German race. Lydekker (101) states its characters in the 
following terms: “ Relatively small, and antlers comparatively simple; palmation 
narrow, much inclined upwards so that much of the outer surface is seen in front 
view ; all the tines small, and those above the bez (v.e. second tine of the nomen- 
clature adopted here) few in number and placed near the summit of palmation.” 

In the Forest Bed and in the Pleistocene of Clacton, Essex, are found the antlers 
of other deer connecting the Fallow Deer and the Giant Deer. Thus, C. brown, 
Dawkins, from Clacton, is clearly an ancestral Fallow Deer. C. belgrandi, originally 
described by Lartet and now well known from the splendid antlers described by 
Harmer (106) from the Forest Bed of Pakefield (Text-fig. 12, c), is a much larger and 
more specialized form, occupying, as its describer claims, a central position between 


Fie. 4.—Young antlers from “ Ireland ” seen from above (Brit. Mus. Nat. Hist.). In this remarkable pair of young 
antlers the brow tine (c) is terminated by two points in the right antler, and by three in the left. There is no 2nd 
tine, but the back tine (e) is well developed; the palm is narrow and tapers rapidly. Only two tines in the right 
antler and three in the left spring from its anterior part, which ends with one point in the right antler and two in the 
left. The length of the pedicel is an indication of the youth of the animal. ‘The left antler is not unlike that of the 
variety ruffi of Nehring. 


C. browni and dama on the one hand and C. giganteus on the other. It differs from 
dama and most early forms in the relatively horizontal position of the beam, which is 
very long, sharply bent forwards and expanded suddenly into a very broad palm, the 
edge of which is marked by many slight irregularities, but bears no definite tines. 

Another type of antler is that from the sands of Mosbach near Biebrich on the 
Rhine, described by Soergel (130) (Text-fig. 12, d) as C. (Megaceros) mosbachensis. 
The beam is short and stout and is continued into a palm which, to judge by the 
figures, suggests a resemblance to that of C. dawkins:. Two tines on the right and 
one on the left antler spring near the burr. 

While the recognition of one or more continental races in addition to the Irish 
race is clearly desirable, attention may be called to the pertinent comment of Scharff, 
that some of the continental authorities do not sufficiently realize the extreme 
variability of the Megaceros antlers, and tend to found races and varieties on 
insufficient grounds. 


3 


is PLEISTOCENE MAMMALIA. 


Pohlig’s table of affinities of the Giant Deer and its allies, as quoted m an 
abbreviated form by Hescheler (120), is as follows (the names in brackets are 
those of related forms not in the table as given by Hescheler) : 


Giant DEER GRovpP. 
C. (Euryceros) hibernicus. 


| C. (Euryceros) italie. 


| 
C. (Buryceros) germame. 


(including C. ruff). 


ELK AND ALLIED GROUPS. Fattow DEER GROUP. 


| 
C. (Euryceros) belgrandt. 
(also C. carnutorum, C. verticornas, 
and C. dawhinst). 


Cervus dicranius (and C. brownz). 


V. DESCRIPTION OF THE SKELETON. 
A. THE SKULL (Plates I and II). 


The characters of the skull in the Cervide are very constant. In the Irish Giant 
Deer, as in others, there is a prominent occipital crest, and the face is less sharply 
bent down on the basicranial axis than in many Artiodactyls. The parietal com- 
pletely fuses with its fellow at an early age. The posterior half of the united frontals 
has a longitudinal ridge which, as Owen points out, bears some resemblance to the 
median prominence of the Giraffe. There is a deep depression, the suborbital or 
lachrymal fossa, in the lachrymal bone, which is large and takes a considerable part 
in the formation of the side of the face in front of the orbit. The post-orbital bar is 
complete, and the orbit is prominent and nearly circular. There are conspicuous 
supraorbital foramina. A vacuity, variable in size, occurs between the anterior end 
of the frontal and the lachrymal, partly bounded in front by the nasal and maxilla. 
The premaxilla is a y-like bone owing to the large size of the anterior palatine foramen. 
The nasal process of the premaxilla does not reach the frontal. The posterior palatine 
foramen is very small. The paroccipital process is long. The external auditory 
meatus 1s conspicuous. 

There are certain differences between the male and female crania in addition to 
those directly concerned with the antlers. In the male there is a ridge crossing the 
posterior part of the frontal, which is lacking in the female. On the other hand, the 
median longitudinal ridge on the frontals in the female is far more conspicuous than 
in the male. The condyles are smaller in the female than in the male. 


THE GIANT DEER. 19 


Stehlin (110) says that a way of distmeuishine the mandible of the Irish Deer 
from that of an Elaphoid Deer of similar size is furnished, not by the dentition, but 
by a section of the mandible. In the Giant Deer a cross-section of the mandible is 
thick and rounded, in an Elaphoid Deer it is flattened, narrower towards the lower 
border and relatively deep. 


(a) MEASUREMENTS OF CRANIUM.! 


Skull of male, | Skull of female. Skull of female, 
= no. 28968 | no. 18095 59, 1915 
| (Brit. Mus.). | (Brit. Mus.). (Nat, Mus., Dublin). 
1. Length from intercondylar notch to. 
anterior end of skull 2 6 | 47-0 43-25 45-2 
| 2. Extreme width across zygomatic arches. 23-4 22:2 21-0 
| 3. Transverse measurement at infraotbital 
| foramen . -| 9-25 | 8-35 
| 4, Measurement across orbits “at fronto- | | 
lachrymal suture : 3] 18-2 | 15-9 | 16-7 
5. Measurement across occipital condyles s| 10-7 10-05 | 9-7 
| 6. Measurement across external auditory | 
| meatus. ae 15:2 16-0 
| 7. Width of palate, measurement taken | 
external to last molar a :| 14-2 13-8 13-7 
8. Length of palate from posterior limit of 
palatine suture to anterior palatine 
foramen . : 21-5 20:9? 19-7 
| 9. Antero-posterior diameter of orbit : 6-3 5:9 5-6 
| 10. Measurement between  supraorbital | 
foramina . 5 5 9 : : 12-15 10-25 10-95 


' In the following tables all measurements are quoted in centimetres. 
> Partly by estimate. 


B. THE ANTLERS. 
(a) Description. 


The astonishing size of the antlers constitutes by far the most remarkable feature 
of the Irish Giant Deer. This is mainly due to the great expansion of the beam into 
a crown or palm, which in a case mentioned by Owen measured 30 inches across. 

Of the tines, three, namely, the brow tine, the second tine and back tine, spring 
from the beam, the remainder from the crown or palm. The first tine to be given off 
is the brow tine from the anterior margin of the beam. This tine, which tends to be 
downwardly directed as in the Reindeer, though to a far less marked extent, is very 
variable. It may sometimes, particularly in young individuals, be simply pointed 
(Text-fig. 3 A, c), it is sometimes forked near its termination, but is as a rule more 
or less flattened and shovel-like, ending with three or four short projections (cf. 
Text-figs. 4 and 9). 

The beam early gives off from its posterior surface a tine which varies much in 
length and point of origin, and may very rarely show a tendency to bifurcate (Text- 
fig. 8, c, e), while occasionally it is absent, It is the only tine which normally springs 


PLEISTOCENE MAMMALIA. 


20 


‘oYEIpVL SOUIY JO Solos B YOrUM uLOIy ued Io Weveq MOLIeU & 
Sey @ pu® 9 Ul pornsy JoyjUe oy], ‘feu st ou puZ 944 (pernsy you) szojyue 4Jo] oy} uP (0) eu Morag oY44 04 quooe[pe Ajoyerpouruit ueeq 044 ulo1f sputids (p) auiy 
pug oy} se pepresor oq 04 Ayquqoad st yeya 4eyy Jovy OYF LOF O[QVYIVOLEI SI | “puo [eystp S91 Woy omg PUB OOvF LOMOJUe 841 WO’, Oy ‘UUped O44 MOA DuLads sour 
imo}y ATU “pues oy} 4e poytoy AYSYS our Modq [eoltpulféo ojduis ve sey wo “Mora quody “ouLeS CYT, @ *eAoqe wLoay uses Jo[yue 4yoT “(EF ON) Q ‘sopjue 44S Jo 
MOTA QUOI “((906T) $93 ‘ON) HX “s9]ZUe qystr Jo Mora quoLy “(76 ON) W “aqua “Unosnyy [PUOTeN YY Ur [Te ‘UTTGn¢ ‘oo ‘yseqoqAleg wosy saopyue BunoK—e “ory 


THE GIANT DEER. 21 


from the posterior surface, while from the anterior surface spring a series of tines 
varying in number according to the animal’s age. 

According to the nomenclature usually accepted at the present day there is no 
bez tine, though Owen and Flower and Lydekker regard the second tine springing 
from the anterior margin as a bez tine. Soergel’s study of the German specimens 
leads him to the conclusion that the tine usually regarded as the brow tine 
(Augsprosse) of the middle and later “‘diluvial”’ Giant Deer is really the downwardly- 
directed equivalent of the second tine (Hissprosse) of early forms. 

In some cases the true brow tine 1s apparently represented by a vestige (“ oriment,”’ 
* offer ’’). 

In normal antlers a series of about five long curving tines follow the second tine 
along the anterior border and distal end of the beam. Their disposition is variable, 
but they tend to curve backwards and inwards over the skull. The last tines of the 
series are the most variable and may be much shorter than the others. Occasionally 
two tines, instead of springing separately from the palm, may arise from a broad 
common base (Text-fig. 9, c), while on other occasions a tine may give off a small 
branch from about half-way up (Text-fig.9, c), or very rarely may fork at the end. 
The posterior face of the beam bears no tines and is normally smooth, but occasionally 
in abnormal antlers irregular projections may occur along it (Text-fig. 10, c). 

The extraordinary variability of the antlers may best be appreciated by an 
examination of the figures. Many irregular and abnormal antlers exist, some of 
the most remarkable being shown in Text-figs. 10 and 11. It has been claimed that 
bifurcation and flattening of the tines affords an indication of advancing age, but in 
the case of the brow tine bifurcation and flattenmg is a normal feature and may 
commence early. The figures accompanying this Monograph show that bifurcation 
may affect any of the tines. 

Although there is no proof, it seems probable that certain very massive antlers 
with a reduced span and shortened tines are those of aged animals. Such antlers are 
represented by Text-fig. 10, p, and one of their features is a tendency to flattening 
and bifurcation of the second tine.' 

One of the most reliable indications of the animal’s age is afforded by the sprmging 
of the antler. In young animals it is borne on a distinct pedicel (Text-fig. 2), but in 
old individuals this is much reduced, the antler springing almost directly from the 
surface of the skull. A strengthening ridge may, as is mentioned by Owen, be 
developed along the under-surface of the beam in exceptionally large antlers, such as 
those at Knowle, near Sevenoaks. 

It has been stated that there is a tendency for the left antler to be larger than the 
right. This claim finds no support from an examination of the large series of antlers 


1 Dr. Scharff agrees that the antlers figured in ‘ Land and Water,’ April 16th, 1892, are those of 
an aged individual. The antlers are stated to be in the Derby Museum, but I am informed by 


Mr, F. Williamson, the Curator, that this is not now the case, 


PLEISTOCENE MAMMALIA. 


22 


‘gfsue onbyqo APYSYs & wLOI, UMeIp Useq SuIAvYy suouTIOds 94} 04 SUIMO SuLUS}10YSeI0F OUMOS SI OY pue @ ‘ywuy ‘sjutod 4104s ve144 Aq poyeuruts.e, 

SI pUv OOF JOMOqUB S}I UO SouTy UO] de14} saveq wed oY, ‘o[qIsIAUT sI ouTy YOeG OUT, “(FI _O8Bo-[[eM oaoge “4s “4eN “SNy “4uE “FzEz ‘W) ., PUPlery ,, Wosy 

THIS JO JopJUB gJoT JO MOIA JOMoyUL-o1oyuy G “wyed oy} Jo oovyz sols0zUe oY} WWOIy Sulads soury suo, Ar9A nog *(gT 9SBO-]]BM BAOGR “ASIF “GUN “SN “4g ‘EzEz IN) 

cc PUBLOIT ,, WOAF [NYS FO JO[JUB 4JoT JO MOIA JOMOFUI-O1OjJUY MH “UMOYS A[IBO[O OLE SpUO II9Y} ZeEU OAIND 04 souTy 94} [[@ Jo Aouepucy Su0d4s oyy pur (9) ou Mosq ayy 

JO Jopowiwyo poeyoueiq pues poueszey oy, “Lojyue oUleS OY} JO MoIA JOMOzUI-O1UY = ‘wayed oy} JO o0¥j JOlI0}Ue O44 WOIz SuIsuIAds SOUly 94} JO YYZue, O44 MoYsS 04 
PoepuozUr st puv “puryod UOd} UoYe} SI MOTA OUT, “(4SIH YEN) Wnoesny yspIg O44 Jo Aro] [V3 oy} ur dn yos ., puvpory ,, Woy Uoyo[ayxs ayy Jo IoyUe 4JoT W—'9 “OIA 
a — 


oD 
N 


THE GIANT DEER. 


