The rabbit in continental Europe




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The rabbit in continental Europe


P. M. Rogers, C. P. A rthur, and R. C. Soriguer
3.1 Palaeontology and history


Lagomorphs have Iived on the continents of Europe, Asia, and North America since the late Eocene or early Oligocene. Members of both families of lago- morphs have occupied the Iberian Peninsula during that time. The family Ochotonidae, once represented by some 15 species in 5 genera, became extinct in Europe (with the exception of Ochofooa pusilla in eastern Russia) at the end of the Pliocene. The Lepor- idae have been reduced from 7 species in 4 genera to the surviving 3 species in two genera, Lepus and Orycfvlagus.

The late Neogene and Quaternary periods saw

many profound ecological changes in Iberia. Late Miocene beds (5-7 million years ago) at Salobrena in the southern Spanish province of Granada have made an important contribution to our understand- ing of those changes. The deposits reveal a fauna which included many forms of direct African origin-giraffes, antelopes, hyaena, and various murids--lying together with Asiatic forms. The data are too sparse to calculate the rate of faunal change, although it seems certain that the African pre-Pleis- tocene immigrants must have bridged across Gibral- tar or the AIboran sea. One European form, Prolagus, must also have made use of that bridge, but in the opposite direction, to colonize North Africa (Table

3.1).


There was a great increase, in both number and kind, of leporids at the end of the Neogene and

during the Quaternary, which was probably related to the development of extensive grasslands, the cool- ing of the climate, and the arrival of Asiatic immi- grants.



3.1.1 The origin of Oryct olagus

the earliest known specimen of Oryctolagus comes from Salobrena. The remains are sparse (one solitary tooth!) and do not permit identification to species (Lopez Martinez 1977b). But they show that Orycto- lagus originated well before the PIeistocene.

There are three plausible hypotheses to explain the ancestry of Orycfolagus. The first is that it is derived from the Asiatic Trischizolagus, but this is difficult to accept as the two genera co-existed during the Upper Miocene of Salobrena (Granada). The second, which suggests an African origin, and the third, an indigenous origin from Aiilepus, seem more likely; but the available evidence does not really favour any of the three hypotheses.

In western Europe, Lepus co-existed with Orycto- lagus, although not constantly. Oryctolagus can survive in any environment, but only as long as the soil is suitable to dig and hide in. Lepus is more widely adaptable, able to live in forest clearings, alpine meadows, or prairie, whatever the soil. Such wide ecological plasticity makes it difficult to deduce much about the habitats available at the time. In Central Europe, Lepus co-existed with Hypvlagus and with Ochofona.



3.1.2 Species of Oryctolagus in the last 2.5 million years

Lopez Martinez (1977h) in an excellent review of Iberian lagomorphs described a new species of Oryctolagus. She also reported new data for



The rabbit in continental Europe 23


Table 3.1 Occurrence of Prolagus sansaniensis in western Palearctic deposits


Country

Miocene

(lower)


Miocene

( middle)



Miocene

(upper)


Pleistocene

Spain France Germany Morocco Portugal Turkey Hungary Greece


X X X
X


X X X


X

X X X


X X X

X X X X


X




O. lacosti, already known from France (Lopez Mar- tinez et al. 1976) and described the oldest known dis- covery of O. cuniculus on the Iberian peninsula, more than half a million years old.

Oryctolagus lacosti, known from Cataluna (Vil- laniense and Bihariense), had a large body size (between that of Lepus and O. laynensis), similar to O. laynensis in Italy and France. Cranial and dental characteristics mark it as an Oryctolagus, although in body size and in the morphology of the premolars (P3) it had similarities with Lepus.

Oryctolagus laynensis appeared in Andalucia, Castilla, and Cataluna in the late Miocene, and is apparently a direct ancestor of present day rabbits. It has also been found in the Pliocene either as a

`descendent' of Trisclrizolagus, also known from the

upper Miocene in Murcia, or as an African immi- grant. It is the oldest known Oryctolagus, with cranial and dental characteristics typical of that genus. Its skeleton appeared to be modified for jumping, but it also had the robust muscular inser- tions, particularly on the humerus and ulna, com- mon in burrowing animals (Donard 1982).

