Allozyme variation and differentiation in African and Indian rhinoceroses

We studied variation at 25 to 31 allozymic loci in African and Asian rhinoceroses. Four taxa in three genera were examined: African Ceratotherium simum simum (northern white rhinoceros), C. s. cottoni (southern white rhinoceros), Diceros bicornis (black rhinoceros), and Rhinoceros unicornis (Indian rhinoceros). Extremely small amounts of intraspecific variation were observed in sample sizes of 2 to 10 presumably unrelated individuals per taxon: P = .00-.10, H = 0.00-0.02. We examined demographic bottlenecks and sampling errors as possible reasons for the low levels of detectable variation. The very small intraspecific genetic distance (D = 0.005) between the two living white rhinoceros subspecies is far less than the distance that has been reported for other mammal subspecies. The mean D value of 0.32 +/- 0.11 between the two African genera was also less than expected given the divergence time of greater than 7 million years suggested by the fossil record. Rhinoceroses may be evolving more slowly at the structural gene loci than are some other mammal groups. The estimate of D = 1.05 +/- 0.24 for the African-Indian split supports this idea, as the lineage diverged at least 26 million years ago. Our results contribute to the currently available scientific information on which management decisions aimed toward saving endangered rhinoceroses should be based.     

Contemporary and historical impacts of megaherbivores on the population structure of tree euphorbias in South African subtropical thicket

African elephant and black rhinoceros – both megaherbivores – impact negatively on the abundance of succulent plants, including tree succulents, in South Africa's subtropical thicket. We sampled 35 sites in subtropical thicket to assess historical and contemporary impacts of African elephant and black rhinoceros on the population structure of three species of succulent tree euphorbia. Population age structures were highly variable and showed no species-specific effects. Almost half the sites had growing populations dominated by young individuals. Sites having contemporary impacts had significantly fewer individuals in the 30–75 years age range, but this effect declined with increasing terrain slope. Eighty-one percent of sites assumed to have been impacted by megaherbivores historically had individuals that predated rhino and elephant extirpation in the region. This and other population age structure data indicate that megaherbivores and tree euphorbias coexisted historically on terrain of low relief. However, in areas not subject to contemporary impacts, euphorbia populations are likely to be much higher now than historically, owing to population relaxation after the extirpation of megaherbivores in the mid 1800s. Given the sensitivity of tree euphorbias to megaherbivore impacts, managers should consider monitoring the population structure of these species in order to identify thresholds of potential concern regarding megaherbivore impacts.     

Testing the interaction between environmental variation and dispersal strategy on population dynamics using a soil mite experimental system

Dispersal can play an important role in both the local and regional dynamics of populations. Empirical studies have shown that the proportion of individuals dispersing is often density dependent, which may have implications for the effect of dispersal on populations. In this study, we manipulate the dispersal strategy of adults within two-patch laboratory populations of soil mites and compare the consequences of fixed (density-independent) and density-dependent dispersal in environments of constant and temporally varying resource availability. Effects of dispersal on population dynamics were dependent on the presence of environmental variation. Both dispersal strategies tended to spatially homogenize the population abundance of adults in a variable environment. However, the effect of environmental variation on mean adult abundance was greater with density-dependent dispersal than with fixed dispersal. Adult dispersal did not affect juvenile or egg abundance. This study demonstrates the potential significance of density-dependent dispersal for population dynamics, but emphasizes the role of the environmental context.     

