Tick treatment practices in the field: Access to, knowledge about, and on-farm use of acaricides in Laikipia, Kenya.

The prevention of tick-borne diseases is a major challenge for livestock production globally. Tick control strategies include the use of acaricides, but the prescribed strategies do not achieve the desired results in several countries, including Kenya. To better understand how tick treatment practices, contribute to reported tick treatment failures, we assessed livestock owners' acaricide procurement, level of knowledge about acaricides and tick resistance, and how they apply acaricides. We also assessed the quality of the commonly available acaricides. We focused on three livestock systems in Laikipia County, Kenya: two private ranches; one community ranch whose members communally graze their cattle and acquire and apply acaricides; and individual livestock owners in two pastoral communities who individually graze their cattle and acquire and apply acaricides. Through interviews and focus group discussions we assessed; access to acaricides, livestock owners' knowledge, and acaricide use practices; interview data were triangulated with participant observations (n = 107). We analysed nine commonly used acaricides to determine the active ingredient concentration and we determined the concentration of active ingredients in acaricide dilutions collected on farms. All livestock owners had access to and used chemical acaricides for tick control, predominantly amitraz-based. Private ranchers bought one amitraz-based acaricide in bulk directly from the manufacturer, while all other livestock owners bought from agrovet shops. The livestock owners acquired knowledge about acaricides from their own experiences and through experience-based recommendations from peers, but not from the technical information provided by the manufacturers and agrovet shops. All pastoral livestock frequently changed acaricide brand and active ingredient class. A large majority of pastoralists (86%) mixed acaricide brands within and across active ingredient classes; a smaller majority (56%) mixed acaricides with crop pesticides and insecticides. Our lab tests confirmed the content description on the labels bought from agrovet shops. However, on-farm acaricide dilutions from all three livestock systems deviated from the level recommended for effective treatment. If too diluted, the acaricide does not kill ticks, promoting resistance development. If too concentrated, this increases environmental contamination and raises public health concerns. Livestock owners lack a technical understanding of the functioning of acaricides, compromising their use and effectiveness. The widely adopted mixing of acaricides with insecticides and pesticides raises serious health concerns.

Associations between monthly rainfall and mortality in cattle due to East Coast fever, anaplasmosis and babesiosis.

Weather conditions can impact infectious disease transmission, causing mortalities in humans, wild and domestic animals. Although rainfall in dry tropical regions is highly variable over the year, rainfall is thought to play an important role in the transmission of tick-borne diseases. Whether variation in rainfall affects disease-induced mortalities, is, however, poorly understood. Here, we use long-term data on monthly rainfall and Boran cattle mortality (1998-2017) to investigate associations between within-year variation in rainfall and cattle mortalities due to East Coast fever (ECF), anaplasmosis and babesiosis in Laikipia, Kenya, using ARIMAX modelling. Results show a negative correlation between monthly rainfall and cattle mortality for ECF and anaplasmosis, with a lag effect of 2 and 6 months, respectively. There was no association between babesiosis-induced mortalities and monthly rainfall. The results of this study suggest that control of the tick-borne diseases ECF and anaplasmosis to reduce mortalities should be intensified during rainy periods after the respective estimated time lags following dry periods.

Will legal international rhino horn trade save wild rhino populations?

Wild vertebrate populations all over the globe are in decline, with poaching being the second-most-important cause. The high poaching rate of rhinoceros may drive these species into extinction within the coming decades. Some stakeholders argue to lift the ban on international rhino horn trade to potentially benefit rhino conservation, as current interventions appear to be insufficient. We reviewed scientific and grey literature to scrutinize the validity of reasoning behind the potential benefit of legal horn trade for wild rhino populations. We identified four mechanisms through which legal trade would impact wild rhino populations, of which only the increased revenue for rhino farmers could potentially benefit rhino conservation. Conversely, the global demand for rhino horn is likely to increase to a level that cannot be met solely by legal supply. Moreover, corruption is omnipresent in countries along the trade routes, which has the potential to negatively affect rhino conservation. Finally, programmes aimed at reducing rhino horn demand will be counteracted through trade legalization by removing the stigma on consuming rhino horn. Combining these insights and comparing them with criteria for sustainable wildlife farming, we conclude that legalizing rhino horn trade will likely negatively impact the remaining wild rhino populations. To preserve rhino species, we suggest to prioritize reducing corruption within rhino horn trade, increasing the rhino population within well-protected 'safe havens' and implementing educational programmes and law enforcement targeted at rhino horn consumers.