“ey[ooRy[eg ‘louvyy woysuBIy) 48 
poAdesoid pue ‘UMO049]}8eD PLOT 03 Sursuojoq ,,‘puvjery ,, WoT, G ‘Ae[ndor11 pue [euIOUqe YeYMoeTIOS SI JopJUe Ajo] OYJ, “Wnesny_ vAoua*) O49 UI MOU ‘YOLIOUIIT 
WO0lg Q ‘MORseTD ‘tnosny AZ1D 04} UI MOU .‘pUBlOIT ,, WOT | “UOWeoInjsiq Jo usIs oUIOS SMOYS OUlZ YOVG FYB 9Y4 pUe soyBoINJIq 9UIy MOIq eyy, “WNIsleg 
‘uIBANOT ‘aseT[OD FInsee oYy4 09 UY Aq pue ‘uUOg jo ‘zqueIy “7 “IG 09 plos ‘HoWoUT]T WOIG W  “yUUIYST[qQeISe ssuIRITIAA “SIssepy 4Je_ Aeyy o10joq uayxe, sydeisojoyd 
WOIJ UMBIP oIv seINnSy oy} pue ‘ul[qnd Jo ‘Uog H smRTTIA\ ‘sussoyy Aq potddns pue poureqqgo [je oom q pue Q “WY soInsy url UMOYS sio[JUe oYT—*) “OIG 


Aiea: 


24 PLEISTOCENE MAMMALIA. 


at Dublin. The left brow tine is, however, in some cases larger than the right, as is 
commonly the case in the Reindeer. 


(6) Young Antlers. 


Cuvier alludes to the lack of knowledge of the young antlers, and Owen (1846) 
makes a somewhat similar remark. It is a singular fact that these statements are 
even now nearly as true as when they were originally made. In the great collection 
at Dublin, although there are a few antlers (cf. Text-figs. 2 and 5) which are not 
fully grown, there are no complete examples of really young ones. Owen, however, 
figures three antlers which he remarks “ appear to indicate progressive epochs in the 
age of the animal.” His figures and descriptions are appended, the lettermg beimg 
altered in accordance with that adopted here : 

“The first (Text-fig. 3, A), which apparently corresponds with the state of the antlers 
in the fourth year in the Fallow Deer, 1s five feet in length and fourteen inches across 
the palm ; it presents a simple cylindrical brow antler (c); a short and simple bez 
antler (d); the hind branch almost straight, and only two long branches from the 
fore-part of the palm, which terminates in three short straight obtuse points, the 
middle being the longest. The second figure (Text-fig. 3,8) shows an expansion and 
flattening of the brow antler, an elongation of the bez antler and of the anterior 
branches of the palm, and the prolongation of the three terminal poits into branches, 
the total number of branches being eight. The length of the antler, followmg the 
curve, is six feet; the greatest breadth of the palm fifteen inches. This form of 
antler corresponds with that at the fifth year in the Fallow Deer. In the third figure 
(Text-fig. 3 c) the brow antler is expanded and bifurcate ; the bez antler is likewise 
expanded and forked, but this is a very rare variety. The palm is much increased 
in breadth and sends off six branches besides the posterior one, the number of 
points in this antler being eleven. The length of the antler, following its curve, is 
seven feet, the breadth of the palm thirty ches. Such an antler would indicate the 
Megaceros to have reached the prime of its age, like the “crowned Hart’ of the 
seventh or eighth year.” ! 


(c) Comparison of the Antlers with those of other Deer. 


As Megaceros antlers have been compared with those of the Elk, the Remdeer 
and the Fallow Deer, it may be well to indicate the points of resemblance and 
difference. 

In the Elk the antlers agree with those of Megaceros in their depressed position 
and in the great expansion of the crown or palm. The antlers differ from those of all 


1 It is probable that the antlers figured by Nehring, on which he founded his race C. ruff, 
may, as he at first concluded, have belonged to a young individual. 


THE GIANT DEER. 


‘YSINGuIpH ‘Wnosnyy YSy{00g [BAOY oY} UI MOU ‘URTT JO ofS] 94} WOLF WOJOTOYS OY} JO JopJUL 4Jo] JO MoIA JAMO] YJoT CG “poyxso} ore souy yor 
pue Moiq Woq “ulqnd ‘eseTjop Ajruly, Jo Wnesny{ [RolsoToes) 049 UI MOT ‘YorIemTyT ‘Inx) YSnoy WoI, UOJoTOYS JO JopjUR 4jo] JO MOIA IOMAzUI-OlOJUW Q “a]qISLAUI 
st “quesord Yysnoyy ‘oul yorq oy, “poYxoy st oun pug oy, “(uNqng “sn yen) Tesuney, Uoorepyory Aq pojuasoad (youeuIy) UOUURoYeYy WOT; UOJeTayS ay Jo 
To[JUV FfoT JO MOTA LOMoJU-O1DjUW | ‘szivd potOysod oYvoIPUI SOUT] poyoq ‘weeq oy} Woy SuIsulids 9uTy pug OU SI a104} PUB poXtOJ SI auIy MOIq Oy, “(UIqnd 
“snl “Y@N) yjeg jo smbaeyy oyg Aq pojuosoad ‘(ueySeuoy_) ssorovmyorey rieou ‘yorpseN YSnoT] Wo WOJoPoYS oY} JO Jo[JUP qf] JO MoLA LOLLoyUI-O1ayUW W—"g “DLT 


z Px) 
PN So Z 
@ — o 
= a —— Fe * t 

a SSN x 
a. F rN ~~ SY 

{ 72) \ Ss a Ga 
~ : Y SL LE >. = 

-—--- =a - 
YA \ = ar 
y \ — tip 


—* -\ ae oe la" | 
_ =\ 


LEE S \ 
A 1 \ 
f, \ \\ 


28 


SS 
—— 
Cn, 


j : 
i 
‘id yi 
=. , 
ne a 
Sth 
y) 
Lop 
2p 


EL 


26 PLEISTOCENE MAMMALIA. 


Fic. 9.—The antlers shown in figs. A, C and D are all from Ireland, but no further information as to the locality 
is available. A Antero-inferior view of the antlers at the College of Science, Dublin. The right brow tine is remark- 
ably flattened, the left back tine is forked. The second tine springs from the commencement of the beam. B Anterior 
view of the antlers belonging to the skeleton from Lough Gur, Limerick, mounted in a crouching attitude in the 
Geological Museum of Trinity College, Dublin. The right brow tine is bifurcated, the left is terminated by three 
short points. C Antero-inferior view of the antlers of the skeleton formerly belonging to Sir Edmund Loder, Bart., 
and now at Selehurst, Horsham (Capt. W. W. Otter). The 2nd tine of the left antler is of exceptional length, and 
those of the right antler tend to arise in pairs from broad outgrowths of the palm. D Anterior view of antlers formerly 
belonging to Sir Edmund Loder, Bart., and now at Selehurst, Horsham (Capt. W. W. Otter). The right antler has 
scarcely any indication of a brow tine, and in both antlers the 2nd tine (d), which is very short, springs close to the 
tines of the beam, which are exceptionally long. 


27 


THE GIANT DEER. 


(61d ,“te0q Ys4tg , Stel Ul pemnstq) *per0jser yonuz ore wayed 044 SueurMs1e, SMOYS UOZRIZSNITIE O44 YOIYA souly Josie joy} JO [eIaAes ynq ‘wed ay} Jo sovy IOLIOjUT 
oy} Woy Sutids suoryoolord 410ys [etoAeg *poyeorpul Ajeoreos st (a) ouly Yowq oY, “poyeoangziq si euO FYDII oy} pue eseq peoasq AT[eMIOUGe UR MOI] SutIds (p) Seu pug 
ouE (19990 “M “A “3d¥p) wreysioy “Gsanyojag ye Mou pue “queg ‘toporT punMIp| IG 07 uygqng jo smeTiA, ‘sassoyy Aq pjos ,, puvfory ,, MOI; S1o[}Ue [eUIOUGE JO 
MOIA JUOTT CE ‘szurod 410ys XIs ynoqe jo solios @ SulTeeq ul [euIouqe Area st wed oY Jo oovzAns IJol0ysod ayy, ‘peyeommFlq Suteq ut [euI0Uqe SI UIP Oya £y30q OF 
WOMIUIOD ose JOYS B WHOI; OSIIe pIg puv pug oy} ‘wped oY} Jo ddvF JOIIOZUY OY} MOIS SuIBUIAds soul] SUT O44 JQ ‘“[eMA0U a1e Sout} pug pu Ao’q ayy, *(SIYo “A 1034") 
uuog jo zquRIy “Iq OF uqnd jo smerIA ‘sassoyy Aq poyddns puvjoty worz sie}Ue Jo SMOIA AJOWOJUI-O1NjJUY Q “Jo]JUB Yove ur (jy) OUT pUZ ayy JO WoryRoINgIG 
pue suruozyey oyg Loz o[qeyreUdod St UuToods oY, “Ul[GuC 479] UoumIoeds oY} orojoq usa sem Ydevasojoyd oy, “Yorn jo AqIsseATUQ oy} JO UNeSN]Y [BOIFO}OOZ eyy 
UL MOU YOMOULIT Wo1f WOYZO[OHS OY} JO S1o]ZUV OY JO MOIA QUOI | ‘Lopsoq 10110980d 841 spavSoer se AT[er1oodso ‘1v[NSodIII SI 1O}JUR Ajo] OY, “sinquIv_ jo Surmumoely “Hf 
07 wry Aq pure ‘uuog jo z7yURIY, “WT “Iq OF UN[GNG JO UoOg w sMTeNIIA ‘sussoyy Aq poyddns ,, purjory,, Wor; sIOUY YW ‘“siopJUe [eUTIOUGE yeyMomIOgG—"YT “OI 


28 PLEISTOCENE MAMMALIA. 


true Deer in the absence of the brow tine and in the expansion of the branch 
corresponding to the bez. They further differ from those of Megaceros in the fact 
that it is from the anterior part of the palm or crown that the tines spring. 

In the Reindeer the antlers agree with or rather exceed those of Megaceros in the 
length and flattened or branched character of the brow tine. The beam, which is 
greatly branched, differs from that of Megaceros in showing no tendency to become 
flattened. 

The Fallow Deer agrees with Megaceros, of which it is the nearest living ally, in 
the expanded character of the beam, and in the fact that the tines spring from the 
posterior face of the beam. It differs from most specimens of Megaceros' m the fact 
that the brow tine is cylindrical, not flattened. It further differs from the Irish race 
of the Giant Deer, and agrees with some of the continental races in having the antlers 
directed upwards and forwards instead of extending horizontally. 


(d) Dimensions. 


Very numerous measurements of the antlers have been published, especially by 
Rowland Ward, Millais and Potter ; and certain measurements obtained from various 
sources are quoted in the descriptions of some of the antlers figured. Only a general 
value, however, can be attached to such measurements, partly because unless they are 
all taken by the same person it is very difficult if not impossible to secure absolute 
consistency, partly because certain measurements, particularly of the span, may 
largely depend on the amount of “ restoration’ to which the tines have been sub- 
jected. Miss D. M. A. Bate kindly allows me to quote the following measurements 
made of specimens in the British Museum : 


M©EASUREMENTS OF ANTLERS OF CERVUS GIGANTEUS BLUMENBACH IN THE 
British Musrum (Naturat History), In Frnt anp INCHES. 


| | 
Circumference | 
“4 : Greatest length of : : : F 
Brit. Mus. No. Maximum span. AGG BHO, demme Lately abexe Left antler. | Right antler. 
On mounted skeleton | 8’ 112” R. 5’ 5” 10” | Tipsof2tines | 3 outertines | 
(? 15282) Treland | i Be IBY | gi” | restored restored 
15602, b, Ireland : 83% Tk, ay | 114” | Complete —- Extreme tip of 
ie By OR? | 104” 1 tine restored. 
M. 2324, Ireland .| 6/ 1” | Ry EY OP | 10” 3 Ditto. | 
| I, fay’ 10” 
| 8085 ‘ Al 9’ 74” | Re oni gi” | Tip of 4 tines Tips of all tines 
| L. 5° 63” gh” restored | restored. 
| 
| M. 2323, Ireland : GORY | R. 6° 33” | 10}” | Complete | Extreme tips of 
L. 6’ 9” 103” 3 tines restored. | 
| | 


* The relatively few specimens with a cylindrical brow tine seem to be in the main young individuals, 


THE GIANT DEER. 29 


lla. 


Fra. 11.—Abnormal antlers. A Antlers from Knocklong, Limerick (No. 83, Nat. Mus., Dublin), seen from above. 
In each antler the 2nd tine (d) springs close to the first tine of the palm. No back tines occur. ‘The left antler is 
relatively normal, but the right antler has the greater part of the anterior border ending in a straight, slightly irregular 
edge instead of being drawn out into tines. B Anterior view of antlers from Lough Gur, Dublin. This abnormal 
specimen, now in the National Museum, Dublin, was figured by V. Ball. Brow and back tines are absent, but in 
other respects the right antler is normal. ‘The blade or palm of the left antler is described by Ball as bifurcated. The 
smaller part, which may represent the 2nd tine, is itself bifurcated, one of the branches showing further subdivisions. 
The larger part has no tines on its anterior face, but terminates in a series of eight points or short tines. C Front 
view of antlers belonging to J. Purefoy Poe, of Harley Park, Callan, co. Kilkenny, and believed to have been found 
in the neighbourhood. The brow tines (c) are remarkably forked, and in the right antler the 2nd tine (d) is long and 
branched and abnormal in arising close to the brow tine. There is a slight indication of a left back tine (e). D Left 
lower view of antlers (No. 79, Nat. Mus., Dublin) from Tipperary. ‘The left antler is very abnormal, particularly as 
regards the length, slenderness and sharply bent character of the beam. There is a small brow tine (broken off) 
but no back or 2nd tine. The right antler, in which the beam is normal, has a second tine (d), but no brow tine, 


30 PLEISTOCENE MAMMALIA. 


Fra. 12.—Antlers of early forms of Giant Deer. a, Cervus (Huryceros) italie ; after Gastaldi, seen from in front. 
b, Megaceros (Huryceros) germanic, Pohlig; from Knielingen (Landessammlung fiir Naturkunde, Karlsruhe); after 
Frentzen and Speyer, seen from below. c, Cervus belgrandi, Harmer; from the Forest Bed, Pakefield, near Lowestoft 
(British Museum), after Harmer, seen from behind. d, Cervus (Megaceros) mosbachensis, Soergel; from Mosbach 
(Naturhistorisches Museum, Mainz); after Soergel, seen from above. 


THE GIANT DEER. 31 


Owen mentions that a skull and antlers in the Museum of the Royal College of 
Surgeons in London weighed 87 lb., and that the average weight of the skull without 
the antlers or mandible is 5 lb. This gives 81? lb. as the weight of the antlers—an 
astonishing amount of bony material to be produced annually and discarded. 


(e) Classificatian of Deer Based on the Antlers. 


Brooke’s classification of the Cervide based on the structure of the manus and 
pes was universally accepted till Cameron' in 1892 proposed to replace it by one 
based on the antlers. He divided the Cervide into three sections : 

1. For the Reindeer with antlers in both sexes. 

2. For the Elk with laterally extended antlers. 

3. For the remainder with erect or suberect antlers found only in the male. This 

group he further subdivided into— 

(a) A group including the typical Old World Deer and the Wapiti, with a 
normal brow tine springing from the outer side of the beam, and— 

(b) A group including the typical New World Deer (except the Wapiti), the 
Roe and one or two others. In these the first tine 1s considered not 
to be comparable to the brow tine as it arises from the inner side of 
the beam. 

Cameron’s classification is accepted by Lydekker. 

Pocock, from a study of the antler development in an American Deer (Odocoileus), 
shows that the first tine arises externally, but gradually assumes an apparently more 
internal position as growth proceeds. Hence he concludes that the term “ brow tine”’ 
is equally applicable in Old World and American Deer. 


c. DentiTIon (Plate II, and Text-figs. 13, 14). 


The teeth in the Cervide, as in all Ruminants, are very uniform in number, and 
those of the Irish Giant Deer conform to the usual formula of— 
ona, = pm. Serer: 2 
3 I 3 3 
The molar and premolar teeth are much less markedly hypsodont than in cattle, 
but have the normal selenodont grinding surfaces characteristic of Ruminants. Owing 
to their large size they are liable to be confused with those of Bos, but Owen points 
out the useful distinguishing feature, that in both upper and lower molars a small 
accessory column on the inner side in the groove between the two main lobes of the 
tooth is confined to the base of the groove in Megaceros while in Bovide it is much 
longer. 
1 « Field,’ 1892, pp. 265, 703, 741, 860. 


32 PLEISTOCENE MAMMALIA. 


Upper Dentition. 


The premolars and molars form a continuous series (Pl. II, fig. 1); all the 
members of which are similar in character, but the premolars are little more 
than half the size of the molars. The character of the grinding surface is better 
realized from figures than from descriptions, but its peculiar irregularity depends 
primarily on a dorsal infolding of the enamel, which after wear forms an 
irregular depression sometimes partly occupied by cement and surrounded by an 
enamel rim. External to this rim is the dentine forming the bulk of the tooth, while 
finally the prominent outer border consists of enamel. In the premolars the infold 


Zz 


ZZ be 


SSS = 


SS 
== 
= : 
IN ~~ z 
= & Se 
TVWAXw a 


Fic. 13.—Anterior end of mandible with incisor and canine teeth. A Left side view; B seen from above; half 
natural size. From Mullingar (Nat. Mus., Dublin). a, mental foramen for passage of a branch of the 5th nerve. 


of enamel has the posterior portion nearly or completely isolated from the remainder 
according to the state of wear of the tooth. In the molar teeth a sharp infold of 
enamel from the inner side greatly modifies the whole grinding surface, and gives the 
dorsal infold the character of a double crescent. The molar teeth have four roots, 
the premolars two on the inner, but one on the outer side. 


Lower Dentition. 


The three incisors and the incisiform canine have long tapering roots and expanded 
chisel-like blades, which together form a continuous cutting edge. The upper surface 
of the blade is marked by a depression. The teeth project conspicuously forward, 


THE GIANT DEER. 33 


and a considerable part of the root is not enclosed in the alveoli in the specimens 
examined. 

There is a gap of eleven or more centimetres between c. and pm. 2, pm. 1 being 
absent. The premolar and molar teeth steadily increase in size throughout the series 
(Pl. II, figs. 2—4). While in the molars a deep transverse groove on the outer side of 
the tooth meeting a less conspicuous one on the inner side almost bisects the tooth, 
the premolars have no such conspicuous grooving. The enamel infoldings into the 
crown of the tooth are crescent-shaped in the worn molars, but not in the premolars. 
The last molar has a conspicuous third lobe, and in all the molars a small cusp springs 
just above the alveolus on the outer side of the tooth. 


Fre. 14.—Milk dentition. A Inner view of left maxilla showing a milk tooth (m. pm. 3) in addition to the permanent 


dentition. From Carlow. B Left lower third milk premolar. From Doneraile. Both half natural size (Nat. Mus., 
Dublin). 


Milk Dentation. 


This is imperfectly known, but a maxilla (‘Text-fig. 14a) in the National Museum, 
Dublin, has m. pm. 3 just about to be shed, and a second fragment has m. pm. 3 
(Text-fig. 148). The former specimen is from Carlow, the latter from Doneraile Cave. 


Dp. THE VERTEBRAL COLUMN. 


Cervical Vertebra.—In the male these are distinguished by their great size and 
massive character in correlation with the weight of the antlers. The cervical vertebra 
of the female, in correlation with the absence of the great antlers, are considerably 
smaller than those of the male. 

The atlas (Text-fig. 154 and B) has specially deep articulating surfaces for the 
condyles of the skull, and rather short but massive transverse processes thickened 

5 


34 PLEISTOCENE MAMMALIA. 


“| 
Z Boars 
7a \ Af . es 
SS Sah RAE f y 
SENG GF ih Ls IY 
SS. tt i y 


\ 
y 


i G a e 


\\ 7 
\ 
A Jha 


4 


48 


Fic. 15.—A Atlas seen from above; B atlas, front view; © axis, front view; D axis, left side view; } natural 
size. From Rockbarton, Kilmallock, Limerick (Nat. Mus., Dublin). a, neural spine ; 5, neural canal; c, surface for 
articulation with condyles of skull; d, odontoid process ; e, transverse process ; f, surface for articulation with atlas; 
g, post-zygapophysis. 


Fia. 16.—Cervical vertebre. A 5th, front view; B 5th, posterior view ; C 6th, right side view; D 7th, right side 
view. All } natural size. All from Rockbarton, Kilmallock, Limerick (Nat. Mus., Dublin). a, neural spine; ), 
neural canal; c, prezygapophysis: d, postzygapophysis; e, transverse process; f, inferior lamella of transverse 
process ; g, vertebrarterial canal; h, anterior face of centrum ; 7, posterior face of centrum. 


to 


THE GIANT DEER. 35 


at the ends. The foramina on the upper surface for the passage of the spinal 
nerves are very conspicuous. The azis (Text-fig. 15c, p) differs less from the other 
cervical vertebree than is usually the case. It has the usual scoop-like odontoid 
process characteristic of Ruminants. The neural spine is not very high. The remain- 
ing five cervical vertebrae (Text-fig. 16) are much alike. The neural spines are short, 
particularly in the case of vertebree 3 and 4; the transverse processes give off a strong 
projection anteriorly, which in the case of vertebrae 3 and 4 is forwardly-directed ; 
in 5, 6 and 7 it becomes more massive and is directed downwards and forwards, and 
becomes an inferior lamella. 

All the measurements of vertebrze given in the following tables are of the male, 
unless otherwise stated. 


MEASUREMENTS OF THE CERVICAL VERTEBR®. 


* Measured from centre of anterior convexity to centre of posterior concavity. 


Female skeleton from Lough Gur, 


Limerick (Nat. 


., Dublin). 


Mus 


Skeleton from | & 
Set M. 2971 mines No, 254, 1904 
Brit. Age _Evniscorthy (Nat. Mus, 
Gamitecicos, |) \Sedewick Muss Dublin), 
oe ‘Ts Cambridge). ‘ 
; iat Pes) ee ey =e eee Gl | 
ATLAS. | 
1. Maximum width . : : : } 21-2 22:5 23-2 
2. Median dorso-ventral diameter a 9°3* 9-3 8-75 
3. Extreme width of condylar articular 
surface . : 5 : ; c 10-8 11-05 
AxIs. 
1. Length from anterior end of odontoid 
process to postero-ventral extremity of | 
centrum . : ; : 2 | 15-4 no | 15-9 
2. Height from base of centrum to top of | 
neural spine. : 6 . . 207 13-5 15-35 
3. Diameter across transverse processes : 16:8 17-5 19-6 
4. Diameter across articulating surfaces for 
atlas ; : : 6 : ; 10-6 11-55 | 11-8 
5. Diameter across post-zygapophyses 8-15 9-9 | 10-45 
* From end of spine to end of hypapophysis. 
MEASUREMENTS OF CERVICAL VERTEBRA. 
3rd cervical, 4th cervical 5th cervical, 6th cervical. 7th cervical, 
Pie | ee es Sma rae 4 | 
} 3 a = S = | | = | | 
-| 2 can ) o- Cy ibys | oll | to) | 
@| BS\e8 | Feo| aa e| 8a/ae Wo eae & 
He | ss eo | A; = Bo} acs | so ey) She 5 4: 
42 \86 Sel Sa \sen. | ne | xe les EeE| 2e | SS]. a 
a 3A do] 2 85 BA |e -f= ae oA gq | 2 35 SE's Sle 2 
a 26\|C214 re |e | Ole laa | aa | OBS eel ey |e Zi 3 
cs AS eat s a% | BS |S]; a9 | ga |g] . a~| SE |S]. | 
Se iar | etl | at (SL eR hea as Sp Sy cers eis Peet a WSS It ceiect |) iat she 
9 Ms |/S8iaqg2 ge/¥%s|/Stima!] oo) 45 |/88laeles| Mu |Selme 
-z | 25 fey tabetha ee ~2 |oa| -& | 270.9 
l= By Aq —-5 ile ay oStao| £m EY of) 5 | x By | oO? 
S 5H | SS ue, || SSeS tS I) caer Ssh ee Ro |H#4| oC |x So 
a fa |@¥S0 go) fa | PKR Oo] po | oa | BSP Sol] ee] ow | t KHIR 
me eS i/o2I"4 Se) go) oSl Al se} so) oet ARISE] zo | oeTH 
‘S. mo |2ol oes | o%| ya |esles| oo] em | eof ss jot) 2A |e olas 
a OC St = Tt) (pene raed o- | Sal 55! o2 mec Fetal | ero eo EGY priae 
> CH | Oat ay | HB Of | Oca | zy CH | Del os |*¥n] SH | Ob aKS 
mM Ga |a4ine n fn |24inm—-!| no Sn | SAP n> | N—| On | Ain 
. Width across trans- |17°25,19°5 19°7 |13°3)17°2 19-9 |19°8 |14:0.19°3)20°1 |20°6 |14:0)16-0 |17-9)18°25 11°3)19°5 16°6 15-1 134 
verse processes | | | | 
. Width across post-| 9°7 /10°9)10:05 T7101 hanna 7710-0. 10°75)10°20} 7°5)10°25)11°3)11°15) 9:0) 98 LOT 1015 84 | 
zy gapophyses | | | | 
. Length of centrum* 105 | .. 10-4] .. 10°05, «. (O75, 2.97) .. (LUT) ..] 9:6 | .. 105) .. 100, ..) 9:36 
| | | 
| | | 


36 PLEISTOCENE MAMMALIA. 


Fie. 17.—Thoracie vertebre. A Ist, front view; B 3rd, right side view; C 11th, right side view; D 15th, 
front view; all } natural size. From Rockbarton, Kilmallock, Limerick (Nat. Mus., Dublin). a, neural spine ; 
b, neural canal; c, prezygapophysis; d, postzygapophysis; e, transverse process; f, surface for articulation with 
head of rib; g, surface for articulation with tubercle of rib; h, anterior face of centrum. 


Fia. 18.—Lumbar vertebrae. A 2nd, right side view; B 2nd, seen from above; © 5th, seen from in front ; 
4 natural size. From Rockbarton, Kilmallock, Limerick (Nat. Mus., Dublin). a, neural spine; 6, neural canal ; 
c, prezygapophysis; d, postzygapophysis; e, transverse process; f, anterior face of centrum; g, posterior face of 
centrum. 


ee eee 


THE GIANT DEER. om 


Thoracic Vertebre (Text-fig. 17).—These number thirteen. The spines increase in 
length from 1 to 3. Vertebrae 4 and 5 have very long spines, but in the succeeding 
vertebrae the length of the spies rapidly decreases. The spines of vertebrae 1-11 
are backwardly-directed to a greater or lesser extent. Those of vertebrae 12 and 13 
are practically vertical and resemble the spines of lumbar vertebrae. The centra 
throughout are short, nearly cylindrical and very uniform in length. 


MEASUREMENTS OF THORACIC VERTEBRA. 


(All No. 254 [1904] from Rockbarton, Kilmallock, co. Limerick, in the National Museum, Dublin.) 


| Ist. 4th. 7th. 9th. 11th. 14th. 


| 

| 1. Height from notch between pre-zygapo- | 
| physes to top of neural spine . : 21-8 36:6 29-0 21-5 20:3 15-4 | 
| 2. Width across transverse processes : 14-25 10-5 10-9 10-3 10-7 8.5 | 
_ 3. Width across pre-zygapophyses . 2 10°5 8:5 7-0 6:5 6:8 | 


MEASUREMENTS OF LUMBAR VERTEBRA. 


