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160 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY
Recent investigations by neontologists into intraspecific
variation in size and allometry, seen across both space and recent
time, emphasize the need for a broader-based approach to
variation seen in fossils. Zink and Remsen's (1986) review of
the relevance of geographic variation to evolution, and case
studies such as those by Johnston and Selander (1964), Grant
(1971, 1986), Fleischer and Johnson (1982), James (1983), and
Dennison and Barker (1991), demonstrate that within-species
morphological differences in birds are frequently observed
across space, and that these differences may evolve over relatively
short spans of time. Analyses of present-day intraspecific
geographic variation therefore give an insight into the spectrum
of variation that may have occurred over recent geological time
in species.
A notable exception to the lack of application of these neontological
studies to fossils or subfossils is the study by Ericson
(1987), wherein subfossil Common Eider {Somateria mollissima
Linnaeus) from Scandinavia were compared to a variety of
different extant subspecies and were shown to differ significantly.
In this instance various environmental criteria were
considered causal, including climatic change and anthropogenic
effects; the author preferred the latter as an explanation.
MATERIALS AND METHODS
The present study has attempted to bring together a widely
distributed body of data on both species of Lagopus from the
present-day western Palearctic. These data were used for biometric
comparison with a broader range of geographically and
temporally separated fossils than has been previously achieved.
All but the British and Belgian fossil samples have been metrically
studied by others, although no one has examined all these
samples together. Those that are newly analyzed herein are
from Pin Hole Cave (Manchester City Museum), Merlin's
Cave (University of Bristol Speleological Society Museum),
Westbury-sub-Mendip (Paleontology Department, Natural History
Museum, London), and Remouchamps (Musee Royaux
d'Art d'Histoire, Bruxelle).
The tarsometatarsus is invariably the most frequently occurring
skeletal element of medium-sized Galliformes in European
cave assemblages (Mourer-Chauvire, 1983) and is thus the
main element dealt with herein. The relative abundance of this
bone is fortunate because it is one of the elements most easily
identified as either Lagopus lagopus or L. mutus (Kraft, 1972;
Mourer-Chauvire, 1975a; Bocheriski, 1985). There appear to
be no details of morphology that can aid determination, and the
two species are identified simply on the basis of size: L. lagopus
is consistently larger than L. mutus (see Figure 2). Many
other postcranial bones present greater problems because they
overlap considerably in size and therefore make analysis more
complicated.
Table 1 lists all the modern skeletal samples of both species,
including two populations of L. lagopus scoticus (Latham), one
from Scotland and one from Derbyshire, England; samples of
TABLE 1.—Mensural data for the tarsometatarsus of modern and fossil samples
of both Lagopus lagopus and L. mutus. See Table 2 for fossil site locations.
Sites that have no mean and have question marks instead of a minimum value
forZ,. lagopus and a maximum value fori, mutus are such because insufficient
tarsometatarsal length difference was present between specimens to define the
respective upper limits of L. lagopus and the lower limits of L. mutus (see Figure
2). (GL=greatest length; KB=shaft width; w=number of specimens.)
Samples
MODERN
Lagopus lagopus lagopus (Scandinavia)
Lagopus lagopus lagopus (Russia)
Lagopus lagopus scoticus (Derbyshire,
England)
Lagopus lagopus scoticus (Scotland)
Lagopus lagopus major (Kazakhstan)
Lagopus mutus mutus (Scandinavia)
Lagopus mutus millaisi (Scotland)
Lagopus mutus helveticus (French Alps)
Lagopus mutus islandorum (Iceland)
FOSSIL
Mamutowa Cave, Lagopus lagopus
Mamutowa Cave, Lagopus mutus
Remouchamps, Lagopus lagopus
Merlin's Cave, Lagopus lagopus
Merlin's Cave, Lagopus mutus
La Balme-les-Grottes, Lagopus lagopus
La Balme-les-Grottes, Lagopus mutus
La Colombiere, Lagopus lagopus
La Colombiere, Lagopus mutus
Pin Hole Cave, Lagopus lagopus
Pin Hole Cave, Lagopus mutus
La Fage, Lagopus lagopus noaillensis
La Fage, Lagopus mutus correzensis
Westbury-sub-Mendip, Lagopus sp.
Rebielice Krolewskie, Lagopus atavus
Minimum-Maximum (n
GL 37.04-42.1 («=11)
KB 2.96-3.34 («= 11)
GL 38.1-42.88 (n=6)
KB 2.84-3.44 (n=5)
GL 38.6-^3.58 (n= 19)
KB 2.94-3.98 (»= 19)
GL 38.38^14.06 («=9)
KB 3.1-3.6 (n=9)
GL 45.32 (n=l)
KB3.64(w=l)
GL 29.44-34.08 («=2)
KB 2.66-2.74 (n=2)
GL 33.22-35.9 (n=5)
KB 2.88-3.12 («=5)
GL 31.8-35.7 (n= 16)
KB 2.46-2.94 (K= 16)
GL 30.18-34.82 (n=2)
KB 2.58-2.86 (n=2)
GL ?-39.84
KB?
GL 32.1-?
KB?
GL 39.44-41.64 («=3)
KB 3.24-3.4 (n=3)
GL ?-40.28
KB?
GL 29.94-?
KB?
GL 36.36-38.8 (n= 10)
KB 3-3.48 (n= 10)
GL 32.24-32.3 (n=2)
KB 3.2 (n=2)
GL 35.2-40.6 (/i=30)
KB 3.04-4.08 («=30)
GL 29.26-33.72 (n=30)
KB 2.6-3.36 (w=30)
GL 36.46-41.32 (n=22)
KB 3.06-3.88 («=22)
GL 30.54-32.98 («=27)
KB 2.74-3.48 (n=27)
GL 36.52-39.3 (n=l)
KB 3.22-3.44 («=7)
GL 31.7-34.8 (n=9)
KB 2.74-3.42 (n=9)
GL-
KB 3.43 (n=2)
GL-
KB3.96(«=1)
Mean
38.95
3.18
40.61
3.19
41.1
3.24
41.1
3.36
-
-
31.76
2.7
34.69
3
33.06
2.71
32.5
2.72
-
-
-
-
40.19
3.3
- --
37.75
3.22
32.27
3.2
38.21
3.4
31.39
3
38.66
3.41
31.85
3.01
38.07
3.33
33.38
3.09
-
-
-
-
L. lagopus lagopus from Scandinavia and Russia; and an individual
skeleton of L. lagopus major Lorenz. Lagopus mutus is
represented by skeletons from Scotland {L. m. millaisi Hartert),
Iceland {L. m. islandorum Faber), Scandinavia {L. m. mutus),