Determining Mhc-DRB profiles in wild populations of three congeneric true lemur species by noninvasive methods.

The major histocompatibility complex (MHC) is a highly polymorphic and polygenic genomic region that plays a crucial role in immune-related diseases. Given the need for comparative studies on the variability of immunologically important genes among wild populations and species, we investigated the allelic variation of MHC class II DRB among three congeneric true lemur species: the red-fronted lemur (Eulemur rufifrons), red-bellied lemur (Eulemur rubriventer), and black lemur (Eulemur macaco). We noninvasively collected hair and faecal samples from these species across different regions in Madagascar. We assessed DRB exon 2 polymorphism with a newly developed primer set, amplifying nearly all non-synonymous codons of the antigen-binding sites. We defined 26 DRB alleles from 45 individuals (17 alleles from E. rufifrons (N = 18); 5 from E. rubriventer (N = 7); and 4 from E. macaco (N = 20). All detected alleles are novel and show high levels of nucleotide (26.8%) and non-synonymous codon polymorphism (39.4%). In these lemur species, we found neither evidence of a duplication of DRB genes nor a sharing of alleles among sympatric groups or allopatric populations of the same species. The non-sharing of alleles may be the result of a geographical separation over a long time span and/or different pathogen selection pressures. We found dN/dS rates > 1 in the functionally important antigen recognition sites, providing evidence for balancing selection. Especially for small and isolated populations, quantifying and monitoring DRB variation are recommended to establish successful conservation plans that mitigate the possible loss of immunogenetic diversity in lemurs.

First records of Hyalomma rufipes and Ixodes neitzi (Acari: Ixodidae) found on large carnivores in South Africa.

Ixodid ticks (Acari: Ixodidae) are important disease vectors for large carnivores, but the composition of the tick communities that parasitize carnivores is poorly understood. We collected ticks from leopards (Panthera pardus) and brown hyenas (Hyaena brunnea) in the Soutpansberg Mountains, South Africa, to determine which species feed on these carnivores. We identified a total of eight tick species belonging to six genera, and recorded Ixodes neitzi and Hyalomma rufipes on P. pardus for the first time.

An experimental approach in revisiting the magnetic orientation of cattle.

In response to the increasing number of observational studies on an apparent south-north orientation in non-homing, non-migrating terrestrial mammals, we experimentally tested the alignment hypothesis using strong neodymium magnets on the resting orientation of individual cattle in Portugal. Contrary to the hypothesis, the 34 cows in the experiment showed no directional preference, neither with, nor without a strong neodymium magnet fixed to their collar. The concurrently performed 2,428 daytime observations-excluding the hottest part of the day-of 659 resting individual cattle did not show a south-north alignment when at rest either. The preferred compass orientation of these cows was on average 130 degrees from the magnetic north (i.e., south east). Cow compass orientation correlated significantly with sun direction, but not with wind direction. In as far as we can determine, this is the first experimental test on magnetic orientation in larger, non-homing, non-migrating mammals. These experimental and observational findings do not support previously published suggestions on the magnetic south-north alignment in these mammals.

Occupancy strongly influences faecal microbial composition of wild lemurs.

The microbiota of the mammalian gut is a complex ecosystem, the composition of which is greatly influenced by host genetics and environmental factors. In this study, we aim to investigate the influence of occupancy (a geographical area of habitation), species, age and sex on intestinal microbiota composition of the three lemur species: Eulemur fulvus, E. rubriventer and E. rufifrons. Faecal samples were collected from a total of 138 wild lemurs across Madagascar, and microbial composition was determined using next-generation sequencing of PCR-amplified 16S rRNA gene fragments. Consistent with reports from other primate species, the predominant phyla were Firmicutes (43 ± 6.4% [s.d.]) and Bacteroidetes (30.3 ± 5.3%). The microbial composition was strongly associated with occupancy in the E. fulvus population, with up to 19.9% of the total variation in microbial composition being explained by this factor. In turn, geographical differences observed in faecal microbiota of sympatric lemur species were less pronounced, as was the impact of the factors sex and age. Our findings showed that among the studied factors occupancy had the strongest influence on intestinal microbiota of congeneric lemur species. This suggests adaptation of microbiota to differences in forest composition, climate variations and correspondingly available diet in different geographical locations of Madagascar.