Oryctolagus cuniculus first appeared in Andalucia, in mid-Pleistocene deposits in Cullar Baza, Granada, co-existing with Lepus. It has also been found in the province of Malaga, and in southern France (de Lumley-Woodyear 1969, 1971; Chaline 1976; Lopez- Martinez 197713; Pages 1980). Dental peculiarities suggest that both O. cuniculus and O. Iacosti are descended from a common ancestor, O. laynensis.

A plot of tooth lengths from Quaternary remains

in the Iberian peninsula shows that the earliest species of Oryctolagus, although smaller than Lepus, were larger than the later O. cuniculus (O. lacosti then O. laynensis; Fig. 3.1a). Over the last 13 000 years the sizes of various osteological variables have also tended to decrease in O. cuniculus (Fig. 3.1b); which was a smaller rabbit than either of the other species, perhaps because it always coexisted with Lepus. The rabbit from Cu]Iar Baza (Granada) was similar to O. cuniculus today.

This is consistent with Donard's (]982) research on

the recent history of the rabbit in France. From bones in numerous caves she was able to show that from

300 000 years (Mindel glaciation) to 15 000 years ago ( Magdalenian epoch) local populations of rabbits showed substantial variation in body size, following changes in climate. Four geological sub-species are thus defined: O. c. lunelleusis, a rabbit of moderate size and temperate climate, dating from 300 to 200 000 years ago; O. c. grenalensis, a large rabbit of very dif- ferent climates, dating from 150 to 70 000 years ago; and O. c. li uxleyi, a small rabbit of hot climates some

40 000 years old, still living on the Canary Islands

( where it was introduced long ago, or was possibly there before people) and Madeira; and O. c. cunculus.

3.1.3 Geographical variation in contemporary

Oryct olagus cuniculus in Europe
Not only has O. cuniculus become smaller in the course of its evolution over the last 1 a millennia; it also exhibits a clear reduction in body size over its

24 P. M. Rogers, C. P. A rthur, and R. C. S origuer


The rabbit in continental Europe 25



present range, from north to south (Table 3.2). The rabbits of North and Central Europe are much larger than their relatives in Mediterranean Europe and North Africa. Even within France or the Iberian peninsula, there is a clear north-south gradient in body size.

Several hypotheses have been suggested to

explain this pattern. One associates selection of body size with various ecological influences (climate, predation etc.). The rabbit apparently conforms to one such hypothesis, Bergmann's law, which pre- dicts larger body size in higher latitudes. Soriguer (1981a, 1983b) has shown that rabbit populations which differ in body size also differ in other respects. The Mediterranean Iberian rabbit grows faster, and as we note later, has smaller litters, breeds younger and dies sooner than its bigger cousins. The final result in terms of the rate of increase 'r' is the same, however, albeit through different strategies. Another explanation suggests the influence of human activities (introductions, 'improvements' etc.) and domestication.

A third possibility is that the northern and south-

ern rabbits represent different evolutionary lines. At the time of the glaciation of Mi.ndel (50 000 BC), U. cuniculus had a fairly restricted range. Studies of the systematics of rabbit parasites, principally fleas, suggest that there were then two relict populations of rabbits: one in Spain, where the rabbit was par- ticularly abundant near Gibraltar, and the second in the south of France (Beaucournu 1980a, 1980b). Indeed while 5pilapsyllus cuniculi, the most common flea on rabbits, is present throughout the rabbits' range, three other fleas, Xenopsylla cunicularis, Caenopsylla lactaevn, and Ddontopsyllus guirosi, are present only in France and Spain (including Morocco for X enopsylla). The morphological differences are clearest between French and Spanish individuals of two of these species of flea, suggesting prolonged separation between the two relict rabbit populations (Beaucournu 1980a).

Studies of mitochondrial DNA and different immunoglobulin alleles support this hypothesis (van der Loo et al. 1991; Biju-Duval et a!. 1991). In general

there is high polymorphism in southern Spain, decreasing towards the north. Phylogenetic relation- ships between the rabbit's mtDNA types suggest that the separation of the two relict populations could have begun even before the Mindel glaciation.

Domestic rabbits appear most similar to northern wild rabbits, and least similar to those of Andalucia (Richardson et al. 1980; Biju-Duval et a!. 1991; Van der Loo et a!. 1991; Van der Loo and Arthur in preparation), This suggests that the foundation stock for domestication came from only one (northern) population. It is interesting to note that the mtDNA of French rabbits is more closely related to that of rabbits in the north of Spain than the south. Almost all the alleles of domestic rabbits are present in the wild genotypes. Rabbit populations could have suf- fered genetic impoverishment the further they have moved from their ancestral home.

Today's wild rabbits still retain traces of their diverse historical origins. Genetic characters distin- guish at least three separate rabbit populations in Europe: one in south, central, and west Spain and Portugal, a central one in northern Spain and south- ern France, and a northern one which includes rab- bits in Belgium, England, Scotland, Wales, and northern France. Rabbits in the last group are genetically similar to those now in Australia and New Zealand (Richardson et al. 1980).


3.1.4 History of Oryct ulagus cuniculus in Europe
Donard's (1982) studies showed no rabbits in all deposits of the Holocene (10 000 years ago) and earlier, north of the Loire. No rabbit bones have been found in all the excavations undertaken in the departements (administrative divisions of France) of Cote d'Or, Eure-et-Loire, the north of Charente, in Seine-et-Marne, Essonne, Yonne, Seine-Maritime and Haute-Savoie, whereas the hare (Lepus) is relat- ively well represented. In contrast, rabbit bones have been found in Charente Maritime, a deposit in the south of Charente, in Gironde, Haute Garonne, Her- ault, Bouches-du-Rhone, Ain, Dordogne, Correze and Ardeche (Fig. 3.2).

Many deposits in Herault and Bouches-du-Rhone

contain rabbit remains, sometimes in abundance and notably of the geological sub-species lunellensis. They confirm that the range of the rabbit has extended progressively northwards from a relict population on the coast of the Mediterranean, and that it was only at the start of the historical epoch (20 000 to 1000 sc) that rabbits invaded other lands to the north of the Loire (Donard 1982).

26 P. M. R ogers, C. P. A rthur, and R. C. 5orrguer

Table 3.2 North-south variation in contemporary rabbit weights (g)


Country dcpartement

(France only)



Sweden

Sex Whole weight ( mean)

Sample size

Maximum weight

Reference




F

1670

521



72000

M

7670

521

72000

F

1500




>1700

M

1530







F

1710

45

72000

M

1725

25

72000

F

1450

47




M

1415

56




1

1450

24

1830

M

1350

16

1760

F

1355

17

72000

M

1370

22

1980

F

1385

29




M

1360

23

-

F

1415

67

1830

M

1355

84

1780

F

1500

100

1930

M

1510

83

1860

F

1360

51

1830

M

1405

63

1760

F

1350

37




M

1360

28




F

1373

28




M

1345

31




F

1360

115

1750

M

1430

90

1650

F

1555

15

72000

M

1355

8

>2000

F

1730

9

>2000

M

1630

9

72000

F

1345

33

1640

M

1310

28

1610

F

1290

12

1370

M

1295

18

1450

F

1245

41

1580

M

1210

28

1510

F

1250

22




M

1235

19




F

1354

101




M

1301

104






Andersson eta!. (1979)


Holland

France' N Nord

(sand dune) W Morbihan h`

(heathland)

Vendee

(bocage`)



Deux-Sevres

(bocage) Centre Loiret"

(bocage) CW Yvelines

(farmland)


Yvellnes

(parkland) Essonne



(woodland)

E Haute-Saones ` (pasture)

CE Saone-et-Loire" (pasture)

SW Landes

(fallow/pasture) Landes

(farmland) Haute-Garonne`` (farmland)

Dordogne

(bocage)


SE Hautes-Alpes` (bocage)

Bouches-du-Rhone

(pasture)

Bouches-du-Rhone

(fallow)

Bouches-du-Rhon' (salt-marsh)

W allage- Drees

(pers. comm.)


Arthur and Guenezan

(unpublished) Arthur (unpublished)


Arthur and Aubineau

(unpublished) Arthur and Aubineau

(unpublished) Arthur and Guenezau

unpublished

Arthur and Guenezau

(unpublished)

Arthur (unpublished) Arthur and AngibauIt

(unpublished)

Arthur (unpublished) Arthur (unpublished)

Arthur and Avignon

(unpublished)

Launay (pers. comm.) Launay (pers. comm.)

Arthur and Garcia

(unpublished) Arthur and Reudet (unpublished) Arthur and Gaudin (unpublished)

Vandewalle {pers. comm.) Rogers (1979)

The rabbit in continental Europe 27

Table 3.2 (cont.)


Country




departement

(France only)


Sex

Whole weight

( mean)

Sample size


Maximum weight


Reference



France SE







Herault'

F

1230

10

1510

Arthur and Tarts







(garrigue/culti v.)

M

1250

10

1430

(unpublished)







Corse

F

1235

12

1430

Arthur and Roux







(maquis)

M

1320

14

1490

(unpublished)







Vaucluse

F

1300

45

1590

Arthur and Gaudin







(fallow)

M

1340

65

1541)

(unpublished)

Portugal







F

1017

18




Lopez Ribeiro (1981)










M

1023

11







Spain NE







1190

4




Soriguer (unpublished)

NE







1158

71




Manosa and Real

(unpublished)






N







1224

97




Ceballos (unpublished)




N







1274

113




CebaIlos (unpublished)

NW







934

6




Soriguer (unpublished)

SE







1043

28




Marquez (unpublished)




S







I011

1044

-

Soriguer (unpublished)




S







910

16

1050

Soriguer (unpublished)

SW







1092

521




Soriguer (1980c, 1981a)

SW







923

78




Soriguer (unpublished)

Morocco










1039







Soriguer (unpublished)


All data except Rogers (1979) are from adult rabbits (>9 months) collected by ferreting in January-February.

Data collected by the Service technique of the Federation Departemettale des Chasseurs.

' Data collected during the hunting season {September/October) from adult rabbits (>9 months). e Includes domestic x wild crosses.

Data collected in all seasons by night shooting; January-February mean weights were 1237 g (QC, n - 22) and 1282 g {dd, n = 28).

` A landscape of small fields (pasture or crop) surrounded by hedgerows on mounds.

By Roman times rabbits had become a problem in Spain. In the Balearic Islands they caused so much damage that the colonists asked the Emperor Augus- tus to send a Roman legion to clear the land of them, or to allocate them land elsewhere (according to Strabo, 58 BC to AD 20). Pliny the Elder, in his Natural History, tells of the ramparts of Tarragone being undermined by rabbit warrens, and recommended even then using ferrets against them. The rabbit was already used as the symbol of Spain by the poet Catullus, and imprinted on the reverse of some coins

by the Emperor Hadrian. The Romans modified the range of the wild rabbit by introducing it to many countries, including to Italy and Corsica from France (Bodson 1978; Zeuner 1963), and to North Africa from Spain (C. Louzis personal communication).

In France, domestication of rabbits began much Iater, by monks in the Middle Ages. Their motivation was that newborn rabbits (the famous laurices) were considered to be aquatic, so were authorized in the Catholic religion for consumption in Lent. From the second to the fifteenth centuries there are numerous

28 P. M. Rogers, C. P. Arthur, and R. C. Sorguer




references to rabbits kept by monks and sometimes by peasants, or by the feudal gentry in enclosures of varying extent and degree of security, sometimes made of stone. Rabbits were often sold in markets, as shown in the book Conejena de Toledo (the Rabbitries of Toledo) of the twelfth century. Trade sometimes spread far afield; there are records of the sale of rab- bits between the abbeys of Corvey (Germany) and Solignac (France) in 1148, and the despatch of 60(X) rabbit pelts from Castile to Devon in 1221 (Delort

1984). However, there is scarcely a mention of rab- bits living free in the wild.

Nevertheless the monarchy in France always attempted to restrict ' warren rights', by forbidding the creation of new warrens or the enlargement of old ones, or the re-establishment of old boundaries (there are examples of such rules in the ordonnances of Jean le Bon in 1356 and of Charles VI in 1413). In

the seventeenth century Colbert (prime minister to King Louis XIV) ordered the destruction of rabbits in all the royal forests, and undertook to compensate for rabbit damage where the royal servants were unsuccessful in their task. The French Revolution abrogated Colbert's ordonnance, and annulled the exclusive right of the gentry to control warrens, but the rabbit in France nevertheless remained enclosed and controlled. Not until the Second Empire, in 1862, did Napoleon II] declare rabbits to be game, and order that they be allowed to range freely in the forest of Compiegne.