Spatial ecology of large herbivore populations

Models of the dynamics of large herbivore populations represent density feedbacks on the population growth rate either directly or indirectly through interactions with vegetation resources. Neither approach incorporates the spatial heterogeneity that is an essential feature of most natural environments, and modifies the population dynamics generated. This is especially true for large herbivores exploiting food resources that are rooted in space but temporally variable in quantity and quality both seasonally and annually. In this review I explore how environmental variation at different spatiotemporal scales influences the abundance of herbivore populations controlled via resources, predators or social mechanisms. Changes in abundance can be spatially disparate and dependent on different resource components at different stages of the seasonal cycle, including buffer resources restricting population crashes in extremely adverse years. GPS telemetry enables movement responses generating spatial patterns to be documented in fine spatiotemporal detail, including migration and dispersal. Models incorporating spatial heterogeneity either implicitly or explicitly are outlined, exemplifying how herbivores cope with temporal variability by exploiting spatial variability in resources and conditions. Global human dominance is generating widened climatic variation while opportunities for herbivore movements are becoming constricted. Theoretical population ecologists need to shift their focus from the workings of demographic structure towards effects of changing environmental contexts, in order to project the likely trajectories of large herbivore populations through the Anthropocene.     

Characterization of microsatellite loci in the black rhinoceros (Diceros bicornis) and white rhinoceros (Ceratotherium simum): their use for cross-species amplification and differentiation between the two species

As the population sizes of the black and white rhinoceroses continues to decline, more efforts are needed in multiple areas to help with the conservation efforts. One area being explored is the use of genetic diversity information to aid conservation decisions. In this study, we designed 21 microsatellite primers for white and black rhinoceroses, 16 and 17 of which amplified bands in the white and black rhinoceros, respectively. Out of these primers all 16 were polymorphic in the white rhinoceros and 12 of the 17 were polymorphic in the black rhinoceros. The mean number of alleles was 3.31 and 2.12, the expected heterozygosities were 0.420 and 0.372, and the observed heterozygosities were 0.436 and 0.322 for the white and black rhinoceroses, respectively. Seven of the primers produced different allele sizes and variations that distinguished between black and white rhinoceroses. Further genetic analyses with larger wild population sample sizes and markers are recommended to obtain a better understanding of the genetic structure of the black and white rhinoceros populations in order to be useful in the conservation efforts of these critically endangered species. A. Kilbourn—In memoriam.     

Environmental niche factor analysis (ENFA) relates environmental parameters to abundance and genetic diversity in an endangered amphibian, the fire-bellied-toad (Bombina bombina)

Increasing attempts are made to understand the factors responsible for both the demographic and genetic depletion in amphibian populations. Landscape genetics aims at a spatially explicit correlation of genetic population parameters to landscape features. Using data from the endangered fire-bellied toad Bombina bombina in Brandenburg (Northeastern Germany), we performed an environmental niche factor analysis (ENFA), relating demographic (abundance) and genetic (diversity at 17 microsatellite loci and partial sequences of the mitochondrial control region in 434 individuals from 16 populations) parameters to ecological and anthropogenic variables such as temperature, precipitation, soil wetness, water runoff, vegetation density, and road/traffic impact. We found significant correlations between road disturbance and observed heterozygosity and between soil wetness and mitochondrial diversity. As the influences of the environmental variables can differ between different indicators for genetic diversity, population size and abundance data, our ENFA-based landscape genetics approach allows us to put various aspects of long- versus short term effective population size and genetic connectivity into an ecological and spatially explicit context, enabling potentially even forecast assessment under future environmental scenarios.     

Mating system and reproductive skew in the black rhinoceros

Only approximately 2600 black rhinoceros survive today, mainly in small, isolated populations of < 100 animals. The management of remaining black rhinoceros populations aims at preserving natural levels of genetic relatedness and optimizing breeding success, which requires an accurate knowledge of the mating system, reproductive skew and effective population size. DNA was extracted from faecal samples from a community of 35 wild black rhinoceros, and microsatellites were used to characterize patterns of paternity of 19 offspring born from eight females in this community. Paternity could be ascribed unequivocally for each offspring. Although our conclusions must be considered tentative, we present the first genetic evidence that black rhinoceros males are polygynous, with a high variance in reproductive success. We also describe a noninvasive management tool that can be used for the genetic management of this critically endangered species, both in the wild and in captivity.     

