© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/

Mamm. biol. 66 (2001) 13-21 Mammalian Biology © Urban & Fischer Verlag http://www.urbanfischer.de/journals/mammbiol Zeitschrift für Säugetierkunde

Review

Evidence for separate specific Status of European {Capreolus capreolus) and Siberian (C pygargus) roe deer

By A. J. M. Hewison and A. Danilkin

Institut de Recherche sur les Grands Mammiferes, Institut National de La Recherche Agronomique, Toulouse, France and Institute of EcoLogy and Evolution, Russian Academy of Sciences, Moscow, Russia

Receipt of Ms. 14. 06. 2000 Acceptance of Ms. 13. 09. 2000

Abstract

Two forms of roe deer, the European {Capreolus capreolus) and the Siberian {Capreolus pygargus), are widely recognised. Some authors consider these two forms as separate species, while others cLassify them as mereLy subspecies or races which are closeLy related. In this study, we compare the geographic distribution, morphoLogical characteristics, karyotypes, biochemical variabiLity, and Potential for hybridisation of European and Siberian roe deer, addressing the question of their phylogenetic Status. For most of historicaL times, the ranges of these two forms have been indepen- dent due to physical barriers such as glaciers or flooding. Overlap occurred for a time in the Middle Ages and again more recently, for the last few decades, but even then, the potential hybrid zone was small and hybrids are not thought to have persisted. The Siberian roe deer is substantially Lar- ger than its European counterpart in all body measurements, with only the very smallest Siberian individuals and the very largest European deer of approximately equivalent size. Furthermore, the two forms can be reliably distinguished on the basis of cranial shape, due to differential rates of growth of the skull, illustrating the hiatus in morphology between the two forms. All European roe have a karyotype of 2n = 70, while Siberian roe possess between 1 and 14 additional accessory B-chromosomes, increasing clinally from west to east. Changes in karyotype seem to occur at phys- ical boundaries, suggesting the differences are due to partial or total absence of gene flow. On the basis of poLymorphism of several enzymes as well as blood and muscle proteins, the genetic dis- tance between the two forms is characteristic of fairly reliable species. A series of hybridisation experiments have illustrated that, although successful crosses can be achieved, they more offen result in stillbirths or birth compLications leading to the death of both

mother and kid, and reduced or complete infertility among Fl hybrid bucks. It is likely therefore that hybridisation in the wild would be rare or absent, and that hybrids would not persist in the face of immigration of either pure form. We concLude that by all the criteria of classical systematics, the European and Siberian roe deer are separate, good, species, albeit phylogenetically closeLy re- lated.

Key words: Capreolus capreolus, Capreolus pygargus, species Status, systematics

1616-5047/01/66/01-013 $15.00/0. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/

14 A. J. M. Hewison etat.

Introduction Geographie ränge

Although roe deer (Capreolus sp.) were once Fossil records suggest that both the Euro- classified as belonging to the Cervinae sub- pean and Siberian roe deer forms have ex- family, it now seems clear that they are in fact isted since the Pleistocene period (Danilk- part of the Odocoileinae (Groves and Grubb IN and Hewison 1996). However, it seems 1987; Grubb 1993). However, taxonomic re- that their geographica! ranges remained in- lationships within this group are far less evi- dependent due to the giaciation of the Rus- dent, in particular the Status of the various sian piains and the Caspian Sea floods geographica! forms of the genus Capreolus. which extended far northward a!ong the Roe deer cover an enormous geographica! Vo!ga. Once these barriers receded, the Si- distribution, ranging from Great Britain and berian roe deer moved west, coionising the Spain to the Far Hast and from Kazal<:hstan plains Up to the Dneiper and possibly and centra! Asia to northern Scandinavia reaching the northern Caucasus in the Mid- and Siberia, and a !arge amount of data has dle Ages (Flerov 1952). Thus, more re- now accumuiated which reveals great Varia- cently, prior to the twentieth Century, the tion of form Over this ränge. This has lead ranges of European and Siberian roe deer certain authors to suggest that the genus con- overlapped in a small part of their overall tains more than one species and perhaps sev- distribution, in the northern Caucasus and eral subspecific forms (Corbet 1978; Da- possibly also in the Dnepropetrovsk, Kiro-

