Spatial Population Structure of Yellowstone Bison

Abstract: Increases in Yellowstone National Park, USA, bison (Bison bison) numbers and shifts in seasonal distribution have resulted in more frequent movements of bison beyond park boundaries and development of an interagency management plan for the Yellowstone bison population. Implementation of the plan under the adaptive management paradigm requires an understanding of the spatial and temporal structure of the population. We used polythetic agglomerative hierarchical cluster analysis of radiolocations obtained from free-ranging bison to investigate seasonal movements and aggregations. We classified radiolocations into 4 periods: annual, peak rut (15 Jul-15 Sep), extended rut (1 Jun-31 Oct), and winter (1 Nov-31 May). We documented spatial separation of Yellowstone bison into 2 segments, the northern and central herds, during all periods. The estimated year-round exchange rate (4.85-5.83%) of instrumented bison varied with the fusion strategy employed. We did not observe exchange between the 2 segments during the peak rut and it varied during the extended rut (2.15-3.23%). We estimated a winter exchange of 4.85-7.77%. The outcome and effectiveness of management actions directed at Yellowstone bison may be affected by spatial segregation and herd affinity within the population. Reductions based on total population size, but not applied to the entire population, may adversely affect one herd while having little effect on the other. Similarly, management actions targeting a segment of the population may benefit from the spatial segregation exhibited.     

Predicting bison migration out of Yellowstone National Park using bayesian models

Long distance migrations by ungulate species often surpass the boundaries of preservation areas where conflicts with various publics lead to management actions that can threaten populations. We chose the partially migratory bison (Bison bison) population in Yellowstone National Park as an example of integrating science into management policies to better conserve migratory ungulates. Approximately 60% of these bison have been exposed to bovine brucellosis and thousands of migrants exiting the park boundary have been culled during the past two decades to reduce the risk of disease transmission to cattle. Data were assimilated using models representing competing hypotheses of bison migration during 1990-2009 in a hierarchal bayesian framework. Migration differed at the scale of herds, but a single unifying logistic model was useful for predicting migrations by both herds. Migration beyond the northern park boundary was affected by herd size, accumulated snow water equivalent, and aboveground dried biomass. Migration beyond the western park boundary was less influenced by these predictors and process model performance suggested an important control on recent migrations was excluded. Simulations of migrations over the next decade suggest that allowing increased numbers of bison beyond park boundaries during severe climate conditions may be the only means of avoiding episodic, large-scale reductions to the Yellowstone bison population in the foreseeable future. This research is an example of how long distance migration dynamics can be incorporated into improved management policies.     

Genetic Analysis of the Henry Mountains Bison Herd

Wild American plains bison (Bison bison) populations virtually disappeared in the late 1800s, with some remnant animals retained in what would become Yellowstone National Park and on private ranches. Some of these private bison were intentionally crossbred with cattle for commercial purposes. This forced hybridization resulted in both mitochondrial and nuclear introgression of cattle genes into some of the extant bison genome. As the private populations grew, excess animals, along with their history of cattle genetics, provided founders for newly established public bison populations. Of the US public bison herds, only those in Yellowstone and Wind Cave National Parks (YNP and WCNP) appear to be free of detectable levels of cattle introgression. However, a small free-ranging population (~350 animals) exists on public land, along with domestic cattle, in the Henry Mountains (HM) of southern Utah. This isolated bison herd originated from a founder group translocated from YNP in the 1940s. Using genetic samples from 129 individuals, we examined the genetic status of the HM population and found no evidence of mitochondrial or nuclear introgression of cattle genes. This new information confirms it is highly unlikely for free-living bison to crossbreed with cattle, and this disease-free HM bison herd is valuable for the long-term conservation of the species. This bison herd is a subpopulation of the YNP/WCNP/HM metapopulation, within which it can contribute significantly to national efforts to restore the American plains bison to more of its native range.     

Detection of mitochondrial DNA from domestic cattle in bison on Santa Catalina Island

In 1924, 14 American bison (Bison bison) were introduced to Santa Catalina Island, California and sporadically supplemented thereafter with additional animals. To reduce the herd and its impact on native vegetation, over 2000 animals have been exported during the past four decades. Today, the herd is estimated to contain around 250 individuals. Genetic analysis was performed on 98 animals removed from the island in 2004. Forty-four samples (45%) had domestic cattle mitochondrial DNA (mtDNA), 12 (12%) had previously reported bison haplotypes and 42 (43%) had a new haplotype differing by one base pair from a previously reported bison haplotype. A complement of five restriction enzymes was found to be useful in identifying bison with domestic cattle mtDNA.     