Demographic responses to weather fluctuations are context dependent in a long‐lived amphibian

Weather fluctuations have been demonstrated to affect demographic traits in many species. In long‐lived organisms, their impact on adult survival might be buffered by the evolution of traits that reduce variation in interannual adult survival. For example, skipping breeding is an effective behavioral mechanism that may limit yearly variation in adult survival when harsh weather conditions occur; however, this in turn would likely lead to strong variation in recruitment. Yet, only a few studies to date have examined the impact of weather variation on survival, recruitment and breeding probability simultaneously in different populations of the same species. To fill this gap, we studied the impact of spring temperatures and spring rainfall on survival, on reproductive skipping behavior and on recruitment in five populations of a long‐lived amphibian, the yellow‐bellied toad (Bombina variegata). Based on capture–recapture data, our findings demonstrate that survival depends on interactions between age, population and weather variation. Varying weather conditions in the spring result in strong variation in the survival of immature toads, whereas they have little effect on adult toads. Breeding probability depends on both the individual's previous reproductive status and on the weather conditions during the current breeding season, leading to high interannual variation in recruitment. Crucially, we found that the impact of weather variation on demographic traits is largely context dependent and may thus differ sharply between populations. Our results suggest that studies predicting the impact of climate change on population dynamics should be taken with caution when the relationship between climate and demographic traits is established using only one population or few populations. We therefore highly recommend further research that includes surveys replicated in a substantial number of populations to account for context‐dependent variation in demographic processes.     

Rainfall and population dynamics of Grey Pileated Finch <Emphasis Type="Italic">Coryphospingus pileatus</Emphasis> (Aves: Passeriformes) in a Neotropical dry forest

In tropical dry environments rainfall periodicity may affect demographic parameters, resulting in fluctuations in bird abundance. We used capture–recapture data for the Grey Pileated Finch from a Neotropical dry forest to evaluate the hypothesis that intra- and inter-annual survival, individuals entrance and population abundance, are related to local rainfall. Sampling occurred across 3 years, with individuals captured, tagged and evaluated for age and presence of brood patch every 14 days. Using the POPAN formulation, we generated demographic models to evaluate study population temporal dynamics. Best-fit models indicated a low apparent annual survival in the first year (16%) compared to other years (between 47 and 62%), with this low value associated with an extreme drought. The abundance of juveniles at each capture occasion was significantly dependent on the accumulated precipitation in the previous 14 days, and the juvenile covariate was a strong predictor of the intra-annual entrance probability (natality). Individuals entrance during the reproductive period corresponded to 53, 52 and 75% of total ingress for each year, respectively. The trend in sampled population size indicated positive exponential growth (N_initial = 50, N_last = 600), with intra-annual fluctuations becoming progressively more intense. Low survival was relevant during population decline at study onset, while at study end intense Individuals entrance promoted rapid population growth. Thus, the indirect effects of rainfall and the combined effect of two demographic rates operated synergistically on the immediate population abundance of Grey Pileated Finch, an abundant bird in a Neotropical dry forest.     

Density‐dependent regulation of the critically endangered black rhinoceros population in Ithala Game Reserve,South Africa

Ensuring the persistence of populations of endangered species requires an understanding of, and response to, the causes of population declines. Species occurring in small populations are vulnerable to stochastic problems that are environmental, demographic, or genetic in nature and can reduce survival as much as the deterministic threats of habitat degradation. Critically endangered black rhinoceros (Diceros bicornis) populations declined throughout Africa since 1960, with numbers steadily increasing at a continental level, but remaining lower than three generations ago. However, size, demographics, trends, and factors affecting these, are poorly known. We used 18 years (1990–2008) of long‐term sightings data from Ithala Game Reserve, KwaZulu‐Natal, South Africa, to determine population estimates, growth rate and fecundity over time, as well as sex and age structure and age‐specific probabilities of survival. Calf survivorship between the ages of 0 and 1 year was 74% for females and 94% for males. Age‐specific survivorship for both sexes was significantly higher from yearling to subadult age‐classes (1–6 years) than for adults (7–30 years). The most frequent cause of mortality was attributed to unknown causes while fighting injuries was recorded as the second most common cause of mortality, particularly among subadult and adult males. There was no significant difference in the sex ratios at birth, although the proportion of females in the population was 0.58. There was strong evidence for density‐dependent regulation, with density in conception year a key driver of population performance (birth rate). The population does not appear to be at ecological carrying capacity; however, social effects are delaying conception. To mitigate density‐dependent social effects, we recommend an adaptive management strategy of pre‐selecting individuals for removal from the reserve, so as to maintain stability in the social organization of the population.     

