COMPOSITE SCORES OF ADAPTIVE UNITS AND GEOGRAPHIC DISTRIBUTION

In Table 11, procyonid species are arranged in descending order with

respect to the number of major climates that are included in their

geographic distributions (Hall and Kelson, 1959:878-897;

Poglayen-Neuwall, 1975; Kortlucke and Ramirez-Pulido, 1982; Nowak and

Paradiso, 1983:977-985). Composite scores ranged from a high of 1.47

for _Procyon lotor_ to a low of 0.39 for _Potos flavus_, whereas

_Nasua nasua_, _Nasua narica_, _Procyon cancrivorus_, and _Bassariscus

astutus_ had intermediate values ranging from 0.64 to 0.79 (Table 12).

Figure 8 demonstrates that there is a direct relationship between the

number of climates these species occupy and their composite scores.

Regression analysis (Y = 2.68.X + 0.24; where Y is number of climates,

and X is composite score) demonstrates a high degree of correlation

between these variables (R = 0.94) and indicates that 89% of the

variance in distribution can be explained by composite scores. The

various combinations of adaptations expressed by these species do,

therefore, play a role in delimiting their climatic (latitudinal)

distributions.

_Procyon lotor's_ normalized scores were higher in all categories than

those of other procyonids. _Procyon lotor_, therefore, possesses those

traits that have allowed it to become the premier climate generalist

of the procyonid family. As an adaptive unit, these traits provide

_Procyon lotor_ with the physiological and behavioral flexibility

required to take full advantage of a wide range of climates and

habitats, and its distribution verifies that it has done so. Even so,

it is probably not fair to assume that this species represents a

perfect physiological match with climate over its entire distribution.

_Procyon lotor_ is, in many respects, still a forest-dwelling species,

and its ability to expand its distribution into other habitats such as

prairie and desert may well be due, in part, to its use of behavior to

take advantage of favorable microclimates in otherwise hostile

environments (Bartholomew, 1958, 1987). This feature of _Procyon

lotor's_ biology needs to be further examined.

TABLE 12.--Normalized and composite scores for selected procyonids.

(H_{br} = ratio of measured to predicted basal metabolism (Table 7),

C_{mwr} = ratio of measured to predicted minimum thermal conductance

(Table 7), D_{dr} = ratio of food categories actually utilized by

each species to total food categories eaten by all six species

(calculated from Table 9), r_{maxr} = ratio of calculated to

expected r_{max} (Table 10).)

----------------------+----------------------------------------------

| Normalized scores

Species |---------------------------- Composite[a]

|H_{br}/C_{mwr} D_{dr} r_{maxr} score

----------------------+----------------------------------------------

_Procyon lotor_ | 0.95 0.95 2.52 1.47

_Bassariscus astutus_ | 0.80 0.33 1.24 0.79

_Nasua nasua_ | 0.48 0.33 1.11[b] 0.64

_Nasua nasua_ | 0.48 0.33 1.11[b] 0.64

_Nasua narica_ | 0.40 0.53 1.11 0.68

_Procyon cancrivorus_ | 0.55 0.33 1.32 0.73

_Potos flavus_ | 0.60 0.11 0.48 0.39

----------------------+----------------------------------------------

[a] Composite score = [(H_{br}/C_{mwr}) + D_{dr} + r_{maxr}]/3.

[b] Value calculated for _Nasua narica_ (Table 10) and used with the

assumption that it must be similar to the value for _Nasua nasua_.

All five species with low [.H]_{b}'s have composite scores less than

1.0 (Table 12; Figure 8). Four of these five, _Nasua nasua_, _Nasua

narica_, _Procyon cancrivorus_, and _Potos flavus_, have

H_{br}/C_{mwr} ratios that are 0.6 or less, which indicates they are

the least cold-tolerant procyonids (McNab, 1966). These four species

also are confined to either tropic, or tropic and subtropic climates

(Table 11). This suggests that these species share a common

thermoregulatory adaptation that represents a specialization to these

climates. Attendant with this adaptation, however, is a high cost of

thermoregulation at temperatures below their T_{lc}, and this must be

an important factor in limiting their distributions to tropic and

subtropic climates. Differences in their distributions within these

climates, therefore, must hinge more on differences in their D_{dr}

and r_{maxr} values than on differences in their H_{br}/C_{mwr}

ratios. This is supported by the fact that _Potos flavus_, which has

the lowest D_{dr} and r_{maxr} values, is confined to a single

climate, whereas _Nasua nasua_, _Nasua narica_, and _Procyon

cancrivorus_ each possess larger D_{dr} and r_{maxr} values and are

found in two climates. Thus, _Potos flavus_, with its highly

specialized diet and low reproductive potential, is the most

ecologically specialized of these procyonids, and its distribution is

limited to the single climate that can provide its requirements.

_Nasua nasua_, _Nasua narica_, and _Procyon cancrivorus_ are less

specialized and thus show more ecological flexibility in their

distributions.

[Illustration: FIGURE 8.--Relationship between number of climates

in which a species is found and its composite score. Symbols for

_Nasua nasua_ overlap at coordinates (0.64, 2). Solid line

represents linear regression of climates (Y) on composite scores

(X): Y = 2.68.X + 0.24; R = 0.94.]

_Bassariscus astutus_, the other species with low [.H]_{b}, is found

in three climates, which indicates that it has greater ecological

flexibility than _Nasua nasua_, _Nasua narica_, or _Procyon

cancrivorus_. D_{dr} and r_{maxr} are comparable for these four

species (Table 12). This suggests that the greater ecological

flexibility of _Bassariscus astutus_ is derived largely from its

greater cold tolerance. _Bassariscus astutus_ has a more insulative

pelt than these other procyonids (C_{mwr} = 0.85; Table 7), so its

H_{br}/C_{mwr} ratio is higher (0.80; Table 12). This, and its greater

capacity for evaporative cooling (Chevalier, 1985), allows

_Bassariscus astutus_ to take advantage of a wider range of thermal

environments than these other species. However, even with its higher

H_{br}/C_{mwr} ratio, the composite score for _Bassariscus astutus_

is not much different than those for _Nasua nasua_, _Nasua narica_,

and _Procyon cancrivorus_ (Table 12). Consequently, _Bassariscus

astutus_ is found in more climates than would be predicted for it on

the basis of its composite score (Figure 8). This suggests that either

the H_{br}/C_{mwr} ratio carries greater weight in determining

distribution than is reflected in this analysis, or as has been

described for some other species (Bartholomew, 1958, 1987),

_Bassariscus astutus_ may extend its distribution farther than

expected via use of its behavior. In either case, for procyonids with

low [.H]_{b}, _Bassariscus astutus_ represents the pinnacle of

adaptation for climate generalization.

Metabolic Adaptation to Climate and Distribution of the Raccoon Procyon Lotor and Other Procyonidae · 第 36 / 127 题