Reproduction and Survival of Yellowstone Bison

ABSTRACT The conservation of bison (Bison bison) from near extinction to >4,000 animals in Yellowstone National Park has led to conflict regarding overabundance and potential transmission of brucellosis (Brucella abortus) to cattle. We estimated survival and birth rates from 53 radiocollared adult female bison during 1995–2001, and we used calf:adult (C:A) ratios to estimate reproduction with the combined effects of pregnancy, fetal loss, and neonatal mortality during 1970–1997. Annual survival of adult females was high (0.92; 95% CI = 0.87-0.95) and constant. Birth rates differed by brucellosis status and age. Birth rates were 0.40 calves per female (95% CI = 0.15-0.65) for brucellosis-positive 3 year olds, 0.63 (95% CI = 0.39-0.87) for individuals testing negative, and 0.10 (95% CI = 0.00-0.24) for individuals contracting brucellosis that birth year (sero-converters). Birth rates were 0.64 (95% CI = 0.52-0.76) for brucellosis-positive individuals ≥4 years old, 0.81 (95% CI = 0.73-0.89) for brucellosis-negative individuals, and 0.22 (95% CI = 0.00-0.46) for sero-converters. Spring C:A ratios were negatively correlated with snow pack (β = −0.01 to −0.03, R² = 0.26-0.60, P < 0.05). Growth rate was highly elastic to adult survival (0.51), and juvenile survival (0.36) was 3 times more elastic than fecundity (0.12). Simulations suggested brucellosis eradication via vaccination would result in increased birth rates and a 29% increase in population growth (γ = 1.09), possibly leading to more bison movements outside the park. Our results will help park managers evaluate bison population dynamics and explore consequences of management actions and disease control programs.     

Body mass and measurements of the European bison during postnatal development

Body mass of the European bisonBison bonasus (Linnaeus, 1758) was determined for 152 animals (86 males, 66 females) from captive breeding centre in Białowieża and for 492 (248, 244) animals from the free-ranging herd in the Białowieża Forest. Body measurements were taken for a total of 472 bison (204, 268). Age-related changes in body mass and measurements of 375 bison (165, 210) were analysed. Bison body mass was significantly correlated with age. Captive bison were heavier than those from a free-ranging herd. Measurements showed the changes in bison body structure in the postnatal development. All measurements were highly correlated with age and mass of the animals and increased most intensively during the first year of life. Sex-related differences in body mass and measurements were most pronounced in bison ≥ 3 years old. Physical development of females ends at the age of 5 years, and that of males at the age of 7 years.     

Resistance to malignant catarrhal fever in American bison (Bison bison) is associated with MHC class IIa polymorphisms

The Rhadinovirus ovine herpesvirus-2 (OvHV-2) is the most common causative agent of malignant catarrhal fever (MCF) in clinically susceptible ruminants including cattle and bison. American bison (Bison bison) are highly susceptible to clinical MCF. Nevertheless, approximately 20% of bison on ranches or in feedlots become infected with the virus without developing clinical disease. Defining the genetic basis for differences in susceptibility between bison could facilitate development of improved control strategies for MCF. One genetic region that influences susceptibility to infectious diseases is the major histocompatibility complex (MHC). In this study, a Bison bison (Bibi) DRB3 oligonucleotide microarray was used to type 189 bison from 10 herds where MCF outbreaks had occurred. Binary logistic regression was used to classify DRB3 alleles as resistant (R), susceptible (S) or neutral (N). Animals were reclassified using six DRB3 genotype categories: N/N, N/R, N/S, R/S, R/R and S/S. Analysis of homogeneity across herds showed that there was a herd effect. Consequently, a penalized logistic regression model was run with herd and genotype categories as the explanatory variables. The R/R genotype was associated with resistance to MCF (P = 0.0327), while the S/S genotype was associated with clinical MCF (P = 0.0069). This is the first evidence that MHC class IIa polymorphism is associated with resistance or susceptibility to OvHV-2-induced MCF.     

Density-Dependent Growth in Invasive Lionfish (Pterois volitans)

Direct demographic density dependence is necessary for population regulation and is a central concept in ecology, yet has not been studied in many invasive species, including any invasive marine fish. The red lionfish (Pterois volitans) is an invasive predatory marine fish that is undergoing exponential population growth throughout the tropical western Atlantic. Invasive lionfish threaten coral-reef ecosystems, but there is currently no evidence of any natural population control. Therefore, a manipulative field experiment was conducted to test for density dependence in lionfish. Juvenile lionfish densities were adjusted on small reefs and several demographic rates (growth, recruitment, immigration, and loss) were measured throughout an 8-week period. Invasive lionfish exhibited direct density dependence in individual growth rates, as lionfish grew slower at higher densities throughout the study. Individual growth in length declined linearly with increasing lionfish density, while growth in mass declined exponentially with increasing density. There was no evidence, however, for density dependence in recruitment, immigration, or loss (mortality plus emigration) of invasive lionfish. The observed density-dependent growth rates may have implications for which native species are susceptible to lionfish predation, as the size and type of prey that lionfish consume is directly related to their body size. The absence of density-dependent loss, however, contrasts with many native coral-reef fish species and suggests that for the foreseeable future manual removals may be the only effective local control of this invasion.     

Body Size Variations in Caribou Ecotypes and Relationships With Demography

ABSTRACT In many vertebrates size is one of the most influential and variable individual characteristics and a strong determinant of reproductive success. Body size is generally density dependent and decreases when intraspecific competition increases. Frequent and long-distance movements increase energy expenditures and, therefore, may also influence body size, particularly in highly mobile species. Caribou (Rangifer tarandus, also known as reindeer) exhibit tremendous variation in size and movements and thus represent an excellent candidate species to test the relationships between body size, population size, and movements. We analyzed body measurements of adult female caribou from 7 herds of the Québec-Labrador Peninsula, Canada, and we related their morphology to population size, movements, and annual ranges. The herds represented 3 ecotypes (migratory, montane, and sedentary). Ecotypes and herds differed in size (length), shape (roundness), and movements. The sedentary ecotype was larger and moved 4 to 7 times less than the migratory ecotype in the 1990s. At the start of a demographic growth period in the early 1960s, migratory caribou from the Rivière-George (hereafter George) herd had longer mandibles than caribou of the sedentary ecotype. Mandible length in the George herd declined in the 1980s after rapid population growth, while individuals performed extensive movements and the herd's annual range increased. Migratory caribou then became shorter than sedentary caribou. After the George herd decline in the 1990s, mandible length increased again near levels of the 1980s. Caribou from the migratory Rivière-aux-Feuilles herd later showed a similar decline in mandible length during a period of population growth, associated with longer movements and increasing annual range. We hypothesize that the density-dependent effect observed on body size might have been exerted through summer habitat degradation and movement variations during herd growth. Our study has 2 important implications for caribou management: the distinctiveness of different populations and ecotypes, and the correlations between population trajectories and changes in body condition and habitat.     

Review of the 2012 Epizootic Hemorrhagic Disease Outbreak in Domestic Ruminants in the United States

An unusually large number of cases of Epizootic hemorrhagic disease (EHD) were observed in United States cattle and white-tailed deer in the summer and fall of 2012. USDA APHIS Veterinary Services area offices were asked to report on foreign animal disease investigations and state diagnostic laboratory submissions which resulted in a diagnosis of EHD based on positive PCR results. EHD was reported in the following species: cattle (129 herds), captive white-tailed deer (65 herds), bison (8 herds), yak (6 herds), elk (1 herd), and sheep (1 flock). A majority of the cases in cattle and bison were found in Nebraska, South Dakota, and Iowa. The majority of cases in captive white-tailed deer were found in Ohio, Iowa, Michigan, and Missouri. The most common clinical sign observed in the cattle and bison herds was oral lesions. The major observation in captive white-tailed deer herds was death. Average within-herd morbidity was 7% in cattle and bison herds, and 46% in captive white-tailed deer herds. The average within-herd mortality in captive white-tailed deer herds was 42%.     

