Quarterly Journal of the Geological Society of London/Volume 34/Contributions to the History of the Deer of the European Miocene and Pliocene Strata

by William Boyd Dawkins

24. Contributions to the History of the Deer of the European Miocene and Pliocene Strata. By W. Boyd Dawkins, Esq., M.A., F.R.S., F.G.S., Professor of Geology and Palæontology in the Owens College. (Read December 19, 1877.)

Contents.

I.	Introduction.
II.	Classification.
III.	The Capreoli. A. Dicroceros elegans, Cervus dicranoceros, C. australis. B. Cervus Matheroni. C. Cervus cusanus.
IV.	The Axeidæ. A. Cervus perrieri. B. Cervus pardinensis. C. Cervus etueriarum. D. Cervus suttonensis. E. Cervus cylindroceros.
V.	Deer incertæ sedis. Cervus tetraceros.
VI.	General Conclusions.

 

I. Introduction.

The Deer of the European Miocene and Pliocene strata have hitherto been a stumbling-block in the path of the palæontologist, from the fragmentary condition in which their antlers are generally preserved, and the difficulty of separating their variations in form, dependent on age, from those which are worthy to rank as of specific value. They are represented, for the most part, by local names without definitions, which in many cases are synonyms so complicated, that very generally I have found it necessary to examine the original specimen before arriving at an opinion as to their value. To add to the confusion, MM. Croizet and Jobert published their work on the Pliocene Cervidæ of Auvergne^[1] without letterpress, and with the names only of the species printed on the outer coloured cover of each part, which, in the natural course of things, has been rejected by the binder. I have only met with them in one out of the many copies which I have seen, in the Jardin des Plantes in Paris. The work of M. Pomel is without plates^[2], and that projected by that author and M. Bravard (which included the Cervidæ) was never published.

In the following essay, based upon materials collected from time to time in France and Italy, I have defined some of those forms of Deer which are most widely spread and most perfectly preserved, confining my attention solely to the antlers. I have merely attempted to treat a portion of the subject, reserving the rest until new evidence may be brought forward about the numerous other species which are only known to me by obscure fragments.

 

II. Classification.

The antlers denned in the following pages may conveniently be grouped together under the head of (1) Capreoli, or Roe-like, (2) Axeidæ or Eastern Deer, of the type of the Axis and Rusa, and (3) Deer incertæ sedis, which I am unable to bring into close relation with any living forms. They are represented by the following species:—

		Name.		Formation.
1. Capreoli.	$\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$	1. Dicroceros elegans,	Lartet.	Middle Miocene.
		⁠=Prox furcatus,	Hensel.	Miocene.
		2. Cervus dicranoceros	Kaup.	Upper Miocene.
		⁠=C. anoceros,	⁠„
		⁠=C. trigonoceros,	⁠„
		3. C. australis,	De Serres	Lower Pliocene.
		4. C. Matheroni,	Gervais.	Upper Pliocene.
		⁠=C. Bravardi,	Bravard, MS.
		5. C. cusanus,	Croizet and Jobert.	Pliocene.
2. Axidæ...	$\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$	6. Cervus perrieri,	Croizet and Jobert.	Upper Pliocene.
		⁠=C. issiodorensis,	⁠„
		⁠=C. pardinensis,	⁠„
		7. O. etueriarum,	⁠„	⁠„
		⁠=C. rusoides,	Pomel.
		⁠=C. perollensis,	Bravard.
		⁠=C. stylodus,	⁠„	⁠„
		8. C. suttonensis,	Dawkins.	Pliocene.
		9. C. cylindroceros,	⁠„	Upper Pliocene.
		⁠=C. gracilis,	⁠„
2. Incertæ sedis	$\scriptstyle {\left.{\begin{matrix}\ \\\ \end{matrix}}\right\}\,}$	10. C. tetraceros,	Dawkins.	Upper Pliocene.

 

III. The Capreoli.

A. Dicroceros elegans, Cervus dicranoceros, C. australis.

The Deer comprised under this head possess antlers similar to those of the living Muntjak (Cervulus) and Roe (Capreolus), which are short, round, and generally perched on a long pedicle. The crown is either simply forked or composed of short confluent tynes.

The first antler-bearing Deer which appears in the geological record is the Dicroceros elegans (Lartet)^[3] of the Middle Miocene of Sansan and Simorre, in which the antler is composed of a simple fork springing close to the burr, and crowning the summit of a long and slender pedicle like that of the Muntjak. Similar antlers have been met with in the Canton of St. Donnat (Brome) and La Grive, St. Albans (lsère), and are preserved in the geological collection in the Palais des Beaux Arts at Lyons, under the care of my friend Dr. Lortet. In Germany the same form may be recognized under the name of Prox furcatus of Hensel, from the Miocene of Steinheim.

