Monitoring and management of the endangered Cape mountain zebra Equus zebra zebra in the Western Cape, South Africa

De Hoop Nature Reserve and a neighbouring conservancy contain the most genetically diverse subpopulation of the Endangered (IUCN) Cape mountain zebra ( Equus zebra zebra Linnaeus 1758). Although vital for the long-term stability of the meta-population, the population had received limited monitoring post-1999. We summarize data obtained during a population monitoring programme established in 2005. Ninety-nine individuals were identified indicating a decline in annual population growth from 6.6% (1995–1999) to 4.5% (1999–2005). The population was male biased and the deficit of females is likely to have prevented additional breeding herd formation resulting in excess nonbreeding males. These animals are currently of limited reproductive value to the meta-population and may be contributing to the decline in reproductive potential at De Hoop by competing for limited resources. One solution may be to translocate 'excess' males to reinforce existing small populations or establish new populations with females from elsewhere provided that a minimum of 78 animals is maintained at De Hoop to limit genetic loss. Population monitoring and effective management strategies for the De Hoop population and the meta-population are vital to ensure the long-term survival of Cape mountain zebra and for the success of other species recovery programmes.     

Genetic markers and sample types for pedigree reconstruction in Grevy's zebra (Equus grevyi)

Effective captive population management relies on maintaining accurate pedigree records in order to direct breeding programmes and maximise conservation of genetic diversity. Where familial relationships or individual identities are unknown, it is now possible to recover this information via genetic analysis. We describe the first application of DNA profiling to reconstruct family groups of the Grevy's zebra (Equus grevyi), in order to identify three unknown stallions. To examine the potential for using non-invasive sampling techniques, DNA recovered from faeces, hair and blood was compared. Microsatellite loci derived from horses were tested for their utility in Grevy's zebra and a set of loci were subsequently employed to successfully identify the three animals via parental exclusion. The study demonstrates the relative ease with which DNA profiling can now be applied to captive management and highlights methods of genetic sampling that minimise stress to the animal and reduce sampling costs.     

Cape mountain zebra in the Baviaanskloof Nature Reserve,South Africa: resource use reveals limitations to zebra performance in a dystrophic mountainous ecosystem

Resource use of Cape mountain zebra was studied in the Baviaanskloof Nature Reserve (BNR) over concerns of the poor performance in population growth. We assessed the seasonal diet, habitat suitability and forage quality of the Bergplaas area in BNR for mountain zebra. Grasses contributed 95.2% to the annual diet of mountain zebra, Tristachya leucothrix contributing the most (39.4%), followed by Themeda triandra (27.6%). Seasonally, T. triandra contributed most to the diet in winter, while T. leucothrix became more important in the summer and also was the only species preferred in all seasons. Mountain zebra concentrated their feeding in Kouga Grassy Fynbos and from our assessment this was the only habitat suitable for mountain zebra. Our analysis of mountain zebra dung indicated that the seasonal nitrogen and phosphorus content was below the threshold values prescribed for grazers, and our study suggests that mountain zebra at Bergplaas are severely resource limited. We emphasize the importance of fire and access to nutrient‐rich lowlands in influencing the nutritional ecology of mountain zebra and provide conservation management recommendations.     

Urinary steroid evaluations to monitor ovarian function in exotic ungulates: VI. Pregnancy detection in exotic equidae

The measurement of serum and urinary estrone sulfate (ES) was evaluated on random single samples for the detection of pregnancy in the Przewalski's horse (PRZ) and Hartmann's mountain zebra (HMZ). Pregnancy was detected by ES analysis of urine or serum in the PRZ and HMZ by at least 8 and 10 months before delivery, respectively. The values of ES in both serum and urine are shown at various stages of gestation. Results indicate urinary ES analysis is a noninvasive early diagnosis for pregnancy in these species.     

Palaeozoological insights into management options for a threatened mammal: southern Africa’s Cape mountain zebra (Equus zebra zebra)