(Details as with thoracic vertebre.) 


1st 3rd 5th 
| 
1. Length of centrum : : ‘ dell 7:1 7:25 
2. Measurement across transverse processes 17-6 25:0 27-0 
3. Height from mid-central point of centrum 
to top of neural spine : ; ; 16-25 16-15 15:3 
| | 


MEASUREMENTS OF SACRUM. 


No. 254, 1904, 

No, 15, 1915, Rockbarton, 
Dublin. Kilmallock, 
co. Limerick. 


1. Length of sacrum. : : : 23-2 25-85 
2. Height to top of neural spine of 2nd 

true sacral . F pi Fi : 10°8 12-9 
2. Measurement across transverse pro- 

cesses of Ist sacral 22:5 24-0 


Lumbar Vertebre (Text-fig. 18).—The five lumbar vertebree all have the neural 
spines abruptly truncated, and the centra very uniform in length. The transverse 
processes are long and straight, taper slightly towards their terminations, and are as 
a rule directed slightly forwards. 

Sacral and Caudal Vertebre (Text-figs. 19, 20).—There are two sacral and three 
pseudosacral vertebra. Both sacral and pseudosacral have rather large neural spines, 


38 PLEISTOCENE MAMMALIA. 


Fic. 19.—Sacrum. A Right side view; B lower surface; 4 natural size. From Rockbarton, Kilmallock, 
Limerick (Nat. Mus., Dublin). a, articular surface for ilium; 5b, anterior face of centrum of Ist sacral vertebra ; 
c, gaps between partially coalesced neural spines of sacral and pseudosacral vertebre ; d, neural spine of last pseudo- 
sacral vertebra; e, foramen for exit of spinal nerve 


Fia. 20.—Free caudal vertebre. A The first; B the second; C the third, (1) seen from above, (2) front view. 
Drawn from casts in the National Museum, Dublin, of the original specimen from Lough Gur, Limerick, now at 
Trinity College, Dublin. % natural size. a, anterior surface of centrum; 6, neural spine; c, prezygapophysis ; 
d, transverse process. 


THE GIANT DEER. 39 


the tops of which merge into one another. The short transverse processes also merge. 
The anterior tail-vertebree have only rarely been obtained and the terminal vertebre 
have apparently never been found. 


E. THE Rips AND STERNUM (Text-fig. 21). 


There are thirteen pairs of ribs, the anterior being rather wide and flattened. 


== - ; Sf =. y) 
( 3 S i 


) 


5 Mine 

ye NO es 

Na eg 

<7! / of 
he . 

Se Gal] t 


» 


Z ? 
gens 

ae a 

We 


oe Zu 


z a 


Fic. 21.—Sternum of the female skeleton from ‘‘ Ireland” set up in the British Museum (Nat. Hist.). 4 natural 
size. a, presternum; 4, first sternebra of mesosternum; c, last sternebra of mesosternum; d, xiphisternum; e, 
first sternal rib; f, last sternal rib. The figure as drawn is somewhat foreshortened. 


F. THE SHOULDER GIRDLE (Text-fig. 22). 


The scapula has a well-marked spine, but the acromion is not prominent. The 
outline is nearly that of an isosceles triangle, as in Ruminants in general. 


MEASUREMENTS OF THE SCAPULA. 


Skeleton from Enniscorthy Rockbarton, Kilmallock, 
(Sedgwick Mus., Limerick, 254, 1904 
Cambridge), (Nat. Mus., Dublin), 
| 
1. Length along line of spine ; 45:02 49-3 | 
2. Maximum width : ; : 28:6 27°3 
3. Width at neck. ; ; : 7:65 7-4 


40 PLEISTOCENE MAMMALIA. 


Fic. 22.—Left scapula seen from outer side. 4 natural size. 


From Rockbarton, Kilmallock, Limerick (Nat. 
Mus., Dublin). 


a, glenoid cavity ; b, spine ; c, coracoid process; d, coracoid border; e, glenoid border ; f, acromion. 


G. THe Anterior Lime (Text-figs. 23—26). 


In the humerus (‘Text-fig. 23) the tuberosities and deltoid ridges are not very 
prominent. The great tuberosity is blunt and not much inwardly-directed. The 
condyle is strongly marked, and the anconeal fossa is deep. There is no supratrochlear 
foramen. 


THE GIANT DEER. 4] 


The radius (Text-fig. 24) is a large bone having the form commonly found in 
swift-footed ungulates. The ulna has a large olecranon, but the lower half, which is 
firmly united to the radius, is reduced to a thin splint-like bone. 


i, / fh fi 4 
pr vif ly 


3 CS 
> ae | 
' 

MA 


¢ aN 


i 2 Ce 
Wins ‘ 
ns 
\ 

\\, 


SS 


Ss 


Otc, vi \) Ss 
/ 


e 


Z 

Y 

XS RSS 
~ 


Fic. 23.—Right humerus. A Antero-internal view; B Outer (right side) view. 4 natural size. From Rock- 
barton, Kilmallock, Limerick (Nat. Mus., Dublin). a, head; b, great tuberosity ; c, lesser tuberosity ; d, deltoid ridge ; 
e, trochlear surface ; f, external condyle; g, internal condyle. 


The carpus (Text-fig. 254) has the arrangement characteristic of the Ungulata 
vera. There is no centrale ; the magnum articulates with the scaphoid and is separated 
; g | | 


from the cuneiform by the meeting of the unciform and lunar. 


All the carpal bones 
6 


42 PLEISTOCENE MAMMALIA. 


strongly interlock. The manus has the normal Ruminant feature of the union of 
metacarpals 3 and 4 to form the cannon-bone. The Irish Giant Deer belongs to the 


ie an 


Ms SN 
{ fy) ig 

Yh 4 
i} ip 
! Y 


itt 


| 


iy 
I if i | 
Ni 


| 


Fie, 24.—A Left radio-ulna seen from right side; B the same seen from in front; } natural size. From Rock- 
barton, Kilmallock, Limerick (Nat. Mus., Dublin). a, olecranon; 6, surface for articulation with trochlea ; G 
proximal end of radius; d, distal end of radius; e, distal end of ulna. 


plesiometacarpal section of the Cervide, in which the proximal part of the second 
and fifth metacarpals persists in the form of a short splint. Distally the second 
and fifth digits are represented by the full series of three small phalanges. 


THE GIANT DEER. 43 


Fra. 25.—A Left manus seen fromin front, and B left pes 
antero-lateral view of the female skeleton from “ Ireland” 
in the British Museum (Nat. Hist.); } natural size. Aa, 
radius ; b, scaphoid ; c, lunar; d, cuneiform; e, magnum 
trapezoid; f, unciform; g, 3rd and 4th metacarpals fused ; 
h, proximal phalanx of 3rd digit; 7, ungual phalanx of 4th 
digit. B k, tibia; 1, fibula; m, caleaneum; 1, astragalus; 
0, navicular portion of naviculo-cuboid; p, cuboid portion 
of naviculo-cuboid; qg, external cuneiform; 7, 3rd and 4th 
metatarsals fused; s, proximal phalanx of 3rd digit; ¢, 
ungual phalanx of 4th digit ; w, sesamoid bone. 


HuMmERUvS. 
1. Extreme length : . : : 
2. Maximum transverse diameter at distal 


end - . c : - ; 
3. Antero-posterior diameter of shaft at 
highest point in deltoid ridge 


XADIUS. 
1. Extreme length 
2. Width at proximal end 
3. Width at distal end 


ULNA. 


Fie. 26.—A Left manus and B left pes, of the skeleton 
from Ireland in the Museum at Selehurst, Horsham (Capt. 
W. W. Otter), antero-lateral view. a, 3rd and 4th meta- 
carpals (or metatarsals), fused; b, vestige of proximal end 
of 5th digit ; c, sesamoid bone; d, ungual phalanx of 5th 
digit; ¢, proximal phalanx of 5th digit. 


TABLE OF MEASUREMENTS. 


| 
Skeleton from Enniscorthy Rockbarton, Kilmallock, | 
Sedgwick Mus., Limerick, 254, 1904 
Cambridge) (Nat. Mus., Dublin) 
7 | 
41-35 40-3 
10-15 | 9-8 
6-9 9-65 
| 
37-7 | 38-45 
10-2 | 10-05 
10-05 9-05 
48-7 47-2 


1. Length (not measured along curvature) 


44 PLEISTOCENE MAMMALIA. 


THE MANUS. 


Skeleton from Enniscorthy 
(Sedgwick Mus., Cambridge). 
| 
| 1. Length of cannon-bone (metacarpals 3 and A 31-4 
| 2. Transverse measurement at proximal end 
of cannon-bone : 7-1 
3. Transverse measurement at distal end of | 
cannon-bone . “| 74 
4. Length of Ist phalanx, measured in middle | 
of outer side. ee 
5. Length of 2nd phalanx, measured in middle. 
of outer side . | 4-9 
6. Length of ungual phalanx, measured ‘along | 
cuter edge. : . : : 2 7-1 


H. THE PELVIC GIRDLE (Text-fig. 27). 


The pelvis has the somewhat narrow form characteristic of Ruminants. The 
ilium is widely expanded anteriorly, but narrows much in front of the acetabulum. 
The pubis is a small bone. There is a long symphysis ischii. The ischial tuberosities 
are very prominent, as is usual in Ruminants. The obturator foramen is oval and of 


moderate size. 
MEASUREMENTS OF THE RIGHT INNOMINATE BONE. 


Skeleton from Rockbarton, Kil- | 
Enniscorthy mallock, Limerick. | 
‘Sedgewick Mus., 254, 1904 
Cambridge). (Nat Mus., Dublin) 
1. Maximum leneth . 51:0 55-0 
| 2. Length from middle of edge of acetabulum to 
middle of supinator border of iliam : : 24-0 25:5 
| 3. Width of expanded portion of ilium. : ‘ 20:0 24-2 
4. Dorso-ventral measurement of ilium at nar- 
rowest point in front of acetabulum . : 5-4 Fel 
| 5. Length of symphysis . : : : 19-9 19-25 
a6 Dorso- ventral measurement “of ischium at 
narrowest point behind acetabulum : : 6-6 6:0 


Fra. 27.—Pelvis. A Seen from below. B Left side view. C Seen from above. +} natural size. From Rock- 
barton, Kilmallock, Limerick (Nat. Mus., Dublin). a, acetabulum; 6, obturator foramen ; c, gluteal surface of ileum ; 
d, iliac surface of ilium; e, ischial tuberosity ; /, pubic symphysis; g, ischial symphysis. 


The femur (Text-fig. 28) has very slight indications of a neck. 


is no 3rd trochanter. 


bent. 


THE GIANT DEER. 


1. THE Posterior Lime (Text-figs. 28—30). 


45 


The great 


trochanter is very large, the lesser very small, and as in other Artiodactyls there 
The upper end of the tzbva (Text-fig. 29) is characteristically 


In the character of the digits the pes (Text-fig. 258) is analogous to the 
manus. The astragalus has both proximal and distal surfaces pulley-hke as in other 


Artiodactyls. 


TABLE OF MEASUREMENTS. 


Skeleton from 
Enniscorthy 
(Sedgwick Mus., 
Cambridge). 


FEMUR. | 

1. Length measured from head to internal condyle. | 

2. Transverse diameter at condyle : | 

| 3. Transverse measurement across head and great | 
trochanter ; ; : : i .| 

TrIpra. 

1. Length é . | 
2. Right to left diameter at proximal end . -| 

3. Antero-posterior diameter at proximal end | 


measured from notch between articular surface 
for femur to top of crest 5 
4, Right to left diameter at distal end . 


9. 


. Length of Ist phalanx me sasured in middle | 


Tue Pus. 

Length of caleaneum | 

Maximum transverse diameter of caleaneum| 
(oblique measurement) . 


. Right to left diameter of astragalus | 


measured at tibial articular surface 


. Length of cannon-bone 


Transverse measurement of cannon- epee | 
at proximal end . ail 


. Transverse measurement of cannon-bone | 


at distal end 
of outer surface . .| 
Length of 2nd phalanx me »asured in middle | 
of outer surface | 
Length of ungual phalanx measured along 
inner edge 


Rockbarton, Kil- 
mallock, Limerick, 


254, 1904 


| 
is Nat. Mus., Dublin). 


Skeleton from 
Enniscorthy 


(Sedgwick Mus., 


Cambridge). 


46 PLEISTOCENE MAMMALIA. 


Fie. 28.—Right femur. A Seen from behind. B Front view. 4 natural size. From Rockbarton, Kilmallock, 
Limerick (Nat. Mus., Dublin). a, head; b, great trochanter; c, lesser trochanter; d, internal condyle; e, external 
condyle; f, trochlear surface. 


THE GIANT DEER. 


Fic. 29.—Right tibia. A Front view: B seen from behind. 1 natural size. 


Limerick (Nat. Mus., Dublin). 


3 


47 


From Rockbarton, Kilmallock, 


a, surface for articulation with condyles of femur; , cnemial crest. 


48 PLEISTOCENE MAMMALIA. 


Fic. 30.—A Left astragalus seen from in front. a, surface (condyle) for articulation with the tibia; b, surface 
for articulation with the naviculo-cuboid. B Left caleaneum seen from in front. c, surface for articulation with 
the upper surface of the astragalus; d, surface for articulation with the outer surface of the astragalus. C Right 
naviculo-cuboid seen from above. e, surface for articulation with the astragalus; jf, surface for articulation with 
the caleaneum. All half natural size and all from ‘“ Ireland” (British Museum (Nat. Hist.), Nos. M. 2339, 18151, 


18038). 


VI. LIST OF IRISH GIANT DEER SKELETONS AND ANTLERS. 


The list of antlers preserved in England is by no means complete, while it is 
impossible to prepare an even approximately complete list of those in Ireland. Dr. 
H. Stokes of Dublin writes : “‘ There are many heads in private houses all over Ireland, 
I would guess several hundreds.” 


SKELETONS. 
A. In the British Isles. 
(1) In publi institutions : 

London, British Museum (Nat. Hist.), S. Kensington (2, one of them 
female) ; Royal College of Surgeons Museum. 

Edinburgh, Royal Scottish Museum. 

Dublin, National Museum of Ireland (4, one disarticulated); Trinity 
College (3) ; College of Science Museum. 

Belfast, City Museum. 

Birmingham, City Museum ; University Museum. 

Cambridge, Sedgwick Museum. 

Cardiff, National Museum of Wales. 


THE GIANT DEER. 49 


Glasgow, City Museum. 

Leeds, City Museum. 

Liverpool, City Museum. 

Oxford, University Museum (a second skeleton in store). 
Sheffield, City Museum. 

Tring, Zoological Museum. 

Warwick, Town Museum. 

York, Yorkshire Museum. 

Douglas Museum, Isle of Man. 


(2) In private possession : 
Marquess of Londonderry, Mount Stewart, co. Down; Mr. J. Gaisford 
St. Lawrence, Howth Castle, co. Dublin; Lord Powerscourt ; 
Dr. H. Stokes, Dublin; Col. Vernon, Clontarf Castle, co. Dublin ; 
Mr. A. Heber-Pe cy, Hodnet Hall, Salop; Capt. W. W. Otter, 
Selehurst, Horsham. 


B. On the Continent. 
In the Museums of Amsterdam, Basel, Berlin, Dijon, Frankfort, Hamburg, 
Lyons, Paris, Stuttgart, Vienna, Ziirich. 


C. In America. 

Berkeley University Museum, California. 

Chicago, Field Museum. 

Cleveland, Ohio, Museum of Natural History. 

Denver, Col. 

Harvard University Museum, Cambridge, Mass. 

New York, Columbia University Museum; American Museum of Natural 
History. 

Yale University Museum, Newhaven, Conn. 


ANTLERS. 
A. In the British Isles. 
(1) In public institutions : 
London, British Museum (Nat. Hist.), S. Kensington (6) ; Royal College 
of Surgeons Museum; Geological Society, Burlington House (2). 
Aberdeen, Marischal College. 
Birmingham, City Museum. 
Brighton, Municipal Museum (2). 
Bristol, City Museum (2). 
Cambridge, Sedgwick Museum. 
Derby, Municipal Museum (2). 
Dublin, National Museum, 32 skulls with antlers, 14 shed antlers. 


50 


PLEISTOCENE MAMMALIA. 


Eton College Museum (imperfect). 
Exeter, Royal Albert Memorial Museum. 
Halifax, Belle Yue Museum (one antler). 
Hull, Municipal Museum (2). 

Leeds, City Museum (2). 

Leicester, Municipal Museum. 

Maidstone, Municipal Museum (imperfect). 
Liverpool, City Museum (2). 

Manchester, University Museum. 
Penzance, R. Geol. Soc. of Cornwall. 
Oxford, University Museum (3). 
Reading, Town Museum. 

Saffron Walden. 

Sheffield, City Museum. 

Warrington, Municipal Museum. 


York, Yorkshire Museum. 


(2) In private possession : 


Duke of St. Albans, Duke of Westminster, Marquess of Londonderry, 
Earl of Besborough, Lord Sackville (Knowle, Sevenoaks), Lord Powers- 
court, Mrs. Donaldson Hudson, Mrs. Graham Lloyd, Mr. J. G. Millais, 
Mr. G. C. Whittaker, Sir Peter Walker, Capt. W. W. Otter, Selehurst, 
Horsham (formerly belonging to Sir Edmund Giles Loder, Bart.). 


The above are quoted by Rowland Ward. 


Sir G. Abercrombie, Forglen, Turriff, Aberdeen (2) ; Lord Rothschild, 
Tring (2); Miss J. Cunnmgham, Bute House, Brook Green, Hammer- 
smith ; Sir Ernest Wills, Littlecote, near Hungerford ; Capt. Desmond 
Buxton, Catton Hall, Norwich. 


The following is a very incomplete list of those in private possession in 
Ireland : 


Lord Castletown; Marquess of Cunningham; Marquess of Headford ; 
Marquess of Londonderry ; Royal Dublin Society ; Lord Dunraven ; 
Lord Dunsany ; General Franks, Ballyseadden, Limerick (2) ; Lord 
Holmpatrick, Abbotstown, Dublin ; Mr. Karkpatrick, Lucan, Dublin ; 
Mr. J. Gaisford St. Lawrence, Howth Castle, Dublin (2); Mr. J. 
Purefoy-Poe, Harley Park, Callan, Kilkenny ; Dr. H. Stokes, Dublin ; 
Capt. Perceval, Hazlewood House, Shgo; Mr. J. Walker, Rathcarrig, 
Sligo; Capt. Bruen, Oak Park, Carlow; Mr. E. Eustace Duckett, 
Castlemore, Tallow, Carlow ; Sir Arthur Ball, Merrion Square, Dublin. 


THE GIANT DEER. 51 


B. On the Continent. 


In the Museums of Berne, Bonn, Brussels, Buda-Pest, Darmstadt, Dijon, 
Dresden, Frankfort, Karlsruhe, Louvain, Mainz, Munich, Paris, Stras- 
burg, Worms. 


VII. CONCLUSION. 


This must take the form of the acknowledgment of the large amount of help 
which I have received. 

To the National Museum of Ireland, Dublin, through Mr. A. W. Stelfox, I am 
greatly indebted for the loan of a large collection of notes, pamphlets and illustra- 
tions which Dr. R. F. Scharff had assembled when in charge of the collection. To 
Mr. Stelfox I am also indebted for much personal help in the Museum. 

Captain W. W. Otter has most kindly helped in every way in the examination of 
the collection made by the late Sir Edmund Loder, Bart., and now at Selehurst, 
Horshan.. I have to thank him for permission to figure three pairs of antlers from 
Selehuxst, Lord Castletown for those at Granston Manor, Ballacolla, and Mr. J. 
Purefoy-Poe those at Harley Park, Callan. 

The Curators of nearly a'l the chief museums of the British Isles together 
with Dr. Revilliol of Geneva. Dr. Wysogorski of Hamburg and Dr. K. Hescheler 
of Ziirich, have kindly replied to inquiries concerning the specimens under their 
respective charge, and Dr. F. Corner has lent me specimens from his collection. Messrs. 
W. Williams & Son, of Dublin, the only exporters of Irish Giant Deer, have allowed me 
to reproduce several photographs taken prior to the specimens leaving their establish- 
ment. Dr. F. Krantz, of Bonn, to whom many of Messrs. Williams’s specimens were 
sold, has most kindly informed me as to their present whereabouts. Dr. Barnum 
Brown has given me information about specimens in American museums. 

Sir Sidney Harmer, Dr. K. Frentzen, Dr. K. Hescheler, Dr. H. Pohhg and Dr. 
W. Soer sel have all allowed me to reproduce illustrations from papers by them. 

Mr. C. Davies Sherborn and Mr. B. H. Soulsby have most kindly helped me with 
many points connected with the bibhography. 

Miss D. M. A. Bate, of the Natural History Museum, South Kensington, and Sir 
Arthur Smith Woodward, have helped me much in various ways. But above all I 
have to thank Dr. R. F. Scharff, our chief authority on the Irish Giant Deer, who has 
taken the keenest interest in the progress of this monograph. I cannot overstate my 
indebtedness to him. 

Mr. James Green has once more placed his artistic skill at my disposal in the 
preparation of the plates and many of the figures, while for the drawings of the Dublin 
specimens I am indebted to Miss Ii, I, Barnes, 


52 PLEISTOCENE MAMMALIA. 


VIII. BIBLIOGRAPHY. 


1. 1697. T. Molyneux, “ A Discourse concerning the Large Horns frequently found 
Under Ground in Ireland. Concluding from them that the Great American 
Deer call’d a Moose was formerly common in that Island,” “ Phil. Trans.,’ 
X1x, p. 485. 
2. 1726. J. Kelly, “ An Account of Fossil Horns Found Underground in Teland,” 
“hill amiss xxcxdve somes 
3. 1726. T. Molyneux, in G. Boate’s ‘ Natural History of Ireland,’ pp. 137—49. 
4.1731. L. D. Hermann,‘ . . . KHlends-thier-corper oder knochen 
in dem Masslicher Pfarr-garten-graben . . . gefunden.’ Budissin. 
5. 1746. T. Knowlton, “An Account of Two Extraordinary Deer’s Horns Found 
Underground in Different Parts of Yorkshire,” ‘ Phil. Trans.,’ xliv, p. 124. 
6. 1758. T. Wright, “ An Account of Fossil Moose-Deer Horns Found in Ireland,” 
‘ Louthiana or Introduction to the Antiquities of Ireland,’ Bk. iu. 
7. 1768. P. 8. Pallas, “ De Ossibus Sibiriz Fossilibus,”’ ‘ Novi Comment. Petropol.,’ 
xi, p. 468 (note). 
8. 1781. F. HE. v. Rochow, “ Nachricht v. emem ungewohnlich grossen, mit Stemrinde 
diinne tiberzogenen Geweih eines zu dem Hirschgeschlecht gehorigen 
Tieres,” ‘ Schrift. d. Ges. naturf. Freunde,’ Berlin, Bd. 2. 
9. 1781. T. Pennant, “ History of Quadrupeds,’ 1, p. 97, and pl. xi, 1. 
10. 1784. P. Camper, “Complementa varia . . . De Alcibus giganteis Hibernie,” 
‘Nova Acta Acad. Scient. Imp. Petropol.,’ i, p. 258. 
11. 1789. G. de Razoumowsky, “Sur une téte et des cornes fossiles d’Irlande,” 
‘ Hist. et Mem. Soc. sc. phys. Lausanne,’ ii, p. 27. 
12. 1811. J. Parkinson, * Organic Remains of a Former World,’ iii, p. 313. 
13. 1821. G. A. Goldfuss, “‘ Ueber den Riesenhirsch (Cervus giganteus),” ‘ Nova Acta 
Acad. Caes. Leopold-Carol,’ x (2), p. 455. 
14, 1822. A. G. Desmarest, “Mammalogie,’ ui, p. 446. 
15. 1823. G. Cuvier, ‘ Recherches sur les Ossemens fossiles,’ nouv. ed., iv, pp. 70—88. 
16. 1825. J. Hart, “ Description of the Skeleton of the Fossil Deer of Ireland,’ Dublin. 
17. 1825. H. R. Oswald, “ Observations Relative to the Fossil Elk of the Isle of Man,” 
‘Edinb. Journ. Sci.,’ ii, p. 28. 
18. 1825. S. Hibbert, “ Account of the Circumstances Connected with the Discovery 
of the Fossil Irish Elk in the Isle of Man,” *‘ Edinb. Journ. Sci.,’ iii, pp. 15—18. 
19. 1825. T. Weaver, “ On the Fossil Elk of Ireland,” * Phil. Trans.,’ exv, pp. 429-35. 
20. 1827. Hamilton Smith in Griffith’s Cuvier, v, p. 306. 


" Notwithstanding the length of this bibliography, it does not include all the literature dealing 
with the Giant Deer. | Other references will be found in the bibliographies of recent continental authors, 
particularly Hescheler (120). 


Ze 


22. 


23. 


24, 


25. 


26. 


27. 


28. 


29. 


30. 


31. 


32. 


33. 


34. 


35. 
36. 


37. 


38. 


39. 


40. 


4]. 
42. 


THE GIANT DEER. 53 


1830. 8. Hibbert, “ Additional Contribution towards the History of the Cervus 
euryceros or Fossil Elk of Ireland,’ ‘Edinb. Journ. Sci.,’ N.s., ii, 
DPa Ola: 

1836. J. Phillips, “ Rep. Phil. Soc., Yorks.,’ p. 13. 

1838. J. Hart, “ Observations on the Fossil Deer of Ireland (Cervus magaceros) ”’ 
(sec), ‘ Journ. Geol. Soc. Dublin,’ i, p. 20. 

1838. J. Scouler, “ Remarks on the Natural History of the Fossil Elk,” ‘ Journ. 
Geol. Soc. Dublin,’ i, p. 197. 

1842. H. Rathke, “ Uber den Riesenhirsch (C. megaceros),” ‘ Preuss. Provincialbl.,’ 
Bd. xxvu. 

1843. R. Owen, “ Report on British Fossil Mammalia,’ ‘Rep. Brit. Assoc.’ 
(Cork), p. 237. 

1845. C. E. vy. Hichwald, “Uber d. Riesenhirsche . . .,” ‘ Erman. Archiv. fiir 
Wissenschaft. Kunde v. Russland,’ v, pp. 158—75 (Moskau). 

1846. EH. Forbes, “ The Geological Relations of the Existing Fauna and Flora of 
the British Isles,” ‘Mem. Geol. Surv.,’ 1, p. 394. 

1846. J. G. Cumming, “ On the Geology of the Isle of Man,” ‘ Quart. Journ. Geol. 
Soc.,’ 1, pp. 344—5. 

1846. R. Owen, ‘ British Fossil Mammals and Birds,’ pp. 444—68. 

1846. H. D. Richardson, ‘ Facts Nat. Hist. Gigantic Irish Deer,’ Dublin. 

1847. C. G. Giebel, “Fauna der Vorwelt,’ 1, p. 145. 

1848. R. Owen, “. . . Fossil Remains of the Megaceros hibermcus : 
in the Pleistocene Deposits . . . at Ilford and Grays Thurrock, Essex,” 
‘Quart. Journ. Geol. Soc.,’ iv, pp. 42—6. 