The simple bifurcating type of antler is met with also in the Upper Miocenes of Eppelsheim, considered by Prof. Gaudry to be older than those of Mont Léberon and Cucuron, in the Deer named by Dr. Kaup^[4] C. anoceros and C. dicranoceros, which seems to me to be an older variety of the same form. In these the fork of the antler is further removed from the burr than in the Dicroceros elegans, and is so far therefore more differentiated. Closely allied to them is the Cervus australis of De Serres^[5], from the Lower Pliocene strata of Montpellier, which is the last fossil representative of Deer of this peculiar type in the European Tertiaries.

The difference between these antlers is so very slight that I feel inclined to view their possessors as homologous species, using the term so happily employed by Dr. Heer to denote a lineal ancestry. In the Dicroceros of the Middle Miocenes we find the antler at a minimum of development, consisting merely of a forked crown springing directly from the burr, while in the Deer of the Upper Miocene the forked crown is separated from the burr by a short beam; and this form is repeated in the Lower Pliocene C. australis. The Middle Miocene type is preserved among the existing Deer by the Muntjak, or Cervulus, of the oriental region of Asia.

 

B. Cervus Matheroni (Gervais). (Fig. 1.)

C. Matheroni, Gervais, Paléont. 1859, p. 149; Gaudry, Animaux fossiles de Mont Léberon, 4to, p. 65, pl. 13.

C. Bravardi, Brit. Mus. Cat. no. 34623.

The Cervus Matheroni of the Upper Miocenes of Cucuron and Mont Léberon is considered by Profs. Gervais and Gaudry to belong to the same division of round-antlered Deer as the Axis. It is, however, certain, from the examination of the nearly perfect antlers (fig. 1) in the British Museum, and their comparison with those figured by Prof. Gaudry, that its affinities are rather with the Capreoli. The specimen termed C. Bravardi in the British Museum is the type selected for publication by MM. Pomel and Bravard in a work which, unfortunately, still remains unpublished.

Definition.—The characters of the two antlers (fig. 1), which belong to the same individual, are as follows:—Pedicle moderate, round; antler erect, deeply grooved, four-pointed at most; burr (A) at right angles to long axis of antler and stout; second tyne, D, given off nearly at right angles to beam, oval, waved, upturned at tip; third tyne, E, upturned, round, and at acute angles to beam; crown, CF, small and two-pointed, palmated.

The only differences to be observed between these antlers and those figured and described by Prof. Gaudry are that in the latter the second tyne rises at a smaller angle to the beam, and the third is larger. These are probably due to varying age and possibly race.

Fig. 1.	Fig. 2.
Fig. 1. Antler of Cervus Matheroni, Gerv., Brit. Mus., one third nat. size. Fig. 2. Antler of Cervus cusanus, Cr. & Job., Brit. Mus., one third nat. size.

The absence of the brow-tyne (see B, figs. 3–5) separates this type of antler from that of the division Axis and Rusa, with which it has been compared.

Size.—The head of the animal was about the size of that of a large Roe-deer, which it probably resembled in outer form.

Measurements (inches).

⁠	Brit. Mus.	Gaudry.
Extreme length from burr to crown ................................................................................................................................................................................................................................................................................................................................................................................................	11⋅4
Length of pedicle ................................................................................................................................................................................................................................................................................................................................................................................................	1⋅5	1⋅2
Basal circumference above burr ................................................................................................................................................................................................................................................................................................................................................................................................	2⋅8
From burr to second tyne ................................................................................................................................................................................................................................................................................................................................................................................................	3⋅0	4⋅75
Length of second tyne ................................................................................................................................................................................................................................................................................................................................................................................................	3⋅0
From second to third tyne ................................................................................................................................................................................................................................................................................................................................................................................................	4⋅2	3⋅9

Formation.—Upper Miocenes of Cucuron and Mont Léberon.

 

C. Cervus cusanus, Croizet and Jobert. (Fig. 2.)

Cervus cusanus, Croizet and Jobert, Les Oss. foss. de Puy-de-Dôme, 4to, pl. viii.; Pomel, Cat. Méthodique, p. 111; Gervais, Paléont. p. 149.

The antlers of this Pliocene species belong to the same round-antlered division as the Roe-deer or Capreoli, and are so closely allied to those of the Cervus Matheroni of the Upper Miocenes that the latter species may have been the ancestor of the former. The antler in the British Museum (No. 34610) from the Pliocene strata of Ardé in Le Puy belongs obviously to the same species as that figured but not described by Croizet and Jobert from Mont Perrier near Issoire.

Definition (fig. 2).—Pedicle long, round; antler rounded below, grooved, erect, three-tyned; burr, A, at right angles to long axis of antler and stout; beam flattened as it approaches second tyne, D; second tyne at acute angles to beam, oval, flattened, pointed; crown composed of two flattened tynes, C, E; no brow-tyne.

These characters, in the specimen in the British Museum, are repeated with but little modification in that figured by MM. Croizet and Jobert.

Measurements (inches).

⁠
Extreme length of specimen ................................................................................................................................................................................................................................................................................................................................................................................................	12⋅0
Length of pedicle ................................................................................................................................................................................................................................................................................................................................................................................................	1⋅5
Basal circumference above burr ................................................................................................................................................................................................................................................................................................................................................................................................	2⋅6
From burr to second tyne ................................................................................................................................................................................................................................................................................................................................................................................................	4⋅4
Length of second tyne ................................................................................................................................................................................................................................................................................................................................................................................................	1⋅7

Relation to Roe-deer.—The animal was probably about the size of a Roe-deer, from which it differed in the antlers being longer and more slender, and having the channelled beam free from knobs. In general form the antlers resemble the third antlers of the Roe^[6], and bear to them the same relation as those of Dicroceros to those of the Muntjak. It seems therefore to me almost certain that the Cervus cusanus was the lineal ancestor of the Roe, which makes its first appearance in the forest-bed of Norfolk, and that through it the Capreoline type may be traced back to the Cervus Matheroni of the Upper Miocenes.

From the description of Cervus Cauvieri given by Prof. Gervais, I should infer that it is closely allied to, if not identical with, C. cusanus.

Formation.—Pliocenes of Cuyssac, near Le Puy (Haute Loire), Ardé, and Etuaire, near Issoire.

 

IV. The Axeidæ.

The fossil species grouped together under this head consist of forms closely allied to the round-antlered Deer of the Oriental Region of Mr. Wallace, which possess one brow-tyne and two or three other tynes, such as the Axis, Rusa, Cervus taëvanus, and C. mantchuricus.

 

A. Cervus perrieri. (Figs. 3 & 4.)

Cervus perrieri, Croizet and Jobert, op. cit. pls. iv., v., vi., viii. figs. 9, 10; Pomel, Cat. Méthod, p. 104.

C. issiodorensis, Pomel, Cat. Méthod, p. 105; Gervais, Paléont. p. 147.

The type specimens of the two forms of Deer from Mont Perrier, described under the name of C. perrieri, are preserved in the Jardin des Plantes at Paris, and are sufficiently perfect to offer a basis for defining one species at least of the Pliocene Cervidæ, which hitherto, owing to the unfortunate accident, before alluded to, of Messrs. Croizet and Jobert's figures of the Cervidæ being without descriptions, have been very imperfectly known.

Definition.—A splendid frontlet bearing two antlers nearly perfect offers the following characters:—Antler (fig. 3) round, grooved, and possessed of four tynes—a brow, B, and a second, D, and two terminal, C, E, which form a fork; pedicle short; burr, A, stout and nearly at right angles to beam; brow-tyne round, given off close to burr, nearly at right angles; beam nearly straight between brow-tyne and second tyne, flattened at basement of latter, thence it sweeps backwards to basement of tyne E, which, with tyne C, constitutes the crown; tynes D, C, and E are round, and form acute angles with the beam, the angle being more open in the case of D than of C and E.

On comparing this antler with that of Cervus issiodorensis (fig. 4) in the same Museum the above definition applies with but slight modifications, which are usually met with in antlers belonging to the same species. In the latter the grooves are not so deep; there is a web, or process of antler, at the interspace between the brow-tyne, B, and the beam; the second tyne, D, is set on at a slightly sharper angle, and the beam forms two gentle curvatures, which are not so strong as in the antler of C. perrieri.

Measurements (inches).

	C. perrieri.	⁠C. issiodorensis.
Total length from burr ................................................................................................................................................................................................................................................................................................................................................................................................	31⋅0⁠	32⋅0⁠
Length of pedicle ................................................................................................................................................................................................................................................................................................................................................................................................	1⋅8
Circumference of pedicle ................................................................................................................................................................................................................................................................................................................................................................................................	5⋅8
Basal circumference of antler ................................................................................................................................................................................................................................................................................................................................................................................................	5⋅8	6⋅3
Distance of brow-tyne from burr ................................................................................................................................................................................................................................................................................................................................................................................................	0⋅8
Circumference of brow-tyne ................................................................................................................................................................................................................................................................................................................................................................................................	3⋅9
Brow-tyne to second tyne ................................................................................................................................................................................................................................................................................................................................................................................................	13⋅0⁠	12⋅0⁠
Second tyne to third ................................................................................................................................................................................................................................................................................................................................................................................................	7⋅5	9⋅0
Basal measurement of second tyne ................................................................................................................................................................................................................................................................................................................................................................................................	..	4⋅25⁠

Fig. 6.	Fig. 5.	Fig. 3.	Fig. 4.
Fig. 3. Antler of Cervus perrieri, Cr. & Job. (Jardin des Plantes, Paris), one ninth nat. size.
Fig. 4. Antler of Cervus issiodorensis, Pomel (Jardin des Plantes, Paris), one eighth nat. size.
Fig. 5. Antler of Cervus pardinensis, Cr. & Job. (Jardin des Plantes, Paris), one ninth nat. size.
Fig. 6. Antler of Cervus etueriarum, Cr. & Job. (Jardin des Plantes, Paris), one ninth nat. size.

