Disease transmission in animal transfer networks.

Infectious diseases transmission is strongly determined by who contacts whom. Bovine tuberculosis (bTB) caused by Mycobacterium bovis is a worldwide burden for animal populations. One of the major transmission mechanism between herd is the transfer of infectious animal. In East Africa, pastoralists may receive or bestow livestock to create and strengthen social relationships. Here, we used a network approach to examine the relative importance of such cattle transfer in the transmission of bTB. First, a total of 2550 cattle from 102 herds were tested using the comparative intradermal tuberculin test to assess the presence of bTB infected cattle in the herd. A herd was considered bTB positive if it had at least one tuberculin reactor animal. Next, we calculated the centrality of each herd in the cattle transfer network using four established measures of social network centralization: degree, betweenness, closeness and fragmentation. The relationships between the network centrality measures and bTB infection were examined using generalized linear mixed models (GLMM). We found that a herd's in-degree in the social network was positively correlated with the risk of being infected with bTB (b=4.2, 95%CI=2.1-5.7; p<0.001). A herd that was close to many others (i.e., had a higher closeness index) had a larger chance of acquiring bTB infection (b=2.1, 95%CI=1.4-2.8; p<0.001). Betweenness centrality was also positively associated with the presence of bTB infection. There was a negative relationship between the fragmentation index and bTB infection (b=-2.7, 95%CI=-4.9-1.3; p<0.001). The study clearly demonstrated that the extent to which a herd is connected within a network has significant implications for its probability of being infected. Further, the results are in accordance with our expectation that connectivity and the probability that a herd will transmit the disease to other herds in the network are related.

The diversity-disease relationship: evidence for and criticisms of the dilution effect.

The dilution effect, that high host species diversity can reduce disease risk, has attracted much attention in the context of global biodiversity decline and increasing disease emergence. Recent studies have criticized the generality of the dilution effect and argued that it only occurs under certain circumstances. Nevertheless, evidence for the existence of a dilution effect was reported in about 80% of the studies that addressed the diversity-disease relationship, and a recent meta-analysis found that the dilution effect is widespread. We here review supporting and critical studies, point out the causes underlying the current disputes. The dilution is expected to be strong when the competent host species tend to remain when species diversity declines, characterized as a negative relationship between species' reservoir competence and local extinction risk. We here conclude that most studies support a negative competence-extinction relationship. We then synthesize the current knowledge on how the diversity-disease relationship can be modified by particular species in community, by the scales of analyses, and by the disease risk measures. We also highlight the complex role of habitat fragmentation in the diversity-disease relationship from epidemiological, evolutionary and ecological perspectives, and construct a synthetic framework integrating these three perspectives. We suggest that future studies should test the diversity-disease relationship across different scales and consider the multiple effects of landscape fragmentation.

Competition between a Lawn-Forming Cynodon dactylon and a Tufted Grass Species Hyparrhenia hirta on a South-African Dystrophic Savanna.

South African savanna grasslands are often characterised by indigestible tufted grass species whereas lawn grasses are far more desirable in terms of herbivore sustenance. We aimed to investigate the role of nutrients and/or the disturbance (grazing, trampling) by herbivores on the formation of grazing lawns. We conducted a series of common garden experiments to test the effect of nutrients on interspecific competition between a typical lawn-forming grass species (Cynodon dactylon) and a species that is frequently found outside grazing lawns (Hyparrhenia hirta), and tested for the effect of herbivore disturbance in the form of trampling and clipping. We also performed a vegetation and herbivore survey to apply experimentally derived insights to field observations. Our results showed that interspecific competition was not affected by soil nutrient concentrations. C. dactylon did show much more resilience to disturbance than H. hirta, presumably due to the regenerative capacity of its rhizomes. Results from the field survey were in line with these findings, describing a correlation between herbivore pressure and C. dactylon abundance. We conclude that herbivore disturbance, and not soil nutrients, provide C. dactylon with a competitive advantage over H. hirta, due to vegetative regeneration from its rhizomes. This provides evidence for the importance of concentrated, high herbivore densities for the creation and maintenance of grazing lawns.

Dilution effect and identity effect by wildlife in the persistence and recurrence of bovine tuberculosis.

Current theories on disease-diversity relationships predict a strong influence of host richness on disease transmission. In addition, identity effect, caused by the occurrence of particular species, can also modify disease risk. We tested the richness effect and the identity effects of mammal species on bovine tuberculosis (bTB), based on the regional bTB outbreak data in cattle from 2005-2010 in Africa. Besides, we also tested which other factors were associated with the regional bTB persistence and recurrence in cattle. Our results suggested a dilution effect, where higher mammal species richness (MSR) was associated with reduced probabilities of bTB persistence and recurrence in interaction with cattle density. African buffalo had a positive effect on bTB recurrence and a positive interaction effect with cattle density on bTB persistence, indicating an additive positive identity effect of buffalo. The presence of greater kudu had no effect on bTB recurrence or bTB persistence. Climatic variables only act as risk factors for bTB persistence. In summary, our study identified both a dilution effect and identity effect of wildlife and showed that bTB persistence and recurrence were correlated with different sets of risk factors. These results are relevant for more effective control strategies and better targeted surveillance measures in bTB.

Hibernation site requirements of bats in man-made hibernacula in a spatial context.