Aim Promoting population growth of genetically distinct subpopulations of Cape mountain zebra (Equus zebra zebra) is crucial to the survival of the subspecies. Several important Cape mountain zebra reserves are dominated by fynbos vegetation, and population growth is limited by a lack of grassland habitat. A fossil ungulate sequence spanning the last c. 18,000 years is examined to understand the long‐term history of this conservation challenge. Location Boomplaas Cave (BPA), South Africa. Methods The fossil sequence from BPA is examined to reconstruct ungulate community dynamics in relation to climate and vegetation change over the last 18,000 years. Results Ungulates from 18,000 to 12,000 years ago suggest an expansion of open grasslands that supported a grazing ecosystem dominated by an extinct caprine antelope and equid remains attributed to E. zebra and E. quagga. At the onset of the Holocene, the grazing ungulate community disappears and small browsers and mixed feeders dominate the assemblage, indicating the loss of open grassland vegetation. Several open‐habitat grazers go extinct at this time, and Equus persists at much lower abundances. This shift can be explained by global climate change across the Pleistocene–Holocene transition. Main conclusions The fossil sequence supports contemporary observations indicating that access to open grassland is crucial to maintaining large Cape mountain zebra subpopulations. Although fynbos is abundant throughout the historic range of the Cape mountain zebra, fossil evidence suggests that such vegetation is unlikely to support dense populations. It has been suggested that the acquisition of agricultural lands that were historically converted to open grasslands for livestock could promote Cape mountain zebra population growth. Results presented here support this management option, as the open grasslands in these converted landscapes likely approximate the vegetation structure during latest Pleistocene, when grasslands were widespread and grazing ungulates abundant.     

Rolling pits of Hartmann’s mountain zebra (Zebra equus hartmannae) increase vegetation diversity and landscape heterogeneity in the Pre‐Namib

Microsites created by soil‐disturbing animals are important landscape elements in arid environments. In the Pre‐Namib, dust‐bathing behavior of the near‐endemic Hartmann''s mountain zebra creates unique rolling pits that persist in the landscape. However, the ecohydrological characteristics and the effects of those microsites on the vegetation and on organisms of higher trophic levels are still unknown. In our study, we characterized the soil grain size composition and infiltration properties of rolling pits and reference sites and recorded vegetation and arthropod assemblages during the rainy season of five consecutive years with different amounts of seasonal rainfall. We further used the excess green vegetation index derived from drone imagery to demonstrate the different green up and wilting of pits and references after a rainfall event. In contrast to the surrounding grassland, rolling pits had finer soil with higher nutrient content, collected runoff, showed a higher infiltration, and kept soil moisture longer. Vegetation in the rolling pits was denser, dominated by annual forbs and remained green for longer periods. The denser vegetation resulted in a slightly higher activity density of herbivorous arthropods, which in turn increased the activity density of omnivorous and predatory arthropods. In times of drought, the rolling pits could act as safe sites and refuges for forbs and arthropods. With their rolling pits, Hartmann''s mountain zebras act as ecosystem engineers, contributing to the diversity of forb communities and heterogeneity of the landscape in the Pre‐Namib.     

Population structuring in mountain zebras (<Emphasis Type="Italic">Equus zebra</Emphasis>): The molecular consequences of divergent demographic histories

The endangered mountain zebra (Equus zebra) is endemic to the semi-arid inhospitable mountainous escarpments of southern Africa. The species is divided taxonomically into two geographically separated subspecies, each with differing recent population histories. In Namibia, Hartmann’s mountain zebra (E. z. hartmannae) is common and occurs in large free-ranging populations, whereas in South Africa, prolonged hunting and habitat destruction over the last 300 years has decimated populations of the Cape mountain zebra (E. z. zebra). In this study, we investigate the consequences of these divergent demographic histories for population genetic diversity and structure. We also examine the phylogeographic relationship between the two taxonomic groups. Genetic information was obtained at 15 microsatellite loci for 291 individuals from a total of 10 populations as well as 445 bp of the mitochondrial control region sequence data from 77 individuals. Both model-based and standard analytical approaches were used to examine the data. Both types of marker returned levels of diversity and structure that were consistent with population history. Low genetic variation within individual Cape mountain zebra populations, the characteristic indicator of population fragmentation and drift, was offset by moderate variation in the entire E. z. zebra sample. This implies that higher levels of diversity still exist within the Cape mountain zebra gene pool. A management strategy that entailed the mixing of aboriginal populations is therefore advocated in order to halt the further loss of Cape mountain zebra genetic diversity. Allele frequencies in Hartmann’s mountain zebra were relatively resilient to demographic fluctuations. Due to the high incidence of mitochondrial haplotype sharing between populations, the hypothesis that Cape and Hartmann’s mountain zebra mitochondrial lineages were reciprocally monophyletic was not supported. However, the presence of private alleles at nuclear loci rendered the two subspecies genetically distinct evolutionary significant units.     

The forgotten Grevy's zebra Equus grevyi population along the Kasigau Corridor ranches,SE Kenya: recent records and conservation issues

Since the 1970s, the globally endangered Grevy's zebra Equus grevyi has suffered a substantial reduction in range and population size. Grevy's zebra was introduced into the Tsavo ecosystem in two translocations: 22 individuals released in Tsavo East National Park in 1964 and 30 individuals released in Tsavo West National Park in 1977. This study focuses on the Tsavo East subpopulation within the Kasigau Corridor REDD+ Project Area. Data were collected from 2011 to 2014 using systematic and ad lib methods, while vegetation stratification was based on tonnes of CO₂e per hectare. Cumulatively, 785 Grevy's zebra were seen in 210 encounters. The largest single group comprised of 24 individuals; mean group size was 4 ± 3. Foals comprised 10% of all individuals seen and were recorded across most months, suggesting year‐round breeding. The range core for the Grevy's zebra in this area was estimated to be 400 km². Its density was estimated at 0.09 ± 0.038 individuals per km², translating to a population of 36 ± 15 individuals (range: 6–67) based on the estimated core range. Grasslands and sparse Acacia–Commiphora vegetation strata were used significantly more than expected. Despite constituting only about 2–3% of the global Grevy's zebra population, this subpopulation could still play a role as a – disconnected – reserve population.     

Environmental transmission of a personality trait: foster parent exploration behaviour predicts offspring exploration behaviour in zebra finches

Consistent behavioural differences among individuals are common in many species and can have important effects on offspring fitness. To understand such ‘personality’ variation, it is important to determine the mode of inheritance, but this has been quantified for only a few species. Here, we report results from a breeding experiment in captive zebra finches, Taeniopygia guttata, in which we cross-fostered offspring to disentangle the importance of genetic and non-genetic transmission of behaviour. Genetic and foster-parents’ exploratory type was measured in a novel environment pre-breeding and offspring exploratory type was assessed at adulthood. Offspring exploratory type was predicted by the exploratory behaviour of the foster but not the genetic parents, whereas offspring size was predicted by genetic but not foster-parents’ size. Other aspects of the social environment, such as rearing regime (uni- versus biparental), hatching position, brood size or an individual''s sex did not influence offspring exploration. Our results therefore indicate that non-genetic transmission of behaviour can play an important role in shaping animal personality variation.     

Development of polymorphic microsatellite markers for the zebra finch (Taeniopygia guttata)

To study the population genetics as well as the mating system of captive zebra finch (Taeniopygia guttata) populations, we developed primers for 12 microsatellite loci and screened them in 529 individuals from two successive generations of a single captive population. All markers were polymorphic with five to 14 alleles per locus. We checked all markers for Mendelian inheritance in 307 offspring whose parents were known for sure. Four markers showed evidence for the presence of null alleles. Once allowing for null alleles, we found no mismatches between offspring and parents, suggesting a very low rate of mutation. Average observed and expected heterozygosities across the eight loci showing no evidence for null‐alleles was 0.819 and 0.812, respectively.     

Using sample aerial surveys to estimate the abundance of the endangered Grevy’s zebra in northern Kenya

The effective management of endangered mammals requires reliable estimates of population size. This is challenging for species such as Grevy’s zebra (Equus grevyi) that are distributed over large areas at low densities. Less than 2500 Grevy’s zebra remain in the wild, scattered across 85,000 km² of savannah in northern Kenya and Ethiopia. An efficient, accurate and repeatable survey method is required to guide conservation planning for the species. Currently, total aerial counts are used to census endangered species within Kenya, but are costly in terms of resources. In this study, we evaluated the suitability of sample survey methods for Grevy’s zebra. We estimated population size using sample aerial counts for a known population of Grevy’s zebra in Lewa Wildlife Conservancy (LWC), providing the opportunity to test the accuracy of sample methods, while comparing resource costs with total count methods. We sampled one‐third of LWC using parallel 500‐ m strip transects at 1500‐ m intervals. The population estimate was comparable to the known population size and was less than half as expensive as the equivalent total count survey. Our results suggest sample aerial surveys provide an accurate and cost‐effective means of monitoring Grevy’s zebra and other endangered species in open habitats.     

Treatment of equine sarcoid in seven Cape mountain zebra (Equus zebra zebra)

Equine sarcoid has been diagnosed in endangered Cape mountain zebra (Equus zebra zebra) in at least two game reserves in South Africa, with prevalence as high as 53% in Bontebok National Park. Seven Cape mountain zebras with sarcoids were treated with either surgical excision, 5-fluorouracil, allogenous vaccine, or a combination of 5-fluorouracil and allogenous vaccine. One of the two sarcoids on one of the 5-fluorouracil-treated zebras was left untreated. The microscopic features of the tumors evaluated showed either all or most of the typical epidermal and dermal histologic features of equine sarcoid. The zebras were examined 2 yr posttreatment to determine outcome. All sarcoids had resolved except on the zebra on which one of the sarcoids was left untreated. The efficacy of the three treatment methods in Cape mountain zebra is encouraging.     

Long-term demography of a zebra mussel (Dreissena polymorpha) population

1. We used long‐term data and a simulation model to investigate temporal fluctuations in zebra mussel populations, which govern the ecological and economic impacts of this pest species. 2. The size of the zebra mussel (Dreissena polymorpha) population in the Hudson River estuary fluctuated approximately 11‐fold across a 13‐year period, following a cycle with a 2–4 year period. 3. This cycling was caused by low recruitment during years of high adult population size, rapid somatic growth of settled animals, and adult survivorship of 50% per year. 4. Adult growth and body condition were weakly correlated with phytoplankton biomass. 5. The habitat distribution of the Hudson's population changed over the 13‐year period, with an increasing proportion of the population spreading onto soft sediments over time. The character of soft‐sediment habitats in the Hudson changed because of large amounts (mean = 34 g DM m^?2) of empty zebra mussel shells now in the sediments. 6. Simulation models show that zebra mussel populations can show a range of long‐term trajectories, depending on the balance between adult space limitation, larval food limitation, and disturbance. 7. Effective understanding and management of the effects of zebra mussels and other alien species depend on understanding of their long‐term demography, which may vary across ecosystems.     

Effects of zebra mussel attachment on the foraging behaviour of a larval dragonfly,Macromia illinoiensis

1. Larvae of Macromia illinoiensis Walsh are often colonised by the zebra mussel, Dreissena polymorpha Pallas, a recent invader to North America. To determine how mussel attachment affects an individual's foraging behaviour, we quantified capture of Hexagenia limbata Hexes mayfly prey and the distance moved by newly‐molted final instars before and after an individual's colonisation with zebra mussels. 2. In night trials, larvae sprawled above the sand, and caught more mayflies than individuals in daytime trials, but the estimated distance travelled did not differ. When resting under a layer of sand with only its eyes exposed during the day, an individual could capture a mayfly prey using a sit‐and‐wait ambush strategy. When sprawled above the sand, some larvae caught prey that rested on their legs. 3. When mussel‐free, individuals captured more prey than they did when carrying zebra mussels, although mussel attachment per se did not affect the estimated distance that a larva moved. 4. During day trials, but not night ones, the increasing mussel load of colonised individuals decreased prey capture and the distance moved in an apparent step‐wise function. Although the number of mussels carried did not differ, night foragers carried a heavier load. Independent of time of the day, the distance an individual travelled when mussel‐free was predictive of the number of prey it caught when colonised, suggesting that the greater general activity of some individuals helped mitigate negative effects that mussel attachment had on prey capture. 5. Our results add to a growing number of negative effects of zebra mussel colonisation on sprawling and hiding dragonfly larvae. Although the impact of these costs on dragonfly populations remains to be determined, a decrease in this guild of predators whose life cycle spans aquatic and terrestrial habitats might have cascading effects across ecosystems.     

Anthrax outbreak among Grevy's zebra (Equus grevyi) in Samburu, Kenya

An anthrax outbreak occurred in the Wamba area of southern Samburu, Kenya, between December 2005 and March 2006. The outbreak affected equids including the endangered Grevy's zebras (Equus grevyi), plain zebras (Equis Burchelli) and donkeys (Equus asinus). Most of the deaths were localized in Nkaroni area just west of Wamba town. The diagnosis of anthrax was rapidly confirmed by bacteriological methods. The relevant government departments, including the Kenya Wildlife Service and Veterinary Department, and other stakeholders were promptly informed. Fifty‐three Grevy's zebra and 26 plains zebras died from anthrax. An equal number (eighteen) of adult male and female Grevy's zebras succumbed to the disease. The outbreak affected immature and mature individuals equally. The dead plain zebras included fifteen adult females, two adult males and nine immature individuals. The Veterinary Department responded by vaccinating livestock while Kenya Wildlife Service vaccinated 620 Grevy's zebras within southern Samburu. Examination of sites at which carcasses of animals which succumbed to the disease were burnt, revealed that unsupervised burning did not eliminate anthrax spores in 42% of the cases (n = 14). There is an urgent need to incorporate strategic wildlife disease monitoring in the struggle to save Grevy's zebras and other endangered species.     