Correction: Risk Factors for Bovine Tuberculosis (bTB) in Cattle in Ethiopia.

[This corrects the article DOI: 10.1371/journal.pone.0159083.].

Effect of host diversity and species assemblage composition on bovine tuberculosis (bTB) risk in Ethiopian cattle.

Current theories on diversity-disease relationships describe host species diversity and species identity as important factors influencing disease risk, either diluting or amplifying disease prevalence in a community. Whereas the simple term 'diversity' embodies a set of animal community characteristics, it is not clear how different measures of species diversity are correlated with disease risk. We therefore tested the effects of species richness, Pielou's evenness and Shannon's diversity on bovine tuberculosis (bTB) risk in cattle in the Afar Region and Awash National Park between November 2013 and April 2015. We also analysed the identity effect of a particular species and the effect of host habitat use overlap on bTB risk. We used the comparative intradermal tuberculin test to assess the number of bTB-infected cattle. Our results suggested a dilution effect through species evenness. We found that the identity effect of greater kudu - a maintenance host - confounded the dilution effect of species diversity on bTB risk. bTB infection was positively correlated with habitat use overlap between greater kudu and cattle. Different diversity indices have to be considered together for assessing diversity-disease relationships, for understanding the underlying causal mechanisms. We posit that unpacking diversity metrics is also relevant for formulating disease control strategies to manage cattle in ecosystems characterized by seasonally limited resources and intense wildlife-livestock interactions.

Risk Factors for Bovine Tuberculosis (bTB) in Cattle in Ethiopia.

Bovine tuberculosis (bTB) infection is generally correlated with individual cattle's age, sex, body condition, and with husbandry practices such as herd composition, cattle movement, herd size, production system and proximity to wildlife-including bTB maintenance hosts. We tested the correlation between those factors and the prevalence of bTB, which is endemic in Ethiopia's highland cattle, in the Afar Region and Awash National Park between November 2013 and April 2015. A total of 2550 cattle from 102 herds were tested for bTB presence using the comparative intradermal tuberculin test (CITT). Data on herd structure, herd movement, management and production system, livestock transfer, and contact with wildlife were collected using semi-structured interviews with cattle herders and herd owners. The individual overall prevalence of cattle bTB was 5.5%, with a herd prevalence of 46%. Generalized Linear Mixed Models with a random herd-effect were used to analyse risk factors of cattle reactors within each herd. The older the age of the cattle and the lower the body condition the higher the chance of a positive bTB test result, but sex, lactation status and reproductive status were not correlated with bTB status. At herd level, General Linear Models showed that pastoral production systems with transhumant herds had a higher bTB prevalence than sedentary herds. A model averaging analysis identified herd size, contact with wildlife, and the interaction of herd size and contact with wildlife as significant risk factors for bTB prevalence in cattle. A subsequent Structural Equation Model showed that the probability of contact with wildlife was influenced by herd size, through herd movement. Larger herds moved more and grazed in larger areas, hence the probability of grazing in an area with wildlife and contact with either infected cattle or infected wildlife hosts increased, enhancing the chances for bTB infection. Therefore, future bTB control strategies in cattle in pastoral areas should consider herd size and movement as important risk factors.

Scale-dependent bi-trophic interactions in a semi-arid savanna: how herbivores eliminate benefits of nutrient patchiness to plants.

The scale of resource heterogeneity may influence how resources are locally partitioned between co-existing large and small organisms such as trees and grasses in savannas. Scale-related plant responses may, in turn, influence herbivore use of the vegetation. To examine these scale-dependent bi-trophic interactions, we varied fertilizer [(nitrogen (N)/phosphorus (P)/potassium (K)] applications to patches to create different scales of nutrient patchiness (patch size 2 × 2 m, 10 × 10 m, or whole-plot 50 × 50 m) in a large field experiment in intact African savanna. Within-patch fertilizer concentration and the total fertilizer load per plot were independently varied. We found that fertilization increased the leaf N and P concentrations of trees and grasses, resulting in elevated utilization by browsers and grazers. Herbivory off-take was particularly considerable at higher nutrient concentrations. Scale-dependent effects were weak. The net effect of fertilization and herbivory was that plants in fertilized areas tended to grow less and develop smaller rather than larger standing biomass compared to plants growing in areas that remained unfertilized. When all of these effects were considered together at the community (plot) level, herbivory completely eliminated the positive effects of fertilization on the plant community. While this was true for all scales of fertilization, grasses tended to profit more from coarse-grained fertilization and trees from fine-grained fertilization. We conclude that in herbivore-dominated communities, such as the African savanna, nutrient patchiness results in the herbivore community profiting rather more than the plant community, irrespective of the scale of patchiness. At the community level, the allometric scaling theory's prediction of plant-and probably also animal-production does not hold or may even be reversed as a result of complex bi-trophic interactions.