It will be seen from this description and the measurements that these two so-called species are really merely individual variations of one species, for which I would retain the name of C. perrieri.

I detected antlers belonging to this species preserved in the collection (examined in 1866) of Mr. Dowson at Beccles, Suffolk, comprising basal and coronal parts, one of the former being identical with the variety C. issiodorensis. The locality, however, of these fragments is uncertain, and it is just as likely that they may have been derived from the Pliocenes of France as from the same horizon in the Crags of Norfolk and Suffolk.

A perfect antler with four tynes, in the Museum at Florence, obtained from the Val d'Arno, which I examined in 1877, also belongs to this form, which therefore is common to the Pliocenes of France and Italy.

Living Representative.—This peculiar type of antler, with four tynes, is identical in form with that of the Cervus taëvanus of the island of Formosa, which Dr. Sclater has figured and described from the animals living in the Gardens of the Zoological Society in Regent's Park (Zool. Trans. 1870, p. 345, pls. xxxiii., xxxiv.). It is also identical with that of Cervus mantchuricus figured and described by Dr. Sclater in the same essay ( = Pseudaxis mantchuricus of British Museum Catalogue, which relates to a young animal with a three-tyned antler).

 

B. Cervus pardinensis.

Cervus pardinensis, Croizet and Jobert, op. cit. pl. xi; Pomel, op. cit. p. 106; Gervais, op. cit. p. 140.

The type specimen bearing this name is in the Jardin des Plantes in Paris, and consists of a shed antler, perfect with the exception of the tips of the three tynes, from Pardines, Mont Perrier.

Definition.—It possesses the following characters:—Antler (fig. 5) grooved, round, but slightly curved, and possessed of three tynes; burr (A) stout and oblique to long axis of beam; brow-tyne (B) basal, short, round, and springing at an acute angle; brow-tyne angle webbed; second tyne (D) forming a forked crown, smaller than third, which it joins at an acute angle; coronal angle webbed.

Measurements (inches).

C. pardinensis (Jardin
des Plantes, 1874).

1. Total length from burr ................................................................................................................................................................................................................................................................................................................................................................................................

   22⋅0

1. Circumference of base of antler ................................................................................................................................................................................................................................................................................................................................................................................................

   4⋅7

1. Burr to fork ................................................................................................................................................................................................................................................................................................................................................................................................

   2⋅8

1. Brow-tyne to coronal fork ................................................................................................................................................................................................................................................................................................................................................................................................

   11⋅0

This form of antler seems to me to be closely related to that of Cervus perrieri, and it is very possible that it is a younger antler of that species, related to it as the young Cervus (Pseudaxis of Gray) mantchuricus in the British Museum with three tynes is related to the older animal with four tynes figured by Dr. Sclater, and living in the Zoological Gardens, London, in 1870. It is evident, from the history of antler-development in the round-antlered Deer, that Cervus perrieri must have had an antler with two and three tynes before it arrived at the number of four, or just such an antler as this in question. For these reasons, and although the brow-tyne is set on at a smaller angle than in the type specimen of Cervus perrieri, I feel inclined to view C. pardinensis as a variety and not a distinct species. All these three antlers (figs. 3–5) are found in the same Pliocene strata at Mont Perrier. Nevertheless it must not be forgotten that Deer of the Axis and Rusa type possessing this form of three-tyned antler live in the Oriental Region along with those possessing four-tyned antlers, C. taëvanus and C. mantchuricus, and that therefore it is possible that C. pardinensis may be a distinct species from C. perrieri. For this reason the name is retained in this contribution to the history of the Cervidæ.

 

C. Cervus etueriarum, Croizet and Jobert. (Fig. 6.)

Cervus etueriarum, Croizet and Jobert, op. cit. pl. vi. figs. 1 & 2, and pls. vii. & viii.; Gervais, op. cit. p. 148.

C. rusoides, Pomel, op. cit. p. 106.

C. stylodus, Bravard, MSS. No. 182.

C. peyrollensis, Bravard, MSS.

The antlers (including one of the typical specimens of Croizet and Jobert preserved in the Museum of the Jardin des Plantes at Paris, and referable to the above species or form) are all small, and are as closely allied to Cervus pardinensis as the Axis is to the Rusa. It is, however, safer, in the present imperfect state of our knowledge, to keep them separate.

Definition.—Antlers (fig. 6) possessed of a graceful double curvature, and with three tynes; grooved basally; burr at right angles to long axis of pedicle; pedicle short; brow-tyne (B) set on at an acute angle, which approaches in some specimens a right angle, round; second tyne (D) round; fork of crown webbed, acute-angled in some, right-angled in others.

The antlers which possess these characters I have met with in the Jardin des Plantes from Mont Perrier, and from the Pliocenes of the Val d'Arno, from which place those in the British Museum (Nos. 28833 & 28834) were obtained by Mr. Pentland. I have also observed the same form in the Museum at Lyons in 1873 from Chagny (Saône et Loire), as well as in the Museum at Florence.

