Tall Grass Invasion After Grassland Abandonment Influences the Availability of Palatable Plants for Wild Herbivores: Insight into the Conservation of the Apennine Chamois Rupicapra pyrenaica ornata.

Invasion of the tall grass Brachypodium genuense was observed in an area of the central Apennines (Italy) where the population size of Apennine chamois (Rupicapra pyrenaica ornata) was in strong decline. Since this dominant tall grass threatens biodiversity and forage quality, our hypothesis was that B. genuense abundance influenced that of palatable species for the chamois, depending on their functional traits and distribution patterns. Our sampling design used plots of 10 × 10 m and 1 × 1 m to investigate the plant community level and fine-scale interactions. We analyzed data using correlation, generalized linear models, and redundancy analyses. We found that B. genuense can reach high abundance values on the deepest soils. Its high cover value influences plant community composition by competitive exclusion of subordinate species and suppression of functional features because of temporal or spatial niche overlap. This leads to low cover of palatable species at a fine scale, and to poor pasture quality for chamois at a wider scale. Therefore, we postulated that B. genuense invasion, enhanced by long-term grazing cessation, may reduce the availability of palatable plants for Apennine chamois, especially below the potential timberline (1900-2000 m a.s.l.). The high abundance of B. genuense may amplify the effect of other negative factors, such as competition with red deer (Cervus elaphus) and climate change, in restricting the suitable habitat of the Apennine chamois to the higher sectors of the central Apennines. Thus, we suggested that B. genuense spread should be monitored carefully and plans to control its invasion should be implemented.

Horn growth patterns in Alpine chamois.

The analysis of horn growth may provide important information about the allocation of metabolic resources to secondary sexual traits. Depending on the selective advantages offered by horn size during intra- and inter-specific interactions, ungulates may show different investment in horn development, and growth variations within species may be influenced by several parameters, such as sex, age, or resource availability. We investigated the horn growth patterns in two hunted populations of Alpine chamois (Rupicapra r. rupicapra) in the Central Italian Alps. We tested the role of individual heterogeneity on the growth pattern and explored the variation in annulus length as a function of different factors (sex, age, hunting location, cohort). We then investigated the mechanisms underlying horn growth trajectories to test for the occurrence of compensatory or recovery growth and their potential differences between sexes and populations. Annulus length varied as a function of sex, age of individuals and, marginally, hunting location; no effect of cohort or individual heterogeneity was detected. Male and female chamois showed compensatory horn growth within the first 5½ years of life, though the partial convergence of horn trajectories in chamois suggests that this mechanisms would best be described as 'recovery growth'. Compensation rates were greater in males than in females, while only compensatory growth rates up to 2½ years of age were different in the two populations. Besides confirming the sex- and age-dependent pattern of horn development, our study suggests that the mechanism of recovery growth supports the hypothesis of horn size as a weakly selected sexual trait in male and female chamois. Furthermore, the greater compensation rates in horn growth shown by male chamois possibly suggest selective effects of hunting on age at first reproduction, while different compensation rates between populations may suggest the occurrence of some plasticity in resource allocation to sexual traits in relation to different environments.

Contrasting life histories in neighbouring populations of a large mammal.

BACKGROUND: A fundamental life history question is how individuals should allocate resources to reproduction optimally over time (reproductive allocation). The reproductive restraint hypothesis predicts that reproductive effort (RE; the allocation of resources to current reproduction) should peak at prime-age, whilst the terminal investment hypothesis predicts that individuals should continue to invest more resources in reproduction throughout life, owing to an ever-decreasing residual reproductive value. There is evidence supporting both hypotheses in the scientific literature. METHODOLOGY/PRINCIPAL FINDINGS: We used an uncommonly large, 38 year dataset on Alpine chamois (Rupicapra rupicapra) shot at various times during the rutting period to test these two hypotheses. We assumed that body mass loss in rutting males was strongly related to RE and, using a process-based approach, modelled how male relative mass loss rates varied with age. For different regions of our study area, we provide evidence consistent with different hypotheses for reproductive allocation. In sites where RE declined in older age, this appears to be strongly linked to declining body condition in old males. In this species, terminal investment may only occur in areas with lower rates of body mass senescence. CONCLUSIONS/SIGNIFICANCE: Our results show that patterns of reproductive allocation may be more plastic than previously thought. It appears that there is a continuum from downturns in RE at old age to terminal investment that can be manifest, even across adjacent populations. Our work identifies uncertainty in the relationship between reproductive restraint and a lack of competitive ability in older life (driven by body mass senescence); both could explain a decline in RE in old age and may be hard to disentangle in empirical data. We discuss a number of environmental and anthropogenic factors which could influence reproductive life histories, underlining that life history patterns should not be generalised across different populations.

Effects of early horn growth on reproduction and hunting mortality in female chamois.

1. Environmental conditions during early development can affect the growth patterns of vertebrates, influencing future survival and reproduction. In long-lived mammals, females that experience poor environmental conditions early in life may delay primiparity. In female bovids, annual horn growth increments may provide a record of age-specific reproduction and body growth. Horn length, however, may also be a criterion used by hunters in selecting animals to harvest, possibly leading to artificial selection. 2. We studied three populations of chamois (Rupicapra rupicapra) in the western Alps to explore the relationships between female horn length and early growth, age of primiparity and age-specific reproduction. We also compared the risk of harvest to reproductive status and horn length. 3. Early horn growth was positively correlated with body mass in pre-reproductive females and with reproduction in very young and senescent adults. Females with strong early horn growth attained primiparity at an earlier age than those with weak early growth. Horn length did not affect hunter selection, but we found a strong hunter preference for nonlactating females. 4. Our research highlights the persistent effects of early development on reproductive performance in mammals. Moderate sport harvests are unlikely to affect the evolution of phenotypic traits and reproductive strategies in female chamois. A policy of penalizing hunters that harvest lactating females, however, may increase the harvest of 2-year-old females, which have high reproductive potential.

Border disease virus shedding and detection in naturally infected Pyrenean chamois (Rupicapra pyrenaica).

Pyrenean chamois (Rupicapra pyrenaica) populations of the central and eastern Pyrenees have been affected by severe outbreaks associated with Border disease virus (BDV) since 2001. Eight Pyrenean chamois (7 males and 1 female) from 1 to 8 years of age with clinical signs consistent with BDV infection were studied. At necropsy, whole blood, tissue samples (skin, brain, prescapular lymph node, thyroid gland, lung, liver, spleen, kidney, small intestine, bone marrow, and testicle), urine, and nasal, oral, and rectal swabs were obtained. The fetus from a pregnant female was also studied. Reverse transcription polymerase chain reaction (RT-PCR) was used to detect the virus in all samples, and virus isolation was performed. Sera and tissue samples were positive to RT-PCR, and the virus was isolated from all chamois. The nasal, oral, and rectal swabs and urine samples were RT-PCR positive in 100%, 85.71%, 71.43%, and 100% of chamois, respectively, confirming the excretion of the virus via these 4 routes. In addition, sera were tested for BDV antibodies using enzyme-linked immunosorbent assay and seroneutralization techniques, with negative results. Sequence analysis of the 5' untranslated region in 7 of the chamois confirmed that the virus is grouped into the BDV-4 genotype, the same BDV previously described in Pyrenean chamois. To the authors' knowledge, this is the first study of naturally infected Pyrenean chamois, providing evidence that infected animals shed BDV through nasal, oral, fecal, and urinary excretion routes.

The factorial decomposition of the Mahalanobis distances in habitat selection studies.

The Mahalanobis distances have been introduced in habitat selection studies for the estimation of environmental suitability maps (ESMs). The pixels of raster maps of a given area correspond to points in the multidimensional space defined by the mapped environmental variables (ecological space). The Mahalanobis distances measure the distances in this space between these points and the mean of the ecological niche (i.e., the hypothesized optimum for the species) regarding the structure of the niche. The map of these distances over the area of interest is an estimated ESM. Several authors recently noted that the use of a single optimum for the niche of a species may lead to biased predictions of animal occurrence. They proposed to use instead a minimum set of basic habitat requirements, found by partitioning the Mahalanobis distances into a restricted set of biologically meaningful axes. However, the statistical approach they proposed does not take into account the environmental conditions on the area where the niche was sampled (i.e., the environmental availability), and we show that including this availability is necessary. We used their approach as a basis to develop a new exploratory tool, the Mahalanobis distance factor analysis (MADIFA), which performs an additive partitioning of the Mahalanobis distances taking into account this availability. The basic habitat requirements of a species can be derived from the axes of the MADIFA. This method can also be used to compute ESMs using only this small number of basic requirements, therefore including only the biologically relevant information. We also prove that the MADIFA is complementary to the commonly used ecological-niche factor analysis (ENFA). We used the MADIFA method to analyze the niche of the chamois Rupicapra rupicapra in a mountainous area. This method adds to the existing set of tools for the description of the niche.