NiLKiN 1986 a; Lehmann and Sägesser vograd and Orel regions (Fig. 1). Hybridisa- 1986). Here, we review pubhshed data on tion may well have occurred here, but due geographic distribution, morphometry, and to reproductive barriers (see below) and genetics of Capreolus to conclude whether the predominance of the European form, this genus is monospecific or not. they almost certainly did not persist. In

Fig. 1. The distribution of Capreolus, showing Variation in chromosome number (70-84) across its present geo- graphicaL ränge (black Line), the extent of historical maximaL overLap in ränge between the European and Siberian forms () and of present day overLap (d). The ränge of the European roe is to the Left of this overLap zone and the ränge of the Siberian roe is to the right. Adapted from Danilkin and Hewison (1996). © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/

Specific Status of roe deer 15

modern times, reduction in ränge and num- lap in size may occur between the very bers of the Siberian roe due to excessive largest individuals of the European form hunting and the abundance of predators re- and the very smallest Siberian roe deer, sulted in discontinuities in geographica! but more generally there is discontinuity in ränge and isolation of the European and Si- average size between adjacent populations berian populations (Danilkin and Hewi- at the ränge limits between these two forms. SON 1996). However, numbers started to re- This discontinuity is due to differential rates cover from the 1930s due to moderation of of early growth and development: kids aver- hunting and a warmer cUmate and the Over- aged 4 kg weight gain per month for Euro- all ränge increased once again. As recently pean roe and 6 kg per month for Siberian as the 1960s, the advance westward of Si- roe when the two were kept together under berian roe deer reached the Volga and sub- identical environmental conditions (Da- sequently the Khoper and Don rivers in nilkin and Hewison 1996). The difference the Volgograd region, bringing European persisted through to adulthood, when the and Siberian deer into contact once more Siberian roe weighed about 20% more in during the last couple of decades (Fig. 1). all seasons. Thus, the geographic ranges of the Euro- In addition to simple size Variation, Euro- pean and the Siberian roe deer have been pean and Siberian roe deer can be distin- largely independent for much of history, guished on the basis of cranial shape. Multi- overlapping only in a restricted area during variate analyses of 905 skulls from certain periods. The complete isolation of populations over the entire geographica! the ranges of these two forms has only very ränge have identified two well-differen- recently been bridged again and the poten- tiated morphs, the Siberian and the Euro- tial hybrid zone remains very small with re- pean (SoKOLOv et al. 1985 a). Again, this spect to the total geographic ränge. Further- discontinuity appears early in life due to more, there is Uttle evidence that hybrids differential growth rates of the skull (Soko- have persisted in any area, probably due to Lov et al. 1985 b). There are also some indi- reproductive isolation between the two cations from this type of analysis that forms. Despite the fact that a large number further discrimination within each main of Siberian deer have been used for intro- group may be possible, particularly for the duction programmes within the European Siberian morph (northern Siberia and the roe deer's ränge, only those introductions Far East), perhaps supporting the designa- that took place where the European form tion of two or more subgroups (Markov et was present in very low numbers or entirely al. 1985 a; Sokolov et al. 1986 a; see also absent have proved successful (Danilkin Hewison 1997). An analogous analysis of and Hewison 1996). antler characteristics was unable to distin- guish clearly between European and Sibe- rian forms, presumably because of the pro- Morphology nounced influence of age, condition and environmental factors on these structures

Despite the fact that there is clearly sub- which are regrown annually (Danilkin and stantial environmental influence on overall Hewison 1996). body size and weight of roe deer (e. g. Gail-

LARD et al. 1996; Hewison et al. 1996 a, b), the European form is markedly smaller Genetic and blochemkal vanabillty than the Siberian form in all body dimen- sions (Fig. 2), including size of antlers (Eu- The karyotypes (chromosomal morphol- ropean: length 17-26 cm, span 7-14 cm, Si- ogy) of European and Siberian roe deer dif- berian: length > 27 cm, span 17-20 cm) and fer dramatically. All populations of the Skulls (condylobasal length: European 180- European form are characterised by an 200 mm, Siberian 201-231 mm). Some over- identical karyotype of 2n = 70, while all Si- © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/