The series of antlers in the British Museum obtained from Peyrolles by M. Bravard, and named Cervus peyrollensis and C. stylodus, are undistinguishable from those of C. etueriarum (Nos. 34516 and 34521, 3, 6, 7, 8 of the Museum Catalogue).

Living Representative.—The Cervus etueriarum is closely allied to the Axis, Chetul, or Spotted Deer of India, some varieties of which possess antlers (Brit. Mus.) of the same slender form and double curvature.

Measurements (inches).

		Jardin des Plantes, Mont Perrier.	Palais des Beaux Arts, Lyons, Chagny (Saône et Loire).	Jardin des Plantes, Paris; Val d'Arno.	Ditto.	Ditto.⁠Brit. Mus. Coll., Pentland, 28833.	C. peyrollensis, Brit. Mus., No. 34516.	Ditto, No. 34526.	Ditto, No. 34527.
Total length ................................................................................................................................................................................................................................................................................................................................................................................................		21+	9+	...	...	...	...	11+
Length of pedicle ................................................................................................................................................................................................................................................................................................................................................................................................		1⋅5
Circumference of pedicle ................................................................................................................................................................................................................................................................................................................................................................................................		3⋅8
Basal circumference of antler ................................................................................................................................................................................................................................................................................................................................................................................................		4⋅5	3⋅9	3⋅9	4⋅0	3⋅5	4⋅5	3⋅4	2⋅8
Burr to fork of brow-tyne ................................................................................................................................................................................................................................................................................................................................................................................................		2⋅0	1⋅5	1⋅15	1⋅8	1⋅4	2⋅4	2⋅5	1⋅5
Fork of brow-tyne to fork of second tyne ................................................................................................................................................................................................................................................................................................................................................................................................	$\scriptstyle {\left.{\begin{matrix}\ \\\ \end{matrix}}\right\}\,}$	9⋅5	6⋅0	...	...	8⋅0
Length of third tyne ................................................................................................................................................................................................................................................................................................................................................................................................		10⋅0

Formation.—Upper Pliocenes of Auvergne, Chagny, and the Yal d'Arno.

D. Cervus suttonensis, Dawkins. (Figs. 7, 8, 9, 10.)

A series of antlers in the British Museum from the Crag presents characters which I am unable to identify with those of any species on record, and which I have met with in nearly every collection of Mammalia from the Crag in Norfolk and Suffolk which I have examined. In spite, therefore, of their fragmentary condition, I have ventured to figure and describe them under the name of Cervus suttonensis, because the two most perfect examples were obtained from the Crag of Sutton. Three out of the four antlers chosen as types are in the British Museum, and are more or less waterworn and stained with peroxide of iron, like most of the remains of the Mammalia with which they were associated. All those which I have seen have been shed, and not torn forcibly away from the head; and all have lost the crown and the distal portion of the beam. The specimens in the British Museum have been obtained at Sutton, Felixstowe, and Woodbridge. Those communicated by Mr. Ransome to Professor Owen, and assigned by him to the Miocene Cervus dicranoceros of Kaup, were derived from the Bed Crag of Sutton and Ipswich; those in Mr. Whincopp's collection from that of Woodbridge; those in the Rev. J. Gunn's from the Norwich Crag of Horstead; and those in Mr. Prestwich's from Sutton.

Definition (figs. 7–10).—The base of the antler is cylindrical, and the burr is very strongly marked and circumscribes the base in a plane

Fig. 7.	Fig. 9.
Fig. 8.
	Fig. 10.
Figs. 7–9. Antler of Cervus suttonensis, Dawkins, Sutton (Brit. Mus.), one half nat. size.
Fig. 10. Simple antler of Cervus suttonensis, Dawkins, Woodbridge (Coll. Whincopp), one half nat. size.

oblique to the axis of the beam, which is traversed by clearly denned deep grooves. These, however, are very generally worn away by the action of water. The beam runs straight away from the burr and is cylindrical, except at the point where the brow-tyne springs immediately above the base. At that point there is a smooth triangular area, slightly convex or flat on the superior surface and slightly concave on the inferior, and which is free from the grooves which occur on the rest of the antler. The brow-tyne (B) is slightly oval in section, and gradually tapers to a rounded point, which is broken away in all the specimens which have passed through my hands; it forms an acute angle with the beam, as in Axis and Rusa, and is very much smaller in every dimension. The beam in figs. 8 & 9 is flattened at the point H on its superior surface, which is an indication that a tyne was about to take its rise. The flattening cannot be a mere accidental variation, because it is found in all the antlers which present 4 or 5 inches of beam. Direct evidence as to the crown is wanting; but the fact that all the antlers (some twenty-six or thirty) are broken in some part of the beam, implies that they possessed a crown which was not simple^[7], and the median flattening renders it very probable that it was forked, as in Axis and Rusa. A fragment of a crown of two points, from the Crag of Sutton, in the possession of Mr. Prestwich, may perhaps be assigned to this species. On the whole, the scanty evidence points in the direction of the Axis and Rusa rather than in any other.