1849. T. Oldham, “ On a Remarkable Group of the Remains of Deer Found near 
Kiltiernan,”’ ‘ Journ. Geol. Soc. Dublin,’ i, p. 252. 

1851. G. A. Mantell, ‘ Petrifactions and their Teachings,’ p. 455. 

1855. K. Peters, “ Kin Vortrag iiber den irischen Riesenhirsch,” ‘ Jahrb. k. k. 
geol. Reichsanstalt,’ 6 Jahrg. (ref. in Petermann’s Mittlg., 1855). 

1856. R. Owen, “ Description of Some Mammalian Fossils from the Red Crag of 
Suffolk,” ‘ Quart. Journ. Geol. Soc.,’ xii, p. 226. 

1858-71. E. Cornalia, ‘‘ Mammiferes fossiles de Lombardie,” * Pal. Lomb.’ (ed. 
Stoppani), pp. 54-64. 

1858. A. Carte, “ Description of the Fossil Bones Discovered by Mr. Brennan at 
Shandon near Dungarvan,” ‘ Journ. Roy. Dublin Soc.,’ 1, p. 351-7. 

1859. H. Denny, ‘“‘ On the Claims of the Gigantic Irish Deer to be Considered 


as Contemporary with Man,” ‘Proc. Geol. Polyt. Soc. Yorkshire,’ ui, 
p. 400. 

1859. J. MacKnery, ‘ Cavern Researches,’ pl. 1. 

1859. W. R. Wilde, “. . . Animal Remains belonging to the Academy,” 


‘Proc. R. Irish Academy,’ vu, p. 195. 


54 


43. 


44, 


45, 


46. 


47. 


48. 


49. 


50. 


ole 


52. 


60. 


1861. 


1861. 


1864. 


1864. 


1861. 


1865. 


1867. 


1867. 


1867. 


1869. 


. 1869. 


5 WES 


PLEISTOCENE MAMMALIA. 


R. Howse, ‘“‘ Notes on the Fossil Remains of Some Recent and Extinct 
Mammalia Found in the Counties of Northumberland and Durham,” ‘ Trans. 
Tyneside Nat. Field Club,’ v, p. 117. 

H. B. Geinitz, ‘“‘ Ueber den Riesenhirsch des Dresdener Museum,” ‘ N. 
Jahrb. f. Mineral.,’ pp. 667-9. 

D. Leslie, “On the Remains of the Megaceros hibernicus in Gypsum in 
Ireland,” “ Geologist,’ vii, p. 165. 

J. B. Jukes, “On Some indentations in Bones of a Cervus megaceros found 
in June, 1863, . . . bog near Legan, co. Longford,” ‘ Journ. Geol. Soe. 
Dulblmnyeyoel27e 

B. Gastaldi, “‘ Intorno ad alcuni fossili del Piemonte e d. Toscana,” ‘ Mem. 
R. Acead. Sci. Torino,’ (2) xxiv, p. 8, pl. i, fig. 14. 

R. A. Couch, “ Notice of the Occurrence of the Horns and Bones of Several 
Species of Deer (Cervus) in the Tin Works of Cornwall,” ‘Trans. R. Geol. 
Soc., Cornwall,’ vii, p. 185. 

A Carte, ““On Some Indented Bones of the Cervus megaceros found near 
Lough Gur, County of Limerick,” ‘ Journ. R. Geol. Soc. Ireland,’ i, p. 151. 
N. Moore, “ Megaceros hibernicus in the Cambridgeshire Fens,’ “ Ann. 
Mag. Nat. Hist.’ (3) xx, p. 77. 

G. V. du Noyer, “ On the Discovery of the Head and Antlers with Some 
of the Bones of Megaceros hibernicus near Kilskeen, co. Meath,” ‘ Journ. 
R. Geol. Soc. Ireland,’ i, p. 247. 

W. Boyd Dawkins, “ On the Distribution of British Post-glacial Mammals,” 
‘Quart. Journ. Geol. Soc.,’ xxv, pp. 192—211. 

Anon., “‘ Cervus meyaceros Hart mit einer Abbildung . . . Schiadels 
mit Geweih,” * Ber. Nat. hist. Gesellsch. in Augsburg,’ xx, pp. 161—7. 

W. Boyd Dawkins, “On the Cervidee of the Forest Bed of Norfolk and 
Suffolk,” ‘ Quart. Journ. Geol. Soc.,’ xxvii, p. 405. 


1872—3. E. J. Ottmer, ‘Uber an bei Olsburg unweit Peine gefundenes Geweih 


» lise. 


. 1874. 


. 1875. 


~ L873: 


1875. 


(C. euryceros),” ‘ Sitzb. Ver. f. Naturw. Braunschweig,’ v, p. 12. 

J. A. Smith, “ Note on the Remains of the Irish Elk (Megaceros hibernicus) 
found in Scotland,” * Proc. Soc. Antiquaries, Edinburgh,’ ix, pp. 51—6. 

P. J. van Beneden, “ découverte du Cervus hibernicus 1 Tamise,” 
‘Ann. Soc. géol. Belg.,” 1, p. Ixv. 

G. Busk, ‘“ List of Mammalian Remains collected by the Rev. J. M. Mello 
in the Rock-fissure Caves in Creswell Crags, Derbyshire,” ‘ Quart. Journ. 
Geol. Soc.,’ xxxi, p. 689. 

W. Dames, “ Cervus megaceros von Rixdorf,” ‘ Zeitschr. deutsch. geol. 
Ges.,” 
B. Gastaldi, “ Cenni sulla giacitura del Cervus Euryceros,” ‘ Atti R, Accad, 
Lincei,’ (2) 11, pp. 436-41. 


XXVH, p. 481. 


61. 


62. 


63. 


64. 


65. 


66. 


67. 


68. 


69. 


70. 


val 


72. 


73. 


74. 


75. 


76. 


ihe 


78. 


19: 


1876 


1877. 
1878. 
1878. 
1878. 
1879. 
1879. 
1879. 


1880. 


1880. 
1880. 
1880. 
1880. 


1880. 


1881. 
1881. 


1881. 


1882. 


1883. 


THE GIANT DEER. 55 


. R. J. Moss, “ Report on the Exploration of Ballybetagh Bog,” ‘ Proc. R. 

Irish Acad.,’ (2) u, Sei., pp. 547—52. 

G. Porte, “On the Recent Discovery of the Remains of the Cervus megaceros 

at Ballybetagh,” ‘ Proc. R. Irish Acad.,’ (2) 1, Sci., p. 738. 

A. Leith Adams, “ Report on the History of Irish Fossil Mammals,” ‘ Proc. 

R. Irish Acad.,’ (2) m1, Sci., pp. 91—2. 

A Savin, “ List of Vertebrata from the Black or Freshwater Bed at West 

Runton, and the Forest Bed near Cromer, ‘Proc. Norwich Geol. Soc.,’ 4, p. 27. 

W. Williams, “An Attempt to Elucidate the History of the Cervus megaceros, 

commonly called the Irish Elk,” ‘ Sci. Proc. R. Dublin Soc.,’ N.s., 1, p. 105. 

M. Canavari, “Sul Cervus euryceros (giganteus) trovato nel dintorni di 

Camerino,” “ Atti Soc. tosc. Sci. Nat., Pisa,’ proc. verb., i. 

W. Williams, “ On Cervus megaceros,’ “ Rep. Brit. Assoc.’ (Dublin, 1878), 

p. 537. 

R. J. Ussher, “ Discovery of an Ossiferous Cavern near Cappagh, co. Water- 

ford,” ‘ Zoologist,’ (3) 11, pp. 331—3. 

G. C. Laube, “ Notiz tiber das Vorkommen von Cervus megaceros Hart 
bei Franzensbad in Bohmen,” ‘ Verh. k. k. geol. Reichsanstalt,’ 

p- 113. 

EK. T. Newton, “‘ Notes on the Vertebrata of the Pre-Glacial Forest Bed 

Series of the Hast of England,” ‘ Geol. Mag.,’ (2) vii, p. 451. 

A. v. Pelzeln, “ Mitteilungen titber den Riesenhirsch (Cervus megaceros),” 

“Verh. zool. bot. Ges. Wien,’ xxix, 8.B., pp. 29-30. 

A. Remelé, ““ Ueber Cervus megaceros aus dem Diluvium von Hohen-Saaten,” 

‘ Zeitschr. deutsch. geol. Ges.,’ xxxui, p. 650. 

A. Leith Adams, “On the Recent and Extinct Irish Mammals,” ‘ Sci. 

Proc. Dublin Soc.” 15 p: 75. 

R. J. Ussher and A. Leith Adams, “ Notes on the Discovery in Ireland of 

a Bone-cave containing the Remains of the Irish Elk, apparently co-existent 

with Man,” ‘Sci. Proc. R. Dublin Soc.,’ N.s., i, p. 234. 

T. Rupert Jones, “ Cervus megaceros in Berkshire,” ‘ Geol. Mag.,’ (2) vin, 

pp. 95 and 480. 

A. Leith Adams, “ Ballynamintra, co. Waterford,’ ‘Trans. R. Dublin 

Soc.,’ (2) 1, p. 200. 

W. Williams, “On the Occurrence of Megaceros hibernicus Owen in the 

Ancient Lacustrine Deposits of Ireland; with Remarks on the Probable 

Age of these Beds,” ‘ Geol. Mag.,’ (2) vii, pp. 354—63. 

K. T. Newton, ‘‘ The Vertebrata of the Forest Bed Series of Norfolk and 

Suffolk,” ‘Mem. Geol. Surv.,’ p. 58. 

G. H. Kinahan, “ Notes on the Cervus megaceros (Megaceros hibernicus),” 

‘Trans. Edinb. Geol. Soc.,’ iv, pp. 3483—5. 


56 


80. 


81. 


82. 


83. 


84. 


85. 


86. 


87. 


88. 
89. 


90. 


Sle 


92. 


93. 
94. 


95. 
96. 
97. 


98. 


99. 


PLEISTOCENE MAMMALIA. 


1883. L. Riitimeyer, “ Beitrage zu eimer natiirlichen Geschichte der Hirsche,” 
‘ Abhandl. Schweiz. pal. Gesellsch.,’ x, pp. 108—11. 

1885. V. Ball, “On the Collection of the Fossil Mammalia of Ireland in the Science 
and Art Museum, Dublin,” ‘ Trans. R. Dublin Soe.’ (2), ui, p. 337, pl. x1. 

1885. R. Lydekker, ‘ Catalogue of the Fossil Mammalia in the British Museum,’ 
ll, pp. 82—9. 

1886. M. Probst, “ Der Riesenhirsch von Ellwangen,” ‘ Jahreshefte d. Vereins f. 
vaterland. Naturkunde v. Wirttemberg,’ 42 Jahrg., pp. 52—57. 

1888. W. H. Potter, “On the Dimensions of the Irish Elk (Cervus megaceros),” 
(letter), ‘ Field,’ Ixxi, p. 935. 

1890. A. Smith Woodward and C. Davies Sherborn, ‘A Catalogue of British 
Fossil Vertebrata,’ pp. 332—4. 

1890. H. Pohlig, “ Die grossen Saiugethiere der Diluvialzeit,”’ ‘ Zool. Vortrage ’ 
(Leipzig), v, pp. 31—3. 

1891. A. Nehring, “ Uber eine besondere Riesenhirsch-Rasse aus der Gegend von 
Kottbus . . .” ‘Sitzb. Ges. naturforsch. Freunde, Berlin,’ pp. 151—62. 

1892. Anon., Letter in “ Land and Water ’ of April 16th, p. 416. 

1892. H. Pohlig, “ Die Cerviden d. thitiring. diluvial Travertins,”’ ‘ Palzonto- 
eraphica,’ xxxix, pp. 215—75. 

1892. E. Harlé, “. . . Sur des restes de Megaceros du sudouest de la France,” 
‘ Bull. Soc. Hist. nat. Toulouse,’ 26™° an. 

1892. A. Nehring, ‘“ Neue Notizen tiber Cervus megaceros var. Ruffi,” ‘ Sitzb. 
Ges. naturf. Freunde, Berlin,’ pp. 3-6. 

1892. A. Nehring, ““ Kin merkwiirdiges Riesenhirsch-Geweih von Worms a. Rh.,” 
‘Deutsch. Jager-Zeitung,’ xvui, pp. 573—5. 

1892. H. N. Hutchinson, “ Extinct Monsters,’ pp. 225—31. 

1894. H. Pohlig, “Sur un important exemplaire du Cervus euryceros hiberme 
(Owen),”’ ‘ Bull. Soc. Belge Géologie,’ vii, pp. 40—5. 

1894. H. Pohlig, “ Die ersten Funde monstroéser Riesenhirschgeweihe,’ “ Verh. 
naturhist. Ver. preuss. Rheinlande,’ 51 Jahresgang, pp. 196—207. 

1895. C. Reid, “ The Orig of Megaceros-marl,” * Irish Naturalist,’ pp. 131-2. 

1896. C. Sterne, ““ Die vorweltlichen Riesenhirsch,”’ ‘ Prometheus,’ vii. 

1897-8. F. Kinkelin, “ Kleine Notizen aus der geol.-paliont. Sektion,” ‘ Bericht 
Senckenberg. naturforsch. Ges. Frankfurt-am-Main,” pp. 193—5. 

1897. J. G. Millais, ‘ British Deer and their Horns,’ London, pp. 5-17. See also 
a review, ‘ Geol. Mag.,’ (4) v, 1898, pp. 183—40. 