16 A. J. M. Hewison et al.

70 160 60 140

50 120 CD o

100 40 3 00

80 30

60 20 40

20 10

6 j ^ ? Body Body Hind foot Body length girth length mass

Character

Fig. 2. Variation in total body Length, body girth, hind foot Length and body mass of European (white bars) and Siberian (bLack bars) maLe and female roe deer (adapted from Danilkin and Hewison, 1996). The central dot de- notes the average vaLue for each group, the bar gives the Limits for popuLation averages and the verticaL Line shows the ränge of extreme vaLues of Single individuals. berian populations have karyotypes which of the Siberian parent, probably due to un- contain 1 to 14 additional accessory B-chro- equal segregation during meiosis. At the mosomes, 2n = 70 + (1-14) (Danilkin 1985; notional boundary between the two forms

SoKOLOv et al. 1986 b). Furthermore, the Si- in the Ukraine and the northern Caucasus, berian roe deer exhibits mosaicism, particu- individuals both with and without B-chro- larly in the Far Hast, where different num- mosomes have been identified (Danilkin bers of B-chromosomes occur within and Hewison 1996). different tissues of the same animal, as well At the biochemical level, electrophoresis of as among individuals of the same popula- certain enzymes has revealed differences in tion. In addition, all 35 pairs of the main protein polymorphism between European set of chromosomes differ in length be- and Siberian roe. Of 14 Systems tested, 3 tween the European and Siberian groups. were Polymorphie in the European sample, The number of B-chromosomes present while only two were Polymorphie in the Si- shows clear clinal Variation, increasing berian sample and frequency differences steadily from west to east (Fig. 1). However, between the two forms were found at one changes in karyotype across the geographic particular enzyme locus (Sokolov et al. distribution of roe deer are abrupt and 1986 c). Isoelectric focusing of blood plasma seem to occur at physical barriers such as proteins has identified differences in the mountain ranges. Hybridisation (see below) pre-albumin zone of the spectrum which leads to inheritance of some B-chromo- are fixed, i.e. all Siberian individuals are somes among offspring, but the number in- different from all European roe. Similarly, herited is usually less than half the number differences are also present in the lEF spec- © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/

Specific Status of roe deer 17

tra of soluble proteins of the muscle tissue. Table 1. Some outcomes of experimental hybridization These different protein fractions probably of Siberian and European roe deer (adapted from Da- nilkin and Hewison 1996) represent the products of alternative alleles for particular loci. Additionally, immuno- Cross^ Result chemical investigations have indicated that the blood serum of European roe deer con- FemaLe x Male n Successful Normal tains certain antigens which are characteris- Mating^ Delivery^ tic of this group only and may also include Sib. X Eur. 19 7 4 two accessory antigens with very different Eur. X Sib. 38 22 8 et al. b). molecular weights (Markov 1985 Fl X Eur. 4 3 2

Fl X Sib. 3 3 3

Eur. X Fl 11 1 1 Hybridisation Sib. X Fl 2 1 1 Fix Fl 10 0 BClSib. X Sib. 2 2 1 A large number of introductions of Siberian BClEur. X Eur. 2 2 deer into areas inhabited by European roe carried in- have been out with the aim of ^Designations for crosses are: Sib. - Siberian roe; Eur. creasing body weights and improving tro- - European roe; Fl - first generation hybrid; BClSib. - phy quality (Danilkin and Hewison 1996). progeny of Fl doe x Siberian buck cross; BClEur. - Indeed, those hybrids that are able to sur- progeny of Fl doe x European buck cross; ^successful vive are heavier and have larger antlers mating indicates embryos were produced; ^normal de- livery indicates unassisted birth of live kids than the pure European form. However, it seems probable that such Operations have proved unsuccessful (see above), with even the introduction of a substantial number that European roe does were unable to bare (several dozen) of Siberian animals result- hybrid offspring, often dying in the process ing in gradual but complete loss of the Si- of giving birth, while all attempts to cross berian form. European bucks with Siberian does were Hybrid populations have not developed in unsuccessful. In yet another set of experi- the wild due to rather high level of repro- ments, Danilkin (1986 b) did succeed in ductive Isolation between the European Crossing European bucks with Siberian and Siberian groups, illustrated by a series does, however, this resulted in a high pro- of experiments on captive deer. In the first portion of Stillbirths. Overall, crosses be- experiments (Stubbe and Bruchholz 1979, tween the two roe deer forms seems to be 1980), two Siberian bucks were mated with possible, but with a much lower level of suc- a group of European does a total of cess than that observed from normal repro- 32 times. Of these matings, 13 did not result duction, with about 20% resulting in the in pregnancy while 19 births were recorded. birth of live offspring without the need of