THE VALIDITY OF CENSUSING BLACK RHINOCEROS POPULATIONS FROM THE AIR

The efficacy of censusing black rhinoceros ( Diceros bicornis Linnaeus) populations from the air was tested. Repeated aerial counts were made of the black rhinoceros population centered on Qlduvai Gorge, Tanzania, where the number of rhinoceros on the ground is known accurately. Even under the most ideal conditions only 50% of the population was detected by observers in an aircraft, and factors accounting for this are considered. It is concluded that the light aircraft is of limited value in providing estimates of black rhinoceros populations, and estimates based purely on aerial counts are subject to considerable variation.     

Demographic population structure of black howler monkeys in fragmented and continuous forest in Chiapas,Mexico: Implications for conservation

For wild primates, demography studies are increasingly recognized as necessary for assessing the viability of vulnerable populations experiencing rapid environmental change. In particular, anthropogenic changes such as habitat loss and fragmentation can cause ecological and behavioral changes in small, isolated populations, which may, over time, alter population density and demographic structure (age/sex classes and group composition) in fragment populations relative to continuous forest populations. We compared our study population of Endangered black howler monkeys (Alouatta pigra) in 34 forest fragments around Palenque National Park (PNP), Mexico (62 groups, 407 individuals), to the adjacent population in PNP, protected primary forest (21 groups, 134 individuals), and to previous research on black howlers in fragments in our study area (18 groups, 115 individuals). We used χ² and Mann–Whitney U tests to address the questions: (a) what is the current black howler demographic population structure in unprotected forest fragments around PNP? (b) How does it compare to PNP's stable, continuous population? (c) How has it changed over time? Compared to the PNP population, the fragment populations showed higher density, a significantly lower proportion of multimale groups, and significantly fewer adult males per group. The population's age/sex structure in the fragmented landscape has been stable over the last 17 years, but differed in a higher proportion of multifemale groups, higher density, and higher patch occupancy in the present. In the context of conservation, some of our results may be positive as they indicate possible population growth over time. However, long-term scarcity of adult males in fragments and associated effects on population demographic structure might be cause for concern, in that it may affect gene flow and genetic diversity. The scarcity of adult males might stem from males experiencing increased mortality while dispersing in the fragmented landscape, whereas females might be becoming more philopatric in fragments.     

Conserving the African rhinoceros

The Southern African rhinoceros population has been dwindling through the years. A lot of effort has been put toward coming up with the most effective way of protecting the black and white rhinoceros population in the world yet still the animals remain endangered despite the enormous global efforts to protect the species. Such conservation efforts include; establishment of sanctuaries, wildlife farming, dehorning, trade and market controls, listing of species as endangered, use of biotechnology and biotelemetry. However useful, each of these methods has its own strengths and weaknesses relative to the success of rhinoceros conservation. It however has been noted in this review that dehorning is the most effective method as it involves removal of the horn hence extinguishing the need for poaching. Despite the high success rate it has also been noted that this requires a great deal of skill and finance hence limiting the application of this technique in developing nations. Sanctuaries provide an ideal breeding environment that is better than the wild and enhanced breeding accounted for a significant increase in rhinoceros population where it has been applied. It was however been conclude that a holistic conservation approach is required to conserve the African rhinoceros.     

Conservation genetics of the black rhinoceros, <Emphasis Type="Italic">Diceros bicornis bicornis</Emphasis>, in Namibia

Poaching and habitat destruