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Intraspecific Variation in Modern and Quaternary<br />

European Lagopus<br />

ABSTRACT<br />

Skeletal proportions of modem European populations of Lagopus<br />

lagopus (Linnaeus) and L. mutus (Montin) from Britain, Iceland,<br />

Scandinavia, northern Russia, and the Alps are compared<br />

with their Quaternary fossil counterparts from Britain, Poland,<br />

France, and Belgium. Lagopus lagopus and L. mutus from most<br />

pre-Holocene deposits are found to differ allometrically from<br />

modem samples. This difference is best seen in the tarsometatarsus,<br />

which often is more robust in both species in the Pleistocene<br />

and in turn may reflect greater body weight. Possible correlations<br />

between this phenomenon and the climatic and ecological conditions<br />

of the past, as well as the possibility that these birds were less<br />

sedentary, are discussed.<br />

Introduction<br />

Previous workers, such as Newton (1924), Mourer-Chauvire<br />

(1975a, 1975b), Janossy (1974, 1976), Bochehski (1974,<br />

1985, 1991), Harrison (1980), Potapova (1986), and Bocheriski<br />

and Tomek (1994) have described differently sized<br />

and proportioned postcranial bones of both Lagopus mutus<br />

(Montin) and Lagopus lagopus (Linnaeus) from the fossil<br />

record of Europe. Specifically, samples of both species from<br />

the Pleistocene were seen to differ in size and allometry from<br />

skeletons of their recent counterparts. These allometric differences<br />

were seen in the relative proportions of their tarsometatarsi,<br />

carpometacarpi, phalanx 1 digit III pedis, and phalanx 1<br />

digit II alae, as well as in cranial elements.<br />

Newton, who was probably the first to note the occurrence of<br />

anomalously proportioned Lagopus fossils of Pleistocene age,<br />

believed that a third species had existed (Newton, 1924). He referred<br />

to this species, found at Merlin's Cave, a late Pleistocene<br />

site in Britain, as a "small ptarmigan" and never named it as a<br />

distinct form, which is how many authors have dealt with these<br />

John R. Stewart, The McDonald Institute for Archaeological <strong>Res</strong>earch,<br />

Downing Street, Cambridge CB2 3ER, England.<br />

John R. Stewart<br />

159<br />

anomalies since. Exceptions are the species Lagopus atavus<br />

Janossy (1974) from the late Pliocene of Poland (Janossy,<br />

1976; Bocheriski, 1991) and the subspecies L. lagopus noaillensis<br />

Mourer-Chauvire (1975a) and L. mutus correzensis<br />

Mourer-Chauvire (1975a) from La Fage, a Rissian (middle<br />

Pleistocene) site in France.<br />

Lagopus lagopus noaillensis and L. mutus correzensis were<br />

described by Mourer-Chauvire as being distinguished by the<br />

robustness of their tarsometatarsi as compared to modern populations.<br />

She also plotted the mean lengths and shaft widths of<br />

tarsometatarsi of both species from La Colombiere and Gigny,<br />

two assemblages of different ages from the last glacial of<br />

France, together with the two named subspecies from La Fage.<br />

This showed that they, too, had relatively robust tarsometatarsi,<br />

although their lengths varied, producing allometric-shape variation.<br />

The most detailed study of Lagopus fossils to date was that<br />

by Bocheriski (1974). In his book he compared a number of<br />

Polish fossil populations of both species from the last glaciation<br />

with samples from much of their modern European distribution.<br />

He concluded that the fossil birds possessed longer carpometacarpi<br />

and humeri but shorter tarsometatarsi than their modern<br />

counterparts, and that the articular ends of the humeri, tarsometatarsi,<br />

and coracoids were more massive. Bocheriski interpreted<br />

the differences in the wing-bone lengths as indicating<br />

that the primary feathers of both species had become shorter,<br />

thus reducing wing-surface area over time. This, together with<br />

the change in bone robustness, especially that of the coracoid,<br />

indicated to him that the two species had maintained, or had<br />

only slightly reduced, their body size over time. Later, Bocheriski<br />

(1985) showed that Pleistocene L. lagopus in Poland<br />

had longer distal-wing bones (carpometacarpi and phalanx 1<br />

digit II alae) and shorter distal-leg elements (tarsometatarsi and<br />

phalanx 1 digit III pedis) when compared to modern birds. Subsequently,<br />

Bocheriski and Tomek (1994) focused on the relative<br />

lengths of postcranial bones of Pleistocene L. mutus in<br />

Austria and showed that they differed from modern alpine birds<br />

in having shorter tarsometatarsi but longer carpometacarpi.

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