Identification of 13 polymorphic microsatellite loci in the zebra finch,Taeniopygia guttata (Passeridae,Aves)

Polymorphic microsatellite loci were identified in order to determine paternity in a captive population of the zebra finch, Taeniopygia guttata. Primer sets from 93 published passerine microsatellite sequences were tested for cross‐species amplification. Thirteen loci were found to be polymorphic, of which, eight displayed null alleles and one locus (Ase50) was found to be Z‐chromosome linked.     

Grass greenness and grass height promote the resource partitioning among reintroduced Burchell's zebra and blue wildebeest in southern Mozambique

Differences in the selection of habitat and specific dietary items support resource partitioning and coexistence of sympatric African grazing herbivores, such as zebra and wildebeest. In Maputo Special Reserve (MSR), southern Mozambique, these two species were extirpated during the civil war (1977–1992); since 2010, they have been reintroduced into the Reserve. Identifying the resource selection by reintroduced species and how these species coexist, while utilising the same resources, is both of ecological interest and important for the management of wildlife communities and parks. This is a key application of our research. Therefore, the present study investigated resource partitioning between Burchell's zebra (Equus burchelli, Smuts 1832) and blue wildebeest (Connochaetes taurinus, Burchell 1823) in the MSR. We conducted the study from July 2016 to June 2017. The data were collected by direct observation, driving the vehicle along the reserve's roads that covered the vegetation communities where zebras and wildebeest are known to commonly occur. The composition of the diet and specific features of the grass grazed by the two species, including greenness, height, and the number of stems, were assessed. The widely available grass, Aristida barbicollis, contributed most to the diet of both herbivores. The dietary overlap between the two herbivores was higher during the dry season (95%) than wet season (86%). Resources partitioning appears to be determined, principally, by the height and greenness of the grass, with the zebra grazing taller grass, which may facilitate the access of the wildebeest to the greener, lower proportion of the forage. That results follow the expectation that, among native herbivores, overlap in resource use is not expected based on evolutionary segregation.     

Genetic variation and differentiation in captive and wild zebra finches (Taeniopygia guttata)

The zebra finch (Taeniopygia guttata) is a small Australian grassland songbird that has been domesticated over the past two centuries. Because it is easy to breed in captivity, it has become a widely used study organism, especially in behavioural research. Most work has been conducted on domesticated populations maintained at numerous laboratories in Europe and North America. However, little is known about the extent to which, during the process of domestication, captive populations have gone through bottlenecks in population size, leading to inbred and potentially genetically differentiated study populations. This is an important issue, because (i) behavioural studies on captive populations might suffer from artefacts arising from high levels of inbreeding or lack of genetic variation in such populations, and (ii) it may hamper the comparability of research findings. To address this issue, we genotyped 1000 zebra finches from 18 captive and two wild populations at 10 highly variable microsatellite loci. We found that all captive populations have lost some of the genetic variability present in the wild, but there is no evidence that they have gone through a severe bottleneck, as the average captive population still showed a mean of 11.7 alleles per locus, compared to a mean of 19.3 alleles/locus for wild zebra finches. We found significant differentiation between the captive populations (F(ST) = 0.062). Patterns of genetic similarity closely match geographical relationships, so the most pronounced differences occur between the three continents: Australia, North America, and Europe. By providing a tree of the genetic similarity of the different captive populations, we hope to contribute to a better understanding of variation in research findings obtained by different laboratories.     

Physiology in ecological niche modeling: using zebra mussel's upper thermal tolerance to refine model predictions through Bayesian analysis

Climate change and human-mediated dispersal are increasingly influencing species’ geographic distributions. Ecological niche models (ENMs) are widely used in forecasting species’ distributions, but are weak in extrapolation to novel environments because they rely on available distributional data and do not incorporate mechanistic information, such as species’ physiological response to abiotic conditions. To improve accuracy of ENMs, we incorporated physiological knowledge through Bayesian analysis. In a case study of the zebra mussel Dreissena polymorpha, we used native and global occurrences to obtain native and global models representing narrower and broader understanding of zebra mussel’ response to temperature. We also obtained thermal limit and survival information for zebra mussel from peer-reviewed literature and used the two types of information separately and jointly to calibrate native models. We showed that, compared to global models, native models predicted lower relative probability of presence along zebra mussel's upper thermal limit, suggesting the shortcoming of native models in predicting zebra mussel's response to warm temperature. We also found that native models showed improved prediction of relative probability of presence when thermal limit was used alone, and best approximated global models when both thermal limit and survival data were used. Our result suggests that integration of physiological knowledge enhances extrapolation of ENM in novel environments. Our modeling framework can be generalized for other species or other physiological limits and may incorporate evolutionary information (e.g. evolved thermal tolerance), thus has the potential to improve predictions of species’ invasive potential and distributional response to climate change.