In Mr. Whincopp's collection, which I examined at Woodbridge in 1866, is a nearly perfect specimen of a simple styliform antler, about 3 inches long, shed and deeply grooved (fig. 10). It is probably the first young antler of this species. It was accompanied by two fragments of similar form.

Two small waterworn fragments of the base of the antler of Cervus suttonensis have been referred by Prof. Owen to the Cervus dicranoceros of Kaup from the Miocene of Darmstadt. If, however, Prof. Owen's figures in the 'Quarterly Journal of the Geological Society' (vol. vii. p. 234, figs. 14a, 16) be compared with those of Dr. Kaup (Oss. foss. de Darmstadt, tab. 24. figs. 3, 3e), it will be seen that the former antler, which is very much waterworn, possesses a beam (op. cit. fig. 14a) which is much larger than the simple bifurcated antler of C. dicranoceros described by Dr. Kaup^[8].

In C. dicranoceros the beam and the brow-tyne were equal in length or nearly so, while in the series of antlers of C. suttonensis the beam was at least as well developed as in Axis and Rusa, and bore a crown. For the same reason also the series of antlers cannot be referred to the C. australis of Marcel de Serres, from the marine sands of Montpellier.

Measurements (inches).

⁠	British Museum.				Whincopp Coll.			Montmerle.
	No. 27516.	No. 28982.	No. 29505.	No. 27858.
Burr to fork of brow-tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	1⋅12	1⋅6	1⋅95	1⋅98	2⋅4	2⋅2	1⋅25	1⋅0
Circumference above brow-tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	2⋅22	3⋅0	3⋅6	3⋅5
Length of beam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	5⋅5	5⋅0	4⋅0	...	8⋅0
Circumference of base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	3⋅0	...	4⋅4	4⋅4	4⋅2	3⋅5	2⋅6	4⋅0

This type of antler is met with also in France. A specimen which I identified in the Museum at the Palais des Beaux Arts at Lyons, in 1873, was derived from the Pliocene strata of Montmerle (Aire), which, according to Dr. Lortet, are of the same age as those of Chagny. It is almost identical with fig. 8. This form of Deer, therefore, is common to the Pliocenes of Norfolk, Suffolk, and of Central France.

Affinities.—The Cervus suttonensis is, in its general form, closely allied to C. pardinensis, of which it may be a small breed or variety; but, considering the fragmentary nature of the specimens referable to it, I think it safe to keep the two series distinct and under different names. It belongs to the section of the Cervidæ now only found in the hot regions of Eastern Asia.

 

E. Cervus cylindroceros, Dawkins. (Figs. 11, 12.)

Cervus cylindroceros, Bravard, MSS.

C. gracilis, Bravard, MSS.

? C. ambiguus, Pomel, op. cit.

The two antlers described under this name were derived from Ardé, Puy de Dome, and are, so far as I know, without any figures or descriptions, the names merely being those attached to the specimens in the British Museum by their discoverer, M. Bravard. The name is selected because the antler to which it belongs is in a better state of preservation than the other, which is crushed and flattened.

Definition.—The antler of Cervus cylindroceros (fig. 11) is, like those of C. etueriarum, possessed of three tynes and a sigmoid curve; it is round and grooved. The pedicle is short ; burr (A) stout, and nearly at right angles to pedicle; brow-tyne (B) round and rising at a distance from the burr; brow-tyne fork nearly at right angles; second tyne (D) and third (C) with rounded tips; fork between them acute-angled and webbed. The third tyne is longer than the second. These characters are repeated in C. gracilis (fig. 12), with the exception that the second antler is very much smaller, and evidently belonged to a younger animal.

These antlers are distinguished from those which have preceded them by the brow-tyne springing at a distance instead of rising directly from the burr.

Fig. 12.

Fig. 11.

Figs. 11, 12. Antlers of Cervus cylindroceros, Dawkins, Auvergne (Brit. Mus.), one eighth nat. size.

The fragmentary remains which form the type specimens of the Cervus ardeus (or ardei) of Messrs. Croizet and Jobert, from Ardé, and are preserved in the Jardin des Plantes in Paris, are also referable in part to this species, together with some other specimens bearing the same name. The fragment, however, with two tynes, assigned by Croizet and Jobert (pl. ii. fig. 3, and pl. iii. fig, 2) to this animal, probably belongs to some other species.

It is very likely that the C. ambiguus of Pomel, from the Pliocenes of Peyrolles, belongs to this species.

Formation.—Upper Pliocenes of Auvergne.

Measurements (inches).