At the end of the nineteenth century, numerous bourgeois hunts were established close to the towns, on agricultural areas which were abandoned after the Industrial Revolution and the phylloxera crisis. The rabbit became their principal quarry. In less than

20 years, from 1845 to 1863, the number of rabbits




The rabbit in continental Europe 29

Fig. 3.3 Number of rabbits shot annually per hectare on a private hunt of 720 ha in I. uiret, France, 1677-1953 (after Ciban

1956).




sold in the markets of Paris rose from 177 000 to nearly 2 million (Gayot 1865). From then on, the extension of the natural range of the rabbit con- tinued progressively. In Sologne, there were so few rabbits in 1880 that it was judged a waste of effort to fence nurseries for reforestation programmes: in

7930, hunters took more than 10 rabbits/hectare in a season from the same area.

At the start of the twentieth century the introduc- tion of winter cereals and the rotation of crops, com- bined with the massive destruction of all predators (aerial or terrestrial) by gamekeepers, greatly favoured population increases in rabbits. Some

gamekeepers even provided food for game (and especially for rabbits) in winter-apples, beetroot, and fodder. Rabbits quickly reached the very high densities recorded from 1920 to 1930 (Fig. 3.3). Bags of over 10 rabbits/hectare were common in northern departements of France from then until 1950-2. In Mediterranean France bags were between 1 and 2 rabbits/hectare (Giban I956).

Damage to farm crops due to the excessive abund- ance of rabbits did not generally cause concern in the north of France, but it was nevertheless the reason that, on 14 June 1952, Dr Delille introduced the myxoma virus on Maillebois, his estate in the departement of Eure-et-Loire. B ecause of previous failures to introduce the virus in other countries, and

because he did not know the mechanism of trans- mission, Dr Delille assumed that any disease would be confined to his 300 ha walled enclosure. But the result of his experiment was a little more spectacular than he expected: by summer 7952 the disease had reached 9 departements. By the end of 1953 the whole of France was affected (Fig. 3.4), and isolated

cases of myxomatosis were reported from Spain, Belgium, Holland, Germany, and England (Joubert et al. 7972).

For hunters, the rabbit population crash in France was catastrophic. In 1953-4 bags were 15 per cent of those before 1952; in 1954-5, 2 per cent, and in 1953-



6 about 7 per rent. Some 90 to 98 per cent of the

French rabbit population was killed by myxomatosis between 1953 and 1955. The pressure of public out-

rage was enormous, and legal proceedings were started against Dr Delille. He was convicted of illegally spreading an animal disease in September

1954, but received only a nominal penalty-a one franc fine. On the other hand, in 1956 the Syndicat

National des Forestiers Francais presented him with a medal 'in recognition of services rendered to agri- culture and sylviculture'l In fact the estimated increase in agricultural and sylvicultural production at that time was in the order of 1000 million francs (using 1992 values for the franc; Siriez 1957).

French hunters quickly attempted to limit the




30 P. M. Rogers, C. P. A rthur, and R. C. S origuer





effects of the myxoma virus, trying to establish

'cordons sanitaires ' around affected areas, and even,in 1956, importing Australian rabbits reputed

to be resistant to the virus. All their efforts failed. Nevertheless, 1956 saw the start of a small increase in bag size. The first attenuated strains of the virus were found in 1955 (Jacotot et a1. 1936), and they

spread gradually until in 1968 more than half the strains isolated from wild rabbits were much attenu- ated, grade I]IB-]V (Fenner and Ratcliffe 1%5). That proportion seems to have been more or less main- tained until 1977-8 (Joubert 1979). Over the last four or five years, however, there has been an increase in the more virulent strains.





3.2 Present distribution and regional variations in numbers


The rabbit is now found over most of western Europe, excluding most of Austria, Italy, and Swit- zerland, and around the coasts of the western Medi- terranean Sea (Fig. 2.2). Its eastern range extends as far as the Crimea. It is also found on the Balearic Islands, Corsica, Sardinia, Sicily, and Crete.