Surviving in a semi-marine habitat: Dietary salt exposure and salt secretion of a New Zealand intertidal skink.

Species that inhabit marine and intertidal ecosystems face osmoregulatory challenges, risking dehydration and increased ion concentrations in the body. Lizards need to either tolerate or regulate these increased ion concentrations. In this study, we aim to understand how Suter's skink (Oligosoma suteri), an intertidal skink restricted to shoreline habitats, is able to cope with the physiological stress of living in an extreme salt environment. We determined the diet, prey species' salt content, and nasal and cloacal salt excretion on Rangitoto and Motutapu Islands in northern New Zealand, where the skinks have contrasting access to terrestrial invertebrates. Analysis of stable isotopes suggests inter- and intra-population variability in Suter's skink diets. Intertidal invertebrates under washed up seaweed appear to compose a major part of the diet of the Rangitoto population, while the Motutapu population showed evidence for a mixed diet of terrestrial and intertidal invertebrates. Sodium content of prey species decreased with an increasing distance from the seawater. Field secretion of cations through nasal glands consisted primarily of Na(+), which is consistent with other marine and intertidal species. Sodium was also the primary cation in urine. In contrast, fecal cations consisted primarily of K(+). This first study into the salt secretion of an intertidal skink species provides evidence of Suter's skink's plasticity in secreting excess ions through nasal salt glands. This likely enables it to deal with the challenges of living in a semi-marine habitat.

Eutrophication of mangroves linked to depletion of foliar and soil base cations.

There is growing concern that increasing eutrophication causes degradation of coastal ecosystems. Studies in terrestrial ecosystems have shown that increasing the concentration of nitrogen in soils contributes to the acidification process, which leads to leaching of base cations. To test the effects of eutrophication on the availability of base cations in mangroves, we compared paired leaf and soil nutrient levels sampled in Nypa fruticans and Rhizophora spp. on a severely disturbed, i.e. nutrient loaded, site (Mahakam delta) with samples from an undisturbed, near-pristine site (Berau delta) in East Kalimantan, Indonesia. The findings indicate that under pristine conditions, the availability of base cations in mangrove soils is determined largely by salinity. Anthropogenic disturbances on the Mahakam site have resulted in eutrophication, which is related to lower levels of foliar and soil base cations. Path analysis suggests that increasing soil nitrogen reduces soil pH, which in turn reduces the levels of foliar and soil base cations in mangroves.

Overlap in nitrogen sources and redistribution of nitrogen between trees and grasses in a semi-arid savanna.

A key question in savanna ecology is how trees and grasses coexist under N limitation. We used N stable isotopes and N content to study N source partitioning across seasons from trees and associated grasses in a semi-arid savanna. We also used (15)N tracer additions to investigate possible redistribution of N by trees to grasses. Foliar stable N isotope ratio (δ(15)N) values were consistent with trees and grasses using mycorrhiza-supplied N in all seasons except in the wet season when they switched to microbially fixed N. The dependence of trees and grasses on mineralized soil N seemed highly unlikely based on seasonal variation in mineralization rates in the Kruger Park region. Remarkably, foliar δ(15)N values were similar for all three tree species differing in the potential for N fixation through nodulation. The tracer experiment showed that N was redistributed by trees to understory grasses in all seasons. Our results suggest that the redistribution of N from trees to grasses and uptake of N was independent of water redistribution. Although there is overlap of N sources between trees and grasses, dependence on biological sources of N coupled with redistribution of subsoil N by trees may contribute to the coexistence of trees and grasses in semi-arid savannas.

Herbaceous forage and selection patterns by ungulates across varying herbivore assemblages in a South African Savanna.