⁠	Cervus cylindroceros, Brit. Mus.	C. gracilis.	C. ardeus, Jardin des Plantes.
Total length from burr . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	25[9]	18⋅8
Length of pedicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	1⋅5	1⋅1	2⋅2
Circumference of pedicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	4⋅4	...	3⋅8
Basal circumference of antler . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	4⋅5	2⋅7	4⋅2	6⋅0	5⋅5	5⋅3
Burr to brow-tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	2⋅0	...	2⋅5	...	3⋅2	3⋅0
Burr to fork . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	4⋅5	3⋅5	4⋅0	5⋅5	5⋅1	5⋅5
Brow-tyne to second fork . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	13⋅5	8⋅9
Length of brow-tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	7⋅0	6⋅0
Length of second tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	7⋅5	2⋅0
Length of third tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	9⋅5	8⋅0

 

V. Deer incertæ sedis.

Cervine antlers have been met with in the Pliocene strata of France and Italy which cannot be brought into close relation with any of those possessed by living Deer: such, for example, as the Cervus ramosus from Auvergne in the Jardin des Plantes in Paris, and the magnificent pair of antlers from the Val d'Arno in the Geological Museum at Florence, termed Cervus dicranios by Nesti, which has not as yet been accurately determined, and many others, among which the folio wing series of antlers deserves a prominent place from their perfection and their number, and the light they throw on the variation in antler-form in proportion to age.

 

Cervus tetraceros, Dawkins. (Figs. 13, 14, 15, 16, 17.)

C. tetracroceros, Bravard, MSS.

The seven shed antlers bearing these names in the British Museum were derived from the Pliocene strata of Peyrolles in the Puy de Dôme, and are remarkable not merely from their forms, but for their fine state of preservation.

They possess respectively two, three, and four tynes, and evidently follow the usual rule of the development of tynes in the Cervidæ, in which the first appears in the second year, and the rest en suite. They belong therefore to animals four, five, and six years old. The two-year old animal probably possessed a simple styliform antler, while at three years a brow-tyne appeared, as is

Fig. 14.

Fig. 13.

⁠

Fig. 15.

Fig. 16.

Fig. 17.

Figs. 13, 14. Cervus tetraceros, Dawkins, first and second antlers restored.

Fig. 15. Fourth antler of Cervus tetraceros, Dawkins, Peyrolles (Brit. Mus.), one eighth nat. size.

Fig. 16. Fifth antler of Cervus tetraceros, Dawkins, Peyrolles (Brit. Mus.), one eighth nat. size.

Fig. 17. Sixth antler of Cervus tetraceros, Dawkins, Peyrolles (Brit. Mus.), one eighth nat. size.

represented in figs. 13 and 14, which are restorations based upon the antlers 15–17.

Definition.—The antler (fig. 15) of the four-year old possesses the following characters:—Beam (A C) rounded, straight, styliform, channelled; burr (A) oblique to long axis of beam; brow-tyne (B) round, and at rather less than right angles to beam; second tyne (D) round, straight, styliform, at right angles to beam, which is flattened at its point of origin; terminal point (C) rounded, styliform.

In fig. 16 of the five-year old these characters are repeated with slight modifications. The brow-tyne (B) is remarkably long and slender, and springs at a right angle. The third tyne (E) like the second, springs at right angles, but is suddenly reflected at a point 5⋅4 inches from the beam to terminate in a round point 7⋅8 inches from the obtuse angle of reflection in the figured specimens. These characters are presented by three antlers. In the antler of six years old (fig. 17) all the tynes spring at right angles to the beam, and the second (D), third (E), and fourth (F) are gently reflected, the curvature being the greatest in the fourth tyne. These characters are presented also by three other antlers.

It will be observed also that there is a progress in size in these three antlers figured, and that the tynes all spring from the upper and outer side of the beam. All also have been more or less crushed and flattened, but evidently belong to the round-antlered section of Deer.

Measurements (inches).

⁠	Four years. Brit. Mus., No. 34405.	Five years. Brit. Mus., No. 34406.	Six years. Brit. Mus., No. 34409.
Total length from burr . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	15⋅5	23⋅0	31⋅0
Basal circumference of antler . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Length of brow-tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	...	13⋅+	16⋅+
Basal diameter of brow-tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	0⋅6	1⋅1	1⋅2
Brow-tyne to second tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	4⋅0	5⋅0	6⋅0
Length of second tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	7⋅5	13⋅0	15⋅+
Basal diameter of second tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	1⋅3	1⋅2	1⋅4
Second tyne to third tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	...	2⋅6	5⋅0
Length of third tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	...	13⋅2	15⋅6
Basal diameter of third tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	...	1⋅3	1⋅4
Third tyne to fourth tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	...	...	3⋅4
Length of fourth tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	...	...	13⋅8
Basal diameter of fourth tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	...	...	1⋅1
Length of terminal tyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .	9⋅3	12⋅4	12⋅5

Affinities.—The only antler capable of being compared with C. tetraceros is that of the Carjacou, or Cariacus virginianus. It is, however, much smaller, and possesses a brow-tyne which rises at a distance from the base and at acute angles to the beam, and is different in other respects. From the comparison I should infer that the antler of Cervus tetraceros was curved forwards, and that the tynes were erect.