Wild rabbits were introduced in southern Ukraine

at the end of the nineteenth century. About 80 years later their range was increased extensively by further introductions to shoots in the Ukraine, Maldavia, and the Pre-Caucasus. In 1979 experiments began on establishing rabbits in Uzbekistan, and later in Lithuania. At each site, 2000-3000 animals were

released. Irrigation development in Central Asia provided opportunities for wild rabbits to become Iocally abundant in desert zones (Skulyatvev 1987).

A detailed picture of the distribution and abund- ance of the rabbit in France was produced by a 1977

survey of all gamekeepers by the Office National de la Chasse (Arthur et al. 1980). It showed, with other data (Arthur and Chapuis 1985), that rabbits are found in every departement (Fig. 3.5). They are less abundant and more dispersed in mountain areas (Jura, Alpes, Vosges, centre of the Massif Central, Pyrenees) and the north-eastern departements, per- haps discouraged by a high snowline, thin soils, too



The rabbit in continental Europe 31



Table 3.3 Density and size of rabbit warrens in various French landscapes

Region

Landscape

Area sampled

(ha)

Warrens per ha


Entrances per warren


Reference



Camargue woodland

60



2.6


8.6


Pages (1980)

Camargue salt-marsh

450

1.4

2.8

Rogers (1979)

Midi vines

40

1.5 *

3.0

Pages (1980)

Midi garriguet

25

0.2

1.3

Pages (1980)

Brittany heath

400

2.0

1.0

Chapuis (1979)

Sologne wood/cultivated

283

0.7**

3.0

Servan (1972)

Ile-de-France cultivated

200

0.4 **

2.2

Panaget (1983)

wood/cultivated

85

1.3 **

2.1

Arthur and Guenezan
















(unpublished)

parkland

250

2.5

24.0

Arthur and Guenezan
















(unpublished)

Vaucluse

fallow/woodland

38

2.6

15.0

Arthur and Guenezan
















(unpublished)


t Shrub-covered calcarious land.

* Warrens mostly in the brush beneath the vines.

** Warrens mostly in woods and hedges.



much woodland and, in the north-east of France, harsh winters. In contrast, they have high popula- tion densities in Brittany, Vendee, along the Rhone valley, and the Mediterranean coast, where there are smaller fields, more uncultivated land (heathland, garrigue-shrub-covered calcarious land) and many hedgerows. Rabbits are also abundant in Nord-Pas- de-Calais, Sologne, and around the cities of Paris and Lyon, where they are encouraged by many private hunts. The pattern of distribution has remained fairly consistent over the years, although abundance has varied.

As might be expected, landscape also determines the distribution and size of warrens. In most of France the density of warrens is less than 1.5 per hectare, each with a maximum of 3 entrances, but may reach 2.5 or more warrens per hectare, some with over 20 entrances (Table 3.3).

Since the introduction of myxomatosis in France

four national surveys have been made to monitor rabbit abundance (Giban 1956; ONC 1976; Arthur et al. 1980; Arthur and Guenezan 1986). Although the survey methods used differed (survey of some private hunts from 1920 to 1956, survey of game- keepers in 1977, sample survey of hunters in 1974 and 1983), the general trend is clear enough.

After the population crash in 1952, hunters ' bags remained small until 1956; then they slowly increased, more rapidly after 1970. In 1977, more than 15 million rabbits (about a quarter of the pre- myxomatosis figure) were shot. Thereafter data are li mited, but only 6.5 million were shot in 1983-a real reduction in total population since 1977 of around 55 per cent. But counting only the area of grassland, cultivated land, hedgerow and woodland edge, the number shot per 100 hectares per year was 36.4 in

1974 and 37.6 in 1977, dropping to 24.4 in 1983; a real reduction of about 35 per cent. However, the pattern of decline varied in different regions of the country (see below).

Variations in abundance of rabbits have been

observed since at least 1880, long before the advent of myxomatosis (Middleton 1934; Giban 1956; Tapper 1985). The explanations offered then were usually coccidiosis or climate; also, natural cycles may be inherent in the population dynamics of rabbits. Part of the reason for the rise and fall in the

1970s and 1980s might be that the years 1970-8 were, in France at least, climatically favourable for the



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