Caesarean delivery was necessary in 9 cases some form of assistance (Tab. 1). and another 3 required manual assistance due to the large size of the kid. The level of reproductive Isolation between the Siberian Discussion and the European roe deer is clearly de- monstrated by the fact that 10 subsequent Speeles are generally distinguished accord- matings between two Fl hybrid bucks and a ing to the independence of their geographi- group of hybrid does did not produce a Sin- ca! distribution, discontinuity in character gle offspring. Indeed, it seems that many hy- Variation and reproductive Isolation. We brid bucks are sterile, however, back-crosses have highlighted clearly here that the Sibe- between hybrid does and pure bucks of rian and the European roe deer have occu- either form did produce viable offspring. Si- pied geographically independent ranges milarly, Sokolov and Gromov (1985) found during the vast majority of historical times. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/

18 A. J. M. Hewison et aL.

The ranges of these two forms have come to When crosses produce sterile offspring sub- overlap again since the 1970s, but this po- genus Status is generally accorded, while

tential hybrid zone is very small with re- when offspring are fertile but have a re- spect to the overall geographic distribution duced probability of survival and/or repro-

and is unhkely to have had substantial im- duction parental forms are considered good pact due to its recent occurrence. Introduc- Speeles. Hybrids of several other cerivd Spe- tions of the Siberian roe to sites within the eles have been reported (Wishart 1980; ränge of the European form have generally Bartos et al. 1981) and these are often fer- proved unsuccessful and are of local impor- tile, forming hybrid populations in the wild tance only. (Harrington 1985). However, the data

There is discontinuity in a wide variety of summarised above clearly show that Euro- morphological or physiological characters pean and Siberian roe deer crosses are asso- between the European and Siberian forms, ciated with a high proportion of stillbirths, notably in body size, craniometry (Sokolov the frequent death of both mother and

et al. 1985 a), including non-metric charac- young due to the inability of European roe teristics (Zima 1989), and basal metabohc does to give birth to large hybrid kids and rates (Grayevskaya et al. 1980), even be- a high level of sterility among hybrid bucks. tween geographically adjacent populations. Thus, in a potential hybrid zone in the wild,

This discontinuity is also found at the tissue we might expect a low rate of successful level as cytogenetic, immunochemical and mixed-pair reproduction and generally low biochemical differences (Sokolov et al. productivity of the hybrid population.

1986 b, 1986 c) and may include a certain Thus, there is overwhelming evidence for degree of histoincompatibility. Combining all the criteria of classical systematics that the results of studies of biochemical Varia- the European and the Siberian roe deer tion suggests that the genetic distance be- are two distinct Speeles, albeit very closely tween the European and the Siberian related. The ecological similarities between groups is characteristic of fairly reliable the European and Siberian forms in feeding Speeles and indicates a rather high degree (differences in diet composition are essen- of reproductive Isolation (see Hartl et al. tially due to contrasting plant availabihties 1998 for comparison of within Speeles gene in Asia and Europe), behaviour (communi- flow for European roe deer). cation, sexual and maternal behaviour, on- The roe deer phenotype seems to vary ac- togeny), social and spatial Organisation cording to the number of B-chromosomes (group size, family group structure, male present, indicating a pivotal role for these territoriahty) and dynamics underlines their accessory structures in roe deer taxonomy extremely close phylogenetic relationship. and providing a defining character for Spe- Siberian roe deer are more adapted to liv- eles designation. Indeed, patterns of B- ing under extreme climatic conditions, par- chromosome distribution may indicate that ticularly deep snow and prolonged periods roe deer originated in central Asia, perhaps of low temperatures (Danilkin and Hewi- in the Altai mountains, and therefore that son 1996). This may be a result, in part, of