 

VI. General Conclusions.

We may gather from the study of the fossil Cervidæ the important fact that in the Middle Miocene age the Cervine antler consisted of a simple forked crown only. In the Upper Miocenes it becomes more complex, but is still small and erect, like that of the Roe. In the Pliocene it becomes larger and longer, and altogether more complex and differentiated, some forms, such as the Cervus dicranios of Nesti, being the most complicated antlers known either in the living or fossil state. These successive changes are analogous to those which are to be observed in the development of the antlers in the living Deer, which begin with a simple point and increase their number of tynes until their limit be reached. It is obvious from the progressive diminution in size and complexity of the antlers in tracing them back from the Pliocenes into the Upper and Middle Miocenes of Europe, that in the latter period we are approaching the zero of antler development. In the Lower Miocenes I have failed to meet with evidence that the Deer possessed any antlers.

It is further evident from the preceding remarks that the Capreoline type of antler is older than any other.

It is also a point of singular interest to observe that the nearest living analogue of the Miocene Deer is the Muntjak, now found only in the Oriental region of Asia along with the tapir. Cervus dicranoceros also coexisted with that animal in the Upper Miocene forests of Germany.

With one exception all the Pliocene Deer which can be brought into relation with living forms are closely allied to the Axes, Rusæ, or others, which also are dwellers in the Oriental Region. They belong to a fauna now met with only in the forests of India, China, Japan, and the Malay archipelago. The exception is the Cervus cusanus, which possessed an antler not very far removed from that of the Roe, an animal now so widely spread over Europe and Northern and Central Asia. I should infer from this that the Oriental Region has offered a secure place of refuge to the Axeidæ, so abundant in the Pliocenes of France and Italy, from those changes in their environment which compelled them to retreat from Europe. The fact of the presence in this quarter of the world of a group of animals now met with only in warm regions, confirms the conclusions as to the warm climate of Pliocene Europe which M. le Vicomte de Saporta has recently arrived at from a study of the vegetation.

 

Discussion.

Prof. Owen said that he thought we ought to be deeply grateful to the author for the interesting, new, and valuable information his paper had imparted to us. He regarded it as the best and most complete analysis that we possessed of the Cervine fossils, showing, as it did, a close and industrious observation of a vast number of specimens. It was especially valuable for the enlarged and beautiful deductions drawn from individual specimens, and from a comparison of extinct with rare living forms.

Prof. A. Leith Adams agreed with the author in comparing the large horns in one of his diagrams to those of Rusa equina. He suggested that the differences in the antlers might be dependent on differences of age, and stated that the antlers described under the name of Cervus Brownii really represented the type of Cervus dama. One of the small horns resembled youthful horns of Cervus elaphus. He had no doubt that the Indian Deer are remnants of Miocene or, it may be, Eocene species.

The President thought the author fully deserved the eulogy passed upon him by Prof. Owen, and remarked that M. Gaudry had just been writing on the same subject. He referred to the appearance of Ruminants with pachydermatoid characters in Miocene times, and inquired whether the antlers of the Miocene Deer were shed or broken off, the horns in Dicranoceros being stated by Prof. Gaudry to undergo separation by fracture. He further referred to researches of his own upon the conditions of blood-supply associated with the growth of the caducous horns of existing Deer, and suggested that perhaps the creeping in of a cold climate might induce a failure of nutrition, and cause originally permanent horns to fall off.

The Author defended his position against the suggestions of Prof. Adams, and remarked that the larger and more highly developed forms did not occur along with the simpler Capreoline types of antlers of the Miocene. There could be no question of mistaking his larger antlers for those of Red Deer; their number and constancy of form rendered this impossible. Cervus Brownii, he admitted, might be a variety of the living Fallow Deer (Cervus dama), but it is certainly not the normal form of the antler of that species. He pointed to a sketch of a specimen from La Grive, which plainly showed that the antlers of Dicranoceros were deciduous, and stated that Cervus Sedgwickii is probably the same as the C. dicranios of Nesti, from the Val d'Arno, in the Florence Museum.

1. Les Oss. foss. du Dép. du Puy-de-Dôme, 4to, 1828.
2. Cat. Méthodique des Vertébrés Foesiles du Bassin Sup. de la Loire (Paris, 1854).
3. Lartet, 'Notice sur la Colline de Sansan,' p. 34; 'Comptes Rendus,' t. iv. p. 88, and t. v. p. 131.
4. Kaup, 'Les Oss. foss. de Darmstadt;' andKarsten, 'Archiv für Mineralogie,' vi. p. 217.
5. Gervais, Paléont. p. 151, pl. vii. fig. 1; Marcel des Serres, 'Cav. de Lunel-Viel,' p. 230.
6. See Blasius, 'Säugethiere Deutschlands,' p. 463.
7. The cause of the fracture is the firm fixture of the crown in the stratum while the rest was exposed to the clash of the waves which washed the antlers out of the fluviatile gravel in which they were imbedded.
8. Prof. Owen's fig. 16 represents an antler viewed on the underside, and in such perspective that the brow-tyne appears to be of nearly equal size with the beam.
9. 27 inches, following curvature.