Herbivores generally have strong structural and compositional effects on vegetation, which in turn determines the plant forage species available. We investigated how selected large mammalian herbivore assemblages use and alter herbaceous vegetation structure and composition in a southern African savanna in and adjacent to the Kruger National Park, South Africa. We compared mixed and mono-specific herbivore assemblages of varying density and investigated similarities in vegetation patterns under wildlife and livestock herbivory. Grass species composition differed significantly, standing biomass and grass height were almost twice as high at sites of low density compared to high density mixed wildlife species. Selection of various grass species by herbivores was positively correlated with greenness, nutrient content and palatability. Nutrient-rich Urochloa mosambicensis Hack. and Panicum maximum Jacq. grasses were preferred forage species, which significantly differed in abundance across sites of varying grazing pressure. Green grasses growing beneath trees were grazed more frequently than dry grasses growing in the open. Our results indicate that grazing herbivores appear to base their grass species preferences on nutrient content cues and that a characteristic grass species abundance and herb layer structure can be matched with mammalian herbivory types.

Shrimp pond effluent dominates foliar nitrogen in disturbed mangroves as mapped using hyperspectral imagery.

Conversion of mangroves to shrimp ponds creates fragmentation and eutrophication. Detection of the spatial variation of foliar nitrogen is essential for understanding the effect of eutrophication on mangroves. We aim (i) to estimate nitrogen variability across mangrove landscapes of the Mahakam delta using airborne hyperspectral remote sensing (HyMap) and (ii) to investigate links between the variation of foliar nitrogen mapped and local environmental variables. In this study, multivariate prediction models achieved a higher level of accuracy than narrow-band vegetation indices, making multivariate modeling the best choice for mapping. The variation of foliar nitrogen concentration in mangroves was significantly influenced by the local environment: (1) position of mangroves (seaward/landward), (2) distance to the shrimp ponds, and (3) predominant mangrove species. The findings suggest that anthropogenic disturbances, in this case shrimp ponds, influence nitrogen variation in mangroves. Mangroves closer to the shrimp ponds had higher foliar nitrogen concentrations.

Optimization of wildlife management in a large game reserve through waterpoints manipulation: a bio-economic analysis.

Surface water is one of the constraining resources for herbivore populations in semi-arid regions. Artificial waterpoints are constructed by wildlife managers to supplement natural water supplies, to support herbivore populations. The aim of this paper is to analyse how a landowner may realize his ecological and economic goals by manipulating waterpoints for the management of an elephant population, a water-dependent species in the presence of water-independent species. We develop a theoretical bio-economic framework to analyse the optimization of wildlife management objectives (in this case revenue generation from both consumptive and non-consumptive use and biodiversity conservation), using waterpoint construction as a control variable. The model provides a bio-economic framework for analysing optimization problems where a control has direct effects on one herbivore species but indirect effects on the other. A landowner may be interested only in maximization of profits either from elephant offtake and/or tourism revenue, ignoring the negative effects that could be brought about by elephants to biodiversity. If the landowner does not take the indirect effects of waterpoints into consideration, then the game reserve management, as the authority entrusted with the sustainable management of the game reserve, might use economic instruments such as subsidies or taxes to the landowners to enforce sound waterpoint management.

Large herbivores may alter vegetation structure of semi-arid savannas through soil nutrient mediation.

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.

The spatial scaling of habitat selection by African elephants.

1. Understanding and accurately predicting the spatial patterns of habitat use by organisms is important for ecological research, biodiversity conservation and ecosystem management. However, this understanding is complicated by the effects of spatial scale, because the scale of analysis affects the quantification of species-environment relationships. 2. We therefore assessed the influence of environmental context (i.e. the characteristics of the landscape surrounding a site), varied over a large range of scales (i.e. ambit radii around focal sites), on the analysis and prediction of habitat selection by African elephants in Kruger National Park, South Africa. 3. We focused on the spatial scaling of the elephants' response to their main resources, forage and water, and found that the quantification of habitat selection strongly depended on the scales at which environmental context was considered. Moreover, the inclusion of environmental context at characteristic scales (i.e. those at which habitat selectivity was maximized) increased the predictive capacity of habitat suitability models. 4. The elephants responded to their environment in a scale-dependent and perhaps hierarchical manner, with forage characteristics driving habitat selection at coarse spatial scales, and surface water at fine spatial scales. 5. Furthermore, the elephants exhibited sexual habitat segregation, mainly in relation to vegetation characteristics. Male elephants preferred areas with high tree cover and low herbaceous biomass, whereas this pattern was reversed for female elephants. 6. We show that the spatial distribution of elephants can be better understood and predicted when scale-dependent species-environment relationships are explicitly considered. This demonstrates the importance of considering the influence of spatial scale on the analysis of spatial patterning in ecological phenomena.