the Siberian form is the more ancient. It physiological differences in energy metabo- seems likely that the modern European kar- lism, including the presence and activity of yotype may have been greatly influenced by regulating hormones such as the catechol- the glaciation of the Russian plains which amines and enzymes involved in metabolic curtailed gene flow, leading to accumula- functions such as glucose-6-phosphatase

tion of genetic differences between the (Grayevskaya et al. 1980). The further divi- European and Siberian forms and even- sion of this taxonomic group into the north-

tually to allopatric speciation and reproduc- ern Siberian form (C. p. pygargus) and the tive Isolation, although the possibihty that Southern Tien Shan form (C. p. tianschani- this simply represents clinal Variation with- cus) representing either separate Speeles or

out speciation should be considered. subspecies is far less researched and can be © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/

Specific Status of roe deer 19

considered rather speculative in view of the the Zone of overlap (e. g. extent and conse- current State of knowledge (Danilkin and quences of hybridisation) in order to ad- Hewison 1996). Further research could use- vance our understanding of their distinc- fully concentrate on the relationships be- tiveness. tween European and Siberian roe deer in

Zusammenfassung

Beweise für den Artstatus von Europäischem {Capreolus capreolus) und Sibirischem

(C. pygargus) Rehwild

Beim Rehwild wird im allgemeinen die Existenz zweier verschiedener Formen, des Europäischen {Ca- preolus capreolus) und des Sibirischen {Capreolus pygargus) Rehs angenommen. Einige Autoren be- trachten diese beiden Formen als eigenständige Arten, andere betrachten sie lediglich als nahe ver- wandte Unterarten oder Rassen. Im Hinblick auf eine Überprüfung des Artstatus werden in der vorliegenden Arbeit Ergebnisse über geographische Verbreitung, morphologische Merkmale, Karyo- typen, biochemisch-generische Variabilität und die Fähigkeit zur Bildung von Hybriden zwischen dem Europäischen und dem Sibirischen Reh zusammenfassend gelistet und verglichen. Über den Großteil ihrer Geschichte hinweg war das jeweilige Verbreitungsgebeit der beiden Formen durch Barrieren wie etwa Gletscher oder überflutete Landstriche separiert. Im Mittelalter und in den letzten Jahrzehnten gab es Überlappungen, aber auch dann war eine potentielle Hybridzone klein und es wird nicht angenommen, daß etwaige Hybriden längeren Bestand gehabt haben. Das Sibi- rische Reh ist in allen Körpermaßen deutlich größer als das Europäische Reh, wobei lediglich die kleinsten Sibirischen Rehe den größten Europäischen Rehen annähernd gleichkommen. Außerdem sind die beiden Formen als Resultat unterschiedlicher Wachstumsraten und aufgrund ihrer Schädel- form verläßlich zu unterscheiden. Alle Europäischen Rehe haben einen Karyotyp von 2n = 70, wäh- rend die Sibirischen Rehe zwischen 1 und 14 akzessorische B-Chromosomen besitzen, in Anzahl klinal von West nach Ost ansteigend. Der Wechsel im Karyotyp scheint an geomorphologischen Bar- rieren aufzutreten, was auf ein partielles oder totales Fehlen von Genfluß zurückzuführen sein dürfte. Auf der Grundlage von Enzympolymorphismen und von genetischer Variation in Blut- und Muskeleiweißen liegt der genetische Abstand in einem Bereich, wie er üblicherweise zwischen vali- den Arten gefunden wird. Eine Serie von Kreuzungsversuchen zeigte, daß trotz des Verkommens er- folgreicher Bastardierung, meist Totgeburten auftreten oder Komplikationen bei der Geburt zum Tod von Mutter und Kind führen. Außerdem gab es eine reduzierte oder vollständige Unfruchtbar- keit bei Fl-Böcken. Das Vorkommen von Hybriden in freier Wildbahn dürfte daher selten oder über- haupt nicht möglich sein, und ein Überdauern von Hybriden wäre angesichts der Überzahl von Indi- viduen der jeweiligen reinen Formen auch nicht wahrscheinlich. Wir schlußfolgern, daß nach allen Kriterien der klassischen Systematik das Europäische und das Sibirische Reh valide, wenngleich stammesgeschichtlich nahe verwandte Arten sind.

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