Bat hibernacula selection depends on various spatial and nonspatial variables that differ widely between sites. However, previous studies have focused mainly on nonspatial variables. This research investigated factors that determined the abundance and species richness of hibernating bats in hibernation objects of the New Dutch Waterline, The Netherlands, and determined the relevant scales over which spatial factors operate using regression techniques and ecological-niche factor analyses. The effects of 32 predictor variables on several response variables, i.e., the total bat abundance, species richness, and abundance and presence of bat species, were investigated. Predictor variables were classified as internal variables (e.g., building size, climatic conditions, and human access) or external variables (e.g., ground and vegetation cover and land cover type) that were measured at different spatial scales to study the influence of the spatial context. The internal building variables (mainly the size of hibernacula and the number of hiding possibilities) affected the hibernating bat abundance and species richness. Climatic variables, such as changes in temperature and humidity, were less important. The hibernation site suitability was also influenced by spatial variables at a variety of scales, thereby indicating the importance of scale-dependent species-environment relationships. The absence of human use and public access enhanced hibernation site suitability, but the internal size-related variables had the greatest positive effect on hibernation site suitability. These results demonstrate the importance of considering the different spatial scales of the surrounding landscape to better understand habitat selection, and they offer directives to managers to optimize objects for hibernating bats and to improve management and bat conservation. The analyses have wider applications to other wildlife-habitat studies.

Spatial autocorrelation and the scaling of species-environment relationships.

Issues of residual spatial autocorrelation (RSA) and spatial scale are critical to the study of species-environment relationships, because RSA invalidates many statistical procedures, while the scale of analysis affects the quantification of these relationships. Although these issues independently are widely covered in the literature, only sparse attention is given to their integration. This paper focuses on the interplay between RSA and the spatial scaling of species-environment relationships. Using a hypothetical species in an artificial landscape, we show that a mismatch between the scale of analysis and the scale of a species' response to its environment leads to a decrease in the portion of variation explained by environmental predictors. Moreover, it results in RSA and biased regression coefficients. This bias stems from error-predictor dependencies due to the scale mismatch, the magnitude of which depends on the interaction between the scale of landscape heterogeneity and the scale of a species' response to this heterogeneity. We show that explicitly considering scale effects on RSA can reveal the characteristic scale of a species' response to its environment. This is important, because the estimation of species-environment relationships using spatial regression methods proves to be erroneous in case of a scale mismatch, leading to spurious conclusions when scaling issues are not explicitly considered. The findings presented here highlight the importance of examining the appropriateness of the spatial scales used in analyses, since scale mismatches affect the rigor of statistical analyses and thereby the ability to understand the processes underlying spatial patterning in ecological phenomena.

Tree cover and biomass increase in a southern African savanna despite growing elephant population.

The growing elephant populations in many parts of southern Africa raise concerns of a detrimental loss of trees, resulting in overall reduction of biodiversity and ecosystem functioning. Elephant distribution and density can be steered through artificial waterpoints (AWPs). However, this leaves resident vegetation no relief during dry seasons. We studied how the introduction of eight AWPs in 1996 affected the spatiotemporal tree-structure dynamics in central Chobe National Park, an unfenced savanna area in northern Botswana with a dry-season elephant density of approximately 3.34 individuals per square kilometer. We hypothesized that the impact of these AWPs amplified over time and expanded in space, resulting in a decrease in average tree density, tree height, and canopy volume. We measured height and canopy dimensions of all woody plants around eight artificial and two seasonal waterpoints for 172 plots in 1997, 2000, and 2008. Plots, consisting of 50 x 2 m transects for small trees (0.20-3.00 m tall) nested within 50 x 20 m transects for large trees (> or = 3.0 m tall), were located at 100, 500, 1000, 2000, and 5000 m distance classes. A repeated-measures mixed-effect model showed that tree density, cover, and volume had increased over time throughout the area, caused by a combination of an increase of trees in lower size classes and a decrease in larger size classes. Our results indicate that the decrease of large trees can be attributed to a growing elephant population. Decrease or loss of particular tree size classes may have been caused by a loss of browser-preferred species while facilitating the competitiveness of less-preferred species. In spite of 12 years of artificial water supply and an annual elephant population growth of 6%, we found no evidence that the eight AWPs had a negative effect on tree biomass or tree structure. The decreasing large-tree component could be a remainder of a depleted but currently restoring elephant population.

Large herbivores that strive mightily but eat and drink as friends.

Grazing in patches of Cynodon dactylon and of Sporobolus spicatus by four large herbivores, and the interaction between these sedentary herbivores was studied in Lake Manyara National Park, northern Tanzania. The herbivores were the African buffalo, Syncerus caffer; the African elephan, Loxodonta africana; the Burchell's zebra, Equus burchelli; and the wildebeest, Connochaetus taurinus. Four different hypotheses of the interactions between the herbivores were tested, viz., increased predator detection/protection through association of species, facilitation of the food intake through the influence of other species, use by other species of the food manipulation strategy of buffalo, and interspecific competition for food. On the level of a single day, zebra and wildebeest were symbiotic, which could have been caused by an increased chance of predator detection. A similar association between buffalo and wildebeest or zebra was also detected on C. dactylon grasslands. There was no indication of facilitation between any of the herbivores. Buffalo had a despotic relationship with elephant, that is the elephant's consumption was lowered when buffalo had visited a patch prior to their arrival. When elephant and buffalo arrived at the same time there appeared to be scramble competition between them.Habitat overlap was calculated for four pairs of species. In conjunction with the analyses of the patch visits, it was concluded that a small overlap was associated with interspecific competition and a large habitat overlap was associated with symbiosis.