100. 1898. R. Lydekker, ‘ The Deer of all Lands,’ London, pp. 134—41. 
101. 1898. “ Report of British Association Committee on Irish Elk Remains,” ‘ Rep. 


Brit. Assoc.’ (Bristol), p. 548. 


102. 1898. P. M. C. Kermode, “‘ The Irish Elk, Cervus giganteus, in the Isle of Man,” 


‘Geol. Mag.,’ (4) v, pp. 116—19. 


THE GIANT DEER. 57 


103. 1900. W. Deecke, “ Vorkommen von bearbeiteten Riesenhirschknochen bei 
Endigen in Vorpommen,” ‘ Globus,’ 78. 

104. 1901. A. Andreae, ‘ Thiere der Vorwelt.-Reconstructionen vorweltlicher Thiere 
entworfen von G. Keller in Miinchen,’ vi, pp. 31—4. 

105. 1901. K. Gorjanovic-Kramberger, “ Der palaolitische Mensch. und seine Zeit- 
genossen aus dem Diluvium von Krapina in Kroatien,” ‘ Mitth. Anthrop. 
Ges. Wien,’ xxxi, p. 180. 

106. 1901. S. F. Harmer, “ On a Specimen of Cervus belgrandi Lartet (C. verticornis 
Dawk.) from the Forest Bed of East Anglia,” ‘ Trans. Zool. Soc. London,’ 
Xa, [2]. VIC). 

107. 1903. H. H. Johnston, ‘ British Mammals,’ pp. 314—18. 

108. 1903. G. W. Lamplugh, “ Cervus giganteus in the Isle of Man,” ‘Mem. Geol. 
Surv., Geology of the Isle of Man,’ pp. 377—89. 

109. 1904. P. M. C. Kermode and W. A. Herdman, “Illustrated Notes on Manx 
Antiquities,” ‘ Proc. Liverpool Biol. Soc.,’ xix, pp. 70—1. 

110. 1905. H. H. Arnold Bemrose and E. T. Newton, ‘‘ On an Ossiferous Cavern of 
Pleistocene Age at the Hoe Grange Quarry, Longclifie, near Brassington 
(Derbyshire), “ Quart. Journ. Geol. Soc.,’ lxi, p. 52. 

111. 1905. A. Laville, “ Megaceros hibernicus, Hart, aux environs de Paris, dans les 
dépots infranéolithiques,”’ ‘ Feuille des jeun. Nat.,’ iv, pp. 23—5. 

112. 1906. EK. Lénnberg, “ Which is the Taxonomic Position of the Irish Giant Deer 
and Alhed Races ?” ‘ Arkiv Zool. (Upsala),” i, pp. 1—8. 

113. 1906. J. G. Millais, ‘ Mammals of Great Britain and Iveland,’ in, pp. 154—6. 

114. 1906. M. Pavlow, “ Hist. paléont. d. Ongulés,” “ Mém. Acad. Imp. Sci. Pétersb.,’ 
(8) xx, pp. 20-39. 

115. 1906. R. Lydekker, “On the Affinities of the Imsh Elk,” ‘Field’ (London), 
cvil, p. 540. 

116. 1907. R. F. Scharff, “‘On the Relationship of the Irish Elk,” ‘Irish Naturalist,’ 
Xvi, p. 145. 

117. 1908. A. Martelli, “ Noti geol. e paleontol. sul travertino di Ascoli, Piceno,” 
‘ Riv. ital. paleont., Perugia,’ xiv, pp. 97—102. 

118. 1908. A. Fritze, “‘ Beschreib. eines Schidels vom Riesenhirsch (Megaceros hiber- 
nicus Blumenb.),” ‘ Jahrb. Prov. Mus., Hannover,’ p. 39. 

119. 1908. G. Hagmann, “ Ein Riesenhirsch aus d. Elsass,’ ‘ Mitt. Geol. Landes- 
anst. Elsass-Lothr., Strassburg,’ vi, pp. 395-8, pl. ix. 

120. 1909. K. Hescheler, “Der Riesenhirsch,’’ ‘ Neujahrsblatt d. naturforsch. 
Gesellsch. in Ziirich,’ 1, pp. 1—41. 

121. 1909. W. Dietrich, ‘‘ Neue Riesenhirschreste aus dem schwabischen Diluvium,”’ 
‘Jahresh. Ver. Vaterl. Naturk.,’ xv, pp. 132—61. 

122. 1910. P. Dierich, ‘ Beitrage z. Kenntniss prahist. Hirsche,’ Bern. 

123. 1910. H. F. Osborn, ‘ The Age of Mammals,’ p. 394. 

8 


58 


124. 


125. 


126. 


127. 


128. 


129. 


130. 


131. 
132. 


133. 


134. 


135. 


136. 


137. 
138. 


139. 


140. 


14]. 


PLEISTOCENE MAMMALIA. 


1911. 8. von Szentpétery, “ Schadelbruchstiick eines Cervus ewryceros (Cuv.) von 
Olaszteleh,”’ ‘ Muz. Fuzetek Mitt. Min. Geol. Sammlung des Siebenbiirgischen 
Nationalmus.,’ 1, pp. 87—9. 

1912. H. G. Stehlin, “ Les Sables de Rosieres, pres Saint-Florent (Cher), II, 
Paléontologie,” “ Bull. Soc. Géol. France,’ (4) xu, p. 201. 

1912. L. Joleaud, “ Sur la position systématique de Cervus pachygenys (Pomel) 
du Quaternaire Algérien,” ‘ Bull. Soc. Géol. France,’ (4) xu, p. 468. 

1912. R. I. Pocock, “On Antler-growth im the Cervide, with Special Reference 
to Elaphurus and Odocoileus (Dorcacephalus),” ‘ Proc. Zool. Soc.,’ p. 773. 

1913. G. Schweder, “ Abwurfstange eines sibirischen Riesenhirsches, C. eury- 
ceros,” ‘ Korrespondenzbl. d. Naturforsch. Verein, Riga,’ lvi, pp. 35—41. 

1913. C. Grevé, “ Kine Megaceros Abwurfstange aus Sibirien,” ‘ Neue Baltische 
Waidmannsblatter,’ ix Jahrg., xv, pp. 34244. Riga. 

1914. W. Soergel, “ Die diluvialen Saugetiere Badens, 1 Teil, Alt. u. Mit. 
Diluvium,” * Mitt. grossh. bad. geol. Landesanst.,’ 1x. 

1914. Rowland Ward, “ Records of Big Game,’ 7th ed., pp. 77—8. 

1914. H. Stokes, “On Irish Elk and other Animal Remains Found at Howth 
and Ballybetagh, co. Dublin,” ‘ Irish Naturalist,’ p. 113. 

1915. I. Chomenko, “‘ Geweit. Diluvium von Nord Kaukasien,”’ ‘ Trd. Obse. 
izoc. sev. Karv. Kraja,’ iti, p. 9. 

1916. L. Joleaud, “ Cervus (Megaceroides) algericus, Lydekker,’ “Recueil et 
Notices . . . Soc. Arch. de Constantine,’ xlix, pp. 1-67. 

1918. R. F. Scharff, H. J. Seymour and KE. T. Newton, “ The Exploration of 
Castlepook Cave, co. Cork,” ‘ Proc. R. Irish Acad.,’ xxxiv (B), pp. 33—72. 

1920. T. Kormos, “ Bericht iiber d. Samml. u. Musealarbeiten im Jahr 1916,” 
‘ Jahresbericht k. ungar. geol. Reichsanstalt.’ 

1924. R. F. Scharff, “ The Irish Elk,” ‘ Irish Naturalists’ Journal,’ pp. 109—11. 

1924. M. Schlosser, “ Ueber die systematische Stellung jungtertiarer Cerviden.” 
‘Centralbl. f. Min.,’ pp. 630—40. 

1927. W. Soergel, “‘ Cervus megaceros mosbachensis n. sp. und die Stammes- 
geschichte der Riesenhirsche,” ‘Abh. Senckenb. Ges. Franktfurt-a.-M.,’ 
XXXIX, pp. 365-408. 

1928. K. Frentzen and C. Speyer, “ Riesenhirsche aus dem Diluvium des Ober- 
rheingebietes,” ‘ Mitt. bad. geol. Landesanst.,’ x, pp. 177—232. 

1928. J. W. Gregory and Ethel D. Currie, ‘ Vertebrate Fossils from the Glacial 
and Associated Post-glacial Beds of Scotland in the Hunterian Museum, 
University of Glasgow,’ p. 7. 


THE FALLOW DEER. 59 


THE FALLOW DEER. 


Cervus dama. 


It is still not very clear when the Fallow Deer became an inhabitant of Britain 
and what claim it has for inclusion in the British Pleistocene fauna. 

Several authors have cited it as occurring in the Forest Bed, but as Newton! 
remarks, the determination is based on fragments which might equally well belong 
to other species. 

It is not the purpose of this memoir to discuss critically the Forest Bed Cervide, 
which have been described in Sir W. Boyd Dawkins’s memoir.” It is, however, clear 
that dama-like deer, such as Cervus savini, are found in the Forest Bed. Whether or 
not any of these forms are to be regarded as actual varieties of Cervus dama is not, 
perhaps, a matter of much importance. The balance of opinion is probably against 
the inclusion of Cervus dama proper as a member of the Forest Bed fauna. 

The antlers from the Middle Pleistocene of Clacton, Hssex, described by Dawkins 
as Cervus brown were separated by him from C. dama owing to the possession of a 
tine on the anterior face of the antler above the 2nd tine.2 Sir V. Brooke* pointed 
out that this tine is occasionally present in Fallow Deer, and figured examples from 
Greece and Sardinia which show it. He also mentioned a case in which one of his own 
Fallow Deer produced antlers bearing this tine. Hence he argued that C. dama and 
C. brown were identical, and that under the latter designation the early occurrence of 
the Fallow Deer in England has been to some extent obscured. Dawkins at first 
accepted Brooke’s® contention, but afterwards considered® that the occasional presence 
of this additional tine in C. dama had best be regarded as due to reversion, and that 
a separate species or variety intermediate between C. savini and C. dama may be 
recognized in C. brown. Millais’ was also of the opinion that C. brownz is a Fallow 
Deer. 


1 “ Geol. Mag.,’ 1880, p. 452. 

* Pal. Soc., “ The Pleistocene Mammalia,” pt. vi (1887). 

8 Ibid., pl. iv, fig. 3e. 

4“ Nature,’ xi, pp. 210—11. 

5 Ibid., p. 226. 

8 Pal. Soc., “The Pleistocene Mammalia,” pt. vi (1887), p. 17. 
7 “Mammals of Great Britain and Ireland ’ (1906), p. 371. 


60 


PLEISTOCENE MAMMALIA. 


Fic. 31.—A Skull and antlers of Cervus megaceros var. ruffi, Nehring, antero-lateral view, from the’ Rhine near Worms; 


Paulus Museum, Worms. 


After Nehring. B Antler of Fallow Deer, Cervus dama, from the7peat of Clapton, Essex 


(M. 4104, Brit. Mus. Nat. Hist.). 


THE FALLOW DEER. 6] 


The evidence for the recognition of C. dama as a member of the Paleolithic fauna 
is inconclusive, and there has been much difference of opinion on the subject. Owen! 
says that there is no decisive evidence of it as coeval with Megaceros and Red Deer. 
He alludes to Buckland’s record of fragments of palmated antlers from Kirkdale and 
Paviland, and to others from the peat at Newbury and from the clay and lignite at 
Bacton, but remarks that such specimens are far from yielding satisfactory grounds 
for identification. 

Dawkins does not include the Fallow Deer in his list of Post-glacial mammals,? 
and Woodward and Sherborn? say of it, “ not a British fossil.” Osborn* does not 
admit it as a member of the 3rd or Upper Pleistocene fauna, including it only in the 
4th or Post-Pleistocene fauna. On the other hand there is evidence of its presence 
in Paleolithic deposits. W.W.Cordeaux® records it from Newnham, near Cambridge ; 
and the Sedgwick Museum, Cambridge, has a series of bones and fragments of antlers 
from the gravel at Newmarket. 

Perhaps the most important piece of evidence as to the occurrence of the Fallow 
Deer in Britain in Pleistocene times is afforded by the remains from the cave at Hoe 
Grange, Derbyshire, described by Bemrose and Newton.® A great number of bones, 
teeth and fragmentary antlers were found which from their small size the authors 
attributed with some hesitation to the Fallow Deer. In the discussion on the paper 
Dawkins expressed scepticism, pointing out that not much importance could be 
attached to measurements of bones and teeth in such a variable group as the 
Cervide. 

The British Museum has an antler (Text-fig. 31, B) said to be from the Neolithic of 
Clapton, Essex. Dr. Frank Corner has two metapodials from the high terrace gravels 
of Dovercourt Park, and it is difficult to avoid the conclusion that they belong to C. 
dama. The evidence at present available as to its occurrence in Britain seems to be, 
that while (as C. brown) it is a member of the Forest Bed fauna and occurs sparingly 
in the Paleolithic deposits, it did not survive to Neolithic times. It was reintroduced 
into Britain possibly, as Millais suggests, by the Phoenicians, possibly, as in the opinion 
of most of those who have alluded to the subject, by the Romans. 

Jeitteles’ assembled a large number of records of C. dama from strata at different 
horizons in various parts of Kurope, and claimed that they proved the wide distri- 
bution of the animal in prehistoric times. Dawkins* commented on the uncritical 


1“ Brit. Foss. Mamm..,’ p. 483. 
2 «Quart. Journ. Geol. Soc.,’ xxiv (1868), pp. 511—16. 
3‘ British Fossil Vertebrata,’ p. 330. 
4“ Age of Mammals,’ 1910, p. 428. 
5 «Proc. Camb. Phil. Soc.,’ iii (1880), p. 348. 
6 * Quart. Journ. Geol. Soc.,’ 1xi (1903), p. 32. 
7 “ Der zoologische Garten,” August, 1874; see also translation by Sclater, ‘ Nature,’ xi, p. 71. 
8 « Nature,’ xi, pp. 112—14. 
9 


62 PLEISTOCENE MAMMALIA. 


acceptance of many of these records, and concluded that the supposed introduction 
by man of the Fallow Deer into Northern Kurope was not disproved by them. 
There seems no doubt as to its having been an inhabitant of the Iberian 
Peninsula at an early date. Busk and Falconer! record it from the caves of Gibraltar, 
Pacheco” from the Spanish Pleistocene, and the latter shows that the wall-figures 
in the caves of Pefio di Candamo indicate its presence there in Magdalenian times. 


1 “Quart. Journ. Geol. Soc.,’ xxi (1865), p. 365. 
2 “Mem. Com. Invest. Paleont. Prehist.,’ xxiv, p. 177 (Madrid). 


PLATE I. 
TrisH Giant DEER. 
Cervus giganteus. 
Cranium. 


(Two-fifths natural size.) 
Fig. 
1. Dorsal ) view of a female cranium (M. 2326) from “ Ireland,” in the British 
2. Palatal) Museum. 
Occipital condyle. 
Anterior palatine vacuity. 
Fronto-lachrymal vacuity. 
Posterior palatine foramen. 
Supra-orbital foramen. 


oS SS Ss 


PALZONTOGRAPHICAL SOCIETY, 1927. 
Reynolds, Giant Deer. PL.I 


eS 2.x 


J : Gre LL, photo. del. C Whittingham & Griggs L'* Imp 


CERVUS GIGANTEUS. 


_ Cranium. 


PLATE II. 
Ir1sH Giant DEER. 
Cervus giganteus. 
Cranvum and mandible. 


(Fig. 3 two-thirds natural size, the remainder two-fifths.) 


Fie. 

1. Cranium (M. 2326) seen from right side. 

. Outer side of left mandibular ramus (M. 2328). 

3. Inner side of part of right mandibular ramus (M. 26124), with roots of the teeth 

exposed. 
4. Palatal view of left lower premolars and molars of M. 2328. 
All from Ireland and in the British Museum (Natural History). 

Occipital condyle. 
Paroccipital process of exoccipital. 
Fronto-lachrymal vacuity. 
Sagittal crest. 
Postorbital bar. 
Coronoid process. 
Mandibular process. 
Mental foramen. 
Alveolus for canine. 


= 


So S&S SSO Gy oS Si 


‘oT Gipuey| pue wintuet*) 


“SNALINVOID SNA 


duy pT shbuap y weybunnym oO [2p ‘oqoyd ‘Use1g CC 


Il'Id a ‘19aq YURID ‘spjoukeay 


1 
f 
Bt) ie : t 
1 
i |! 
iayl 
| 
fy / 
Hy \ 
| . 
i : 
ive ‘ 
1 . 
i 
a | 
1a 
ey 
T 
. ( 
{ 
qt 
hit 
i 
{ 
‘al 
en ) 
H | 
H \) 
* 


* ¢ 
ee 
‘ » 


a ve 


g 


pila oy 


e 


\ 
f 


fet? oe = aay te 77) NN iy Ta RE . 2 ae 
IVINOSHLINS S3tu¥vWugI? LIBRARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS Sa IyvVusiIT LIBE 
id Zz z = x Zz 
eS w ul < on = SA ul 
~ = : : ze : G NS z 
SNe = <x on < 4 <x = 
NN Ps c we = o = \ o = 
om = [2] = a = oO = 
: wll a pa =z a Zz = Zz ; 
SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS S3t¥vVYsiT LIBRARIES NO! 
r- = ine = - < = z 
== 5 _ _ B = oO 
t, —_ ow — ow _ ; wo — 
OO : : : : 5 MS : 
UT fe 7 a = Eat re a f= 2 YP a = 
> m w m Ww m w . - m wn” 


VVINOSHLINS S3IMYUSIT LIBRARIES SMITHSONIAN INSTITUTION NOILONLILSNI NVINOSHLINS S31uyvugIT_LIBF 


wo ; 78 ” 5 ” =z n” . 
= i = : Es < = : 
= Yr. 5 = N\ = as rs . 
wo "hE DP NX wo ~ WW a kD wo 

oO hy? fi = Ww 8 oO * oO = oO ¥ 
2 e Zz \S Z = z 
>" y > . oe > P 
Fe 79) XS z S = = 


SMITHSONIAN INSTITUTION NOIMLNLILSNI_ NVINOSHLIWS Saiuvyuagi LIBRARIES SMITHSONIAN INSTITUTION NOIL 


LIBRARIES 
NOILNLILSNI 


NOLL LILSNI 
NOILALILSNI 
LIBRARIES 
NOILALILSNI 


WVINOSHLINS S3JIYVYSIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31NVUSIT LIBI 


S3iNVY9!IT LIBRARIES SMITHSONIAN 


NVINOSHLINS S3IYVYSIT LIBRARIES SMITHSONIAN 


= - z z = z 
Ni ro) o ° ° wo Xs ° 
SE = = =: get ee = = Se - 
ER = > - GG = et Sage 
Lay a = by G rg a = 
a m n Vfl o m ,? 
ee 7) z ; z ps te 
SMITHSONIAN INSTITUTION NOILOLILSNI NVINOSHLINS S31¥VYSIT LIBRARIES SMITHSONIAN 
z e on = 622) Fe vee ” = 
= . = S = oe ¢ = \ WN = = 
z St ee War 4 wl fy = NS | = a 
3 2 ify & 5 GY 3X 3 = 
ae 0 GY, i So Uys 2 \. fo) = ; 
ee z Yj, E z Yi) = NN 2 E Y 
S PS bey le = = a = = | 
” : » a a. 
NVINOSHLINS S3IYVYSIT LIBRARI ES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS Sa 1yYVYsIT LIB 
A Le = A = 
NOG wW a iw a a 5 NES Gi Fr 
Ke = 4 =) se . % WE = S 
ae = oe = = = \ = 
SY 2 = oe S : S SY = = 
> 6 = © = 0 a * = 
Seed 75 _ z = Fad Ss eee a 
SMITHSONIAN INSTITUTION NOILNLILSNI S3tyYvVYuYgiIT LIBRARIES SMITHSONIAN INSTITUTION NOIJ 
a z Ee = a 3 iE = 
= a 
a = oa > = o = 1 ies) — 
2 5 2 Qy = = - 4% > = 
i> = > SAN = > = V4 i> m4 
8) = ) > WN = bs) = (Pp C4. 2 = 
. — on SY NS — Pe 4 — 
= w = ~ \ SS <2) _ ” m ” 
77) s wo ie = wo z 7) z 
NVINOSHLINS S32 1yVvyuattT a Li B RARI ES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLIWS | S3 tyvUuEsit Pid 
3 Yyp = a = cae = a z 
z Yi 5S 5 5 NWS x x es 
[e) hy Fi aks oO at mS IW an © | oO = ‘ 
Sy Eee 2Z E Wr" 2. 2, Sy 
2 a 2 = » = 3 a 


SMITHSONIAN INSTITUTION NOILALILSNI_ NVINOSHLIWS S3INVYGII- LIBRARIES SMITHSONIAN INSTITUTION NOId 


LIB RARIES SMITHSONIAN 


z a Z a z = ” 

Yjn,® = z : : 3 : 

a fa < a < = ia < 

7% 3 Siz = foal 3 faa} = = fee] 

oO ~- Oo S —_ ie) oO - 

z | Zz wl =z -— al 
NVINOSHLINS S3ZIYVYYUSIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3INVYUSIT LIB 

N oO a so) Oo Oo oe \ Oo o 

\ ° 5 gh ty ® r TAN EF : 

WES 2 > = re. > = > - woe > 

AS = 2 = % a = a = saad 

r ~ + GG : - t a 

2 s z a z ye ae o 


SMITHSONIAN INSTITUTION NOILONLILSNI NVINOSHLINS S3IHYVNSIT LIBRARIES SMITHSONIAN INSTITUTION. NOI. 


Brateaertn oe, 77) A hte Pete le tery ik. Sree ee 7) =F a et wo 
MITHSONIAN INSTITUTION NOILMLILSNI_NVINOSHUWS S3IYVYSIT_LIBRARIES SMITHSONIAN _ INSTITUTION  NOILN4I 
= us z Nae a re 2 uJ 
i E GN = z 3 = 
=i < a Ws = < a < 
NS < 
c ex c NN oc S oc = ox 
= rae] = < ao aa faa) a aa) 
ae i Sa 2 ah itera 5 
4 | 
VINOSHLINS S3!YVYaIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS_SAluYVvVudi7_ LIBRA! 
. re oi = i Es = weuy S 
. = o = lp OD == o = w 
: : 5 $42 : : : : 
2 & 2 GEE? = > = > 
AY 5 2 = Ake 2 Z 2 : 2 
oy 2 a z ui Z o z o 
MITHSONIAN INSTITUTION NOILOLILSNI_NVINOSHLINS SAluVUaIT LIBRARIES, SMITHSONIAN INSTITUTION NOILNL 
= < = ete oe = < A ey 
= z =| YY = = aN = Vy 
zi A 3 a fp pi ® zo s \wi y/ 
S + ro! I}: x . . =) Se x WY ro) YG 
=p = =. = 8 = FS WNW z: 
2 7) = Yn oS z ” a, Zz 


LIBRARIES SMITHSONIAN NOILMLILSNI NVINOSHLINS S3luvudil LIBRAI 


Z 
22) 
‘ex 
=f 
c 
= 
2) 
= 


= 
= 
= 
oO 
w 
aS 
E 
= 
” 
” S wn = ” Ss ” = 
= 4 a Vin 4 ze = oe = 
OF = < fy Zaz F : AS 2 5 
AS s a a pares a a ENS a = 
- _ o es ro) _ 1S) “SS = fe) 
4 2 _! Faz 2 = Zz 
MITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3I1YVYGIT LIBRARIES SMITHSONIAN INSTITUTION NOILNL 
e = rc «: = Es z iS z 
. tale. = e 3 = z 
“S SO a = 5 = 
> — F . \. = = ==) 
- > rs > ke 5 > ke 
2 = 2 - - e. = F 
5 Z 5 Pie D zZ D 2 
WINOSHLIWS SAINVNEIT_LIBRARIES SMITHSONIAN INSTITUTION | NOILNLILSNI poaleWealT LIBRA 
= ier = «&< = 2 2 Be a 
= z Ww et at = = a SY = = if fie KX 
=. o AY. ew ro) 1) ee ve) LU “YY SD WN 
& 2M 3 2 i AM % bY 2% 
2 Ee » 2 i 2 Gy = S Gg = “XN 
hh ee eee |e : 2 
” y ; w” 
MITHSONIAN INSTITUTION NOILMLILSNI_NVINOSHLINS Sa1UVYSIT_LIBRARIES SMITHSONIAN _ INSTITUTION  NOILNL 
= ME Zz coe = Zz 
G = o ASS w 0 = <o = 
= & pas Rs Nw ~ _ [owe _ a 
Pa < = Was < = oan A = 
5 oe = \\, fc & oc 5 a 
= oO = ‘ re) = fan) = oO 
2 3 g cs $ APOE Washes oi ee = 
IVINOSHLINS_ S31YVUGIT_ LIBRARIES. SMITHSONIAN_ INSTITUTION NOILMLILSNI_NVINOSHLINS_S3IYVYUGIT_ LIBRA 
ae : : = : Sa: : 
— wo _ Yy o — > ea @ 
= = GLi, = \, = 
Fe) Lily, a \ 2 
= : 2 He : SW 2 : 
ASS = z — Ski? : NEE x 
BS = = ” m os S\ 2 my 
= 7) < 7) = = a) 


MITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS S31uvuUgIT_LIBRARIES SMITHSONIAN INSTITUTION NOILNL 


NVINOSHLINS S31uvudl 


wae ” ” = = nw 
< = = ; <x ae = 
= or 4 Y re Pra x 4 "4 
ie x = YG.5 WS S Wo: F 
ao an a i ‘ ae \ aio BSS v Vi 
E NO" z. ha aN \ ze AW. 
= SY >" > aa = \ > 
“a Fn Z “a Es a vs = 


WWINOSHLINS S3IYVYEIT LIBRARIES INSTITUTION NOILNLILSNI_ NVINOSHLINS S3lYvVudiT LIBRA 


LIBRARIES SMITHSONIAN 


wl a 5 w a Y w = 

a _ wean a ae ~ ea 

>: , “ a xd : eC % SS wc = 

. =] a 4 SYQSO& 4 

alle 2 e - pe 2 5 

~ = Zz =] 4 af z 
SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S31YVYYSIT LIBRARIES SMITHSONIAN INSTITUTION NOILNJ 

a = FE z e z r z 

o = o e © = .D 2 

wD be a] 27 ke 

> 2 >! " > e > = 

a = 5) _ ee) — 7 2 jis 

- yess = cea - rel e 

2 m ” m ea m n 

wn” = “ . £ ” = ” = 
NVINOSHLINS SAIYVYAIT_LIBRARIES SMITHSONIAN INSTITUTION | NOILALILSNI NVINOSHLINS SaluyWvuall_LIBRA 
a “wy. = we 


N 


| 
ll 


\ 


| 


00611 69 


TION LIBRARIES 


il 


ll 


| 


| 


3 wu 


ih 


SMITHSONIAN INST! 


| 


a teed 
ee eens oem ge 
oe oan nen penn =e ew ant 
pain ae we ene 

Seige name 08 ede 


