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American Bison - Buffalo Field Campaign

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season: Wolfe and Kimball (1989) reported an increase in the<br />

percentage of calves from 10.2% in late May to 12.2% in late<br />

July (i.e., count too early and you may miss some).<br />

Similarly, segregation of age and sex classes may influence<br />

estimates of population composition. Meagher (1973) reported<br />

that calves formed 20% of mixed age and mixed sex bison<br />

herds, but 11% of the total Yellowstone bison population.<br />

Other biases are also possible. Carbyn et al. (1998) reported<br />

an unweighted average of 36 calves per 100 adult females<br />

for bison in Delta Area of WBNP for 1989-1996 (Table 6.3),<br />

while others reported between 20 and 30 calves per 100 adult<br />

females for the same area and during the same time period<br />

(Bradley and Wilmshurst 2005). Similarly, Carbyn et al. (1998)<br />

reported an average of 20 yearlings per 100 adult females for<br />

this time period, while others reported more than 10 yearlings<br />

per 100 adult females for only one of those years (Bradley and<br />

Wilmshurst 2005). Thus, composition estimates need to be<br />

interpreted with considerable caution and would benefit by<br />

inclusions of confidence intervals.<br />

Few data sets permit evaluation of reproductive success and<br />

survival of young in relation to population densities (Table 6.4).<br />

The higher ratios of calves and yearlings per 100 adult females<br />

in the Mink Lake area of WBNP compared to MBS (Table 6.4)<br />

reflect differences between increasing and declining populations<br />

(Larter et al. 2000). Lower calf and yearling to adult female ratios<br />

were linked to a period of population decline at WBNP (Bradley<br />

and Wilmshurst 2005). Reynolds et al. (2003) reported density<br />

dependent fecundity in bison at EINP.<br />

Information on the age structure of free-ranging bison<br />

populations not subjected to regular culling is limited. Wood<br />

bison at the MBS were assigned to age and sex classes in July<br />

1993: calves and yearlings were not assigned to sex classes,<br />

all females two or more years old were assigned to a single<br />

category, and males more than two years old were assigned to<br />

one of four age categories following Komers et al. (1992). Here,<br />

the population age structure is presented with an assumption<br />

of an equal sex ratio in calves and yearlings (Figure 6.2).<br />

Irrespective of the sex, the relatively low numbers of calves and<br />

yearlings suggest a low recruitment rate (Figure 6.2).<br />

6.3.2 Reproduction<br />

The age of first reproduction is sensitive to nutritional condition<br />

and, therefore, highly variable. The proportion of females calving<br />

as two-year-olds (conceiving as one-year-olds) ranges between<br />

4-12% (Table 6.5). However, female bison typically enter oestrus<br />

as two-year-olds, and give birth to their first calf at three years<br />

(Table 6.5). Mature females in some populations reproduce<br />

each year (Rutberg 1984; Shaw and Carter 1989; Wolff 1998),<br />

although in other populations mature females may not breed<br />

in some years (Fuller 1962; Green 1990; Halloran 1968; Soper<br />

1941; Van Vuren and Bray 1986; Wolfe et al. 1999). This is<br />

particularly true of females breeding as two- to four-year-olds<br />

(Green 1990). Fuller (1962) noted that for wood bison in the Hays<br />

Camp area of WBNP, 21% of the females more than three years<br />

old at the time of parturition were lactating, but non-pregnant,<br />

while the same was true for 9% of the females in the Lake Claire<br />

area of the park. This proportion may vary within the same<br />

population at different densities of bison and other ungulate<br />

species relative to forage conditions (Halloran 1968; Shaw and<br />

Carter 1989). The young born to females following a year of not<br />

breeding were larger and more fecund than the young of females<br />

who bred the previous year (Green and Rothstein 1991). Females<br />

continue to breed until more than 16 years of age (Green 1990).<br />

<strong>Bison</strong> are typically monoparous, with twinning reported only<br />

occasionally (Reynolds et al. 2003).<br />

Male bison maintained on supplemental feed are physiologically<br />

capable of breeding as early as 16 months of age (Helbig et al.<br />

2007), and those not receiving diet supplements may breed at<br />

two to three years old (Maher and Byers 1987). However, males<br />

generally do not breed until they are five or six years old and<br />

large enough to compete with older and more experienced bulls<br />

(Fuller 1960; Komers et al. 1994; Meagher 1973; Rothstein and<br />

Griswold 1991).<br />

The age of first successful reproduction may be modified<br />

by disease in bison of the Jackson, Yellowstone and GWBE<br />

populations. More than 90% of the first pregnancies were<br />

lost in brucellosis infected captive female bison (Davis et al.<br />

1990; 1991). In free-ranging bison, the impact of brucellosis<br />

on the age of first successful reproduction will vary with<br />

the proportion of first time breeders in the population, the<br />

proportion of those breeders infected with brucellosis, and<br />

the severity of the infection (Bradley and Wilmshurst 2005).<br />

Diseases may also modify reproductive performance of older<br />

females. At WBNP, both tuberculosis (BTB) and brucellosis<br />

may impact the reproductive success of females of all age<br />

classes within select population segments (Joly and Messier<br />

2004; 2005). In two population segments of wood bison at<br />

WBNP, infection with brucellosis or BTB alone did not impact<br />

pregnancy status, but infection with both diseases reduced the<br />

probability of pregnancy by 30% (Joly and Messier 2005). In a<br />

third population segment, infection with BTB alone reduced the<br />

probability of pregnancy by 75% (Joly and Messier 2005).<br />

6.3.3 Mortality factors and survival<br />

Proximate causes of mortality in contemporary wood bison<br />

herds include wolf predation and the exotic diseases brucellosis<br />

and BTB (Fuller 1962; Calef 1984; Carbyn et al. 1993; Joly and<br />

Messier 2001, 2004; 2005; Wilson et al. 1995 in Bradley and<br />

Wilmshurst 2005). In addition, some wood bison succumb to<br />

irregular outbreaks of anthrax (Bacillus anthracis) (Gates et al.<br />

<strong>American</strong> <strong>Bison</strong>: Status Survey and Conservation Guidelines 2010 49

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