Interchange and Overlap Among Four Adjacent Arctic Caribou Herds

Barren ground caribou (Rangifer tarandus granti) are distributed in herds that seasonally use specific geographic regions within an annual range, with varying levels of fidelity during different periods (e.g., calving, insect relief, wintering). As a result, caribou management is generally tailored to individual herds that often range across administrative boundaries. Herd ranges can shift over time, seasonal ranges of adjacent herds often overlap, herds merge, and there is often little genetic differentiation among adjacent herds. If substantial herd interchange occurs, it would have important management implications by influencing estimates of herd size, herd composition, and harvest rates. We compiled satellite telemetry data from 2003–2015 for 4 large arctic caribou herds to quantify herd interchange rates. We calculated a metric of herd interchange based on the relationship of caribou locations to typical weekly herd ranges (all yrs combined) and the distance to other radio-collared caribou from each of the 4 herds (yr specific). Although herd membership cannot always be clearly defined based on location, this metric provides an objective measure of the strength of evidence of herd membership that can be used to make comparisons among herds and time periods. We also calculated herd overlap and quantified how it varied throughout the year. Herd interchange was rare in the 2 larger herds, generally occurring when caribou overwintered with an adjacent herd, whereas herd interchange from the 2 smaller herds was more frequent and could last longer than a year. Although sample sizes were limited, there were no clear patterns in herd interchange with year or annual herd size. The 2 smaller herds had large seasonal overlap with adjacent herds, suggesting that herd interchange may be related to spatiotemporal herd overlap and relative herd size. Our results can help managers understand herd interchange and overlap to make management decisions, interpret research results, and develop more accurate population models. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Patterns of genetic variation in US federal bison herds

Like many wide‐ranging mammals, American bison (Bison bison) have experienced significant range contraction over the past two centuries and are maintained in artificially isolated populations. A basic understanding of the distribution of genetic variation among populations is necessary to facilitate long‐term germplasm preservation and species conservation. The 11 herds maintained within the US federal system are a critically important source of germplasm for bison conservation, as they include many of the oldest herds in the USA and have served as a primary resource for the establishment of private and public herds worldwide. In this study, we used a panel of 51 nuclear markers to investigate patterns of neutral genetic variation among these herds. Most of these herds have maintained remarkably high levels of variation despite the severe bottleneck suffered in the late 1800s. However, differences were noted in the patterns of variation and levels of differentiation among herds, which were compared with historical records of establishment, supplementation, herd size, and culling practices. Although some lineages have been replicated across multiple herds within the US federal system, other lineages with high levels of genetic variation exist in isolated herds and should be considered targets for the establishment of satellite herds. From this and other studies, it is clear that the genetic variation represented in the US federal system is unevenly distributed among National Park Service and Fish and Wildlife Service herds, and that these resources must be carefully managed to ensure long‐term species conservation.     

Contrasting Results of Culture-Dependent and Molecular Analyses of Mycobacterium avium subsp. paratuberculosis from Wood Bison

Reduced to near extinction in the late 1800s, a number of wood bison populations (Bison bison athabascae) have been re-established through reintroduction initiatives. Although an invaluable tool for conservation, translocation of animals can spread infectious agents to new areas or expose animals to pathogens in their new environment. Mycobacterium avium subsp. paratuberculosis, a bacterium that causes chronic enteritis in ruminants, is among the pathogens of potential concern for wood bison management and conservation. In order to inform translocation decisions, our objectives were to determine the M. avium subsp. paratuberculosis infection status of wood bison herds in Canada and to culture and genetically characterize the infective strain(s). We tested fecal samples from bison (n = 267) in nine herds using direct PCR for three M. avium subsp. paratuberculosis-specific genetic targets with different copy numbers within the M. avium subsp. paratuberculosis genome. Restriction enzyme analysis (REA) and sequencing of IS1311 were performed on seven samples from five different herds. We also evaluated a panel of different culture conditions for their ability to support M. avium subsp. paratuberculosis growth from feces and tissues of direct-PCR-positive animals. Eighty-one fecal samples (30%) tested positive using direct IS900 PCR, with positive samples from all nine herds; of these, 75% and 21% were also positive using ISMAP02 and F57, respectively. None of the culture conditions supported the growth of M. avium subsp. paratuberculosis from PCR-positive samples. IS1311 REA and sequencing indicate that at least two different M. avium subsp. paratuberculosis strain types exist in Canadian wood bison. The presence of different M. avium subsp. paratuberculosis strains among wood bison herds should be considered in the planning of translocations.     

Marker genotypes and population admixture and their association with body weight, height and relative body mass in United States federal bison herds

Elucidating genetic influences on bison growth and body composition is of interest, not only because bison are important for historical, cultural, and agricultural reasons, but also because their unusual population history makes them valuable models for finding influential loci in both domestic cattle and humans. We tested for trait loci associated with body weight, height, and bison mass index (BMI) while controlling for estimated ancestry to reduce potential confounding effects due to population admixture in 1316 bison sampled from four U.S. herds. We used 60 microsatellite markers to model each phenotype as a function of herd, sex, age, marker genotypes, and individual ancestry estimates. Statistical significance for genotype and its interaction with ancestry was evaluated using the adaptive false discovery rate. Of the four herds, two appeared to be admixed and two were nonadmixed. Although none of the main effects of the loci were significant, estimated ancestry and its interaction with marker loci were significantly associated with the phenotypes, illustrating the importance of including ancestry in the models and the dependence of genotype-phenotype associations on background ancestry. Individual loci contributed approximately 2.0% of variation in weight, height, and BMI, which confirms the utility and potential importance of adjusting for population stratification.     

The feral goats of Kielderhead Moor

Herds of feral goats are present along the high ground of the Scottish Borders at the College, Hindhope, Kielderhead Moor (the study area), Bewcastle Fell and Roan Fell.
In the study area the nannies remained solitary, or in the company of their yearlings on the steep ground below the high moor throughout the winter. After the birth of their kids in March two female herds were formed each with a herd billy which came from the two male herds which were present through the winter on the high moor. The male herds left the study area in the spring and their ranges are not known. The two female herds maintained separate ranges on the high moor until the end of August when they combined into one large herd and were joined by a strange herd of billies which had not previously been seen.
After the first three or four days of its life the kid is left in a hiding place for most of the day whilst its mother ranges widely grazing with the herd. The kids join the herd when they are about six weeks old.
The Kielderhead Moor goats had a recruitment rate of 0–29/nanny/year and the females have a further expectation of life of 3–7 years. Figures for the billies are unreliable as disappearances of males from the moor are more likely to be the result of emigration than of deaths. While the herds of nannies have fairly restricted home ranges there is some evidence to suggest that many of the billies wander freely along the Borders.
In the early part of the summer the sheep and goats are eating very similar proportions of the plants present on the moor. Later on in the season there is a divergence with the sheep eating less Calluna while the goats eat less Eriophorum , but as the Trichophorum becomes available both animals include this in their diet.     

Environmental Stochasticity and Long-Term Livestock Viability—Herd-Accumulation as a Risk Reducing Strategy

This study tests the hypothesis that herd accumulation can be a risk reducing strategy aimed at increasing security in an unpredictable environment. Saami reindeer husbandry in Norway is characterized by environmental unpredictability and occasionally harsh winters can have dramatic negative effects on reindeer population densities. While herd accumulation has been found to be an adaptive risk reducing strategy in stochastic environments (i.e., individually rational), the accumulation of large herds may also result in collectively negative density dependent effects, which may negatively affect individual herders (i.e., collectively irrational). We found that individual husbandry units’ strategies, such as accumulating reindeer, have a larger effect on individual husbandry units’ herd size than a negative density-dependent effect.     

Home Range and Movement Patterns of White-lipped Peccary (Tayassu pecari) Herds in the Northern Brazilian Amazon1

Two white-lipped peccary (Tayassu pecari) herds were radio-tracked for 5 and 13 mo, respectively, on Maracá Island Ecological Reserve, Roraima, Brazil. Home range size was 109.6 km² for the larger herd (130 animals) and 21.8 km² for the smaller herd (53 animals). Neither herd migrated or moved nomadically during the study period. The herd with the longer observation period increased its home range size during the flooded season by including new sites not used in the nonflooded season and continuing to use the nonflooded season sites. This pattern of simultancously using distinct seasonal ranges also occurred between the rainy and dry periods and the herd regularly and predictably returned to favored feeding sites. Population declines and disappearances in this study and others may have been caused by in situ mortality rather than by long-distance movements.     

Conflicting management objectives on the Colorado Plateau: Understanding the effects of bison and cattle grazing on plant community composition

The American Bison (Bison bison Linnaeus) in the Henry Mountains are one of the last free-roaming, genetically pure herds of bison remaining in North America. Anecdotal evidence indicates that this herd is utilising a cattle winter range during the summer and fall, creating a conflict between the state agency that manages the bison, and the Bureau of Land Management (BLM) and local ranchers. In theory, the addition of bison grazing pressure could reduce forage availability in the short term and lead to undesired changes in the plant community in the long term. Our objective was to determine whether bison have altered the plant species composition of the cattle winter range. We characterised plant species composition, percent cover, and grazing intensity on three adjacent, geomorphologically similar mesas. Grazing regimes were different on the three mesas, one with bison and cattle present, one with cattle only present, and the third with neither cattle nor bison present. Vegetation surveys were accompanied by a 28-year remote sensing time series to test for temporal shifts in an index of primary productivity. We found a higher grazing intensity on two dominant forage species on the bison plus cattle grazed mesa in fall, before the cattle were turned out to winter pasture. Despite this difference in grazing intensity, we found few differences in species composition, percent cover, or NDVI across the three grazing regimes. Our results suggest that high intensity summer bison grazing, while likely creating short-term reductions in forage availability, has not caused differences in plant community composition or productive potential. Shifts in community composition can take years to unfold and just as long to correct; therefore, continued monitoring of the combined effects of cattle and bison is needed.     

Escaping parasitism in the selfish herd: age, size and density-dependent warble fly infestation in reindeer

It has been suggested that animals may escape attack from mobile parasites by aggregating in selfish herds. A selfish herd disperses the risk of being attacked among its members and the per individual risk of parasite infection should therefore decrease with increasing animal density through the encounter–dilution effect. Moreover, in a selfish herd, dominant and agile animals should occupy the best positions and thereby receive fewer attacks compared to lower ranked animals at the periphery. We tested these predictions on reindeer ( Rangifer tarandus tarandus ) parasitized by warble flies ( Hypoderma tarandi ). Warble flies oviposit their eggs on reindeer during summer and induce strong anti-parasitic behavioural responses in the herds. In this period, reindeer are sexually segregated; females and calves form large and dense herds while males are more solitary. After hatching, the warble fly larvae migrate under the skin of their host where they encyst. In the present study encysted larvae were counted on newly slaughtered hides of male calves and 1.5 year old males from 18 different reindeer herds in Finnmark, northern Norway with large contrasts in reindeer density. In reindeer, body mass is correlated with fitness and social status and we hypothesized that individual carcass mass reflected the animal's ability to occupy the best positions within the herd. Larval abundance was higher among the 1.5 year old males than among the calves. For calves we found in accordance with the selfish herd hypothesis a negative relationship between larval abundance and animal density and between larval abundance and body mass. These relationships were absent for the 1.5 year old males. We suggest that these differences were due to different grouping behaviour where calves and females, but not males, aggregated in selfish herds where they escaped parasitism.     

Buffalo Herd Structure and its Repercussions for Condition of Individual African Buffalo Cows

The African buffalo Syncerus caffer was studied in Lake Manyara National Park, Tanzania. Emphasis was placed on the study of (individual) buffalo cows, which live in mixed herds. Buffalo herds are discrete social units and females were never observed in another herd than their own. The herd showed a structure with respect to the distribution of sex-age classes. Individual cows generally kept the same location within the herd. The location in the herd appeared to be coupled to food intake and was strongly related to physical condition. The best location (highest intake and best condition) was between the front and the centre of the herd, the worst location was the rear of the herd (when moving or grazing). Females with calves appeared to have the highest position in the hierarchy as determined from the rate of displacement over food; adult bulls did not interact with cows. Few births were observed during the late dry season and it appears that there is a calving peak at the end of the long rains. Conception rate increased when cows increased in condition and dropped when cows lost condition. Cows showed a strong seasonality in condition but bulls on average hardly changed in condition, except for a loss in condition during the inferred conception peak. Buffalo herds in Manyara showed a fusion-fission pattern independent of season but strongly influenced by the size of the herd: large herds split more often than smaller ones. In large herds, buffalo grazed closer together than in small herds and it appeared likely that competition was more severe in large herds. Animals in the rear of a large herd lost condition faster during the dry season than animals in the best location in the herd, and especially cows in the rear split off most frequently from the herd to graze in a smaller fragment. From the literature on cattle, it is inferred that the reproductive success of cows in the rear of the herd will be lower than of cows in the best location, and this differential is confirmed by the behaviour of adult bulls. It is as yet unclear what the advantage is for adult cows in the rear of a large herd to stay in that herd but the sharing of information with more successful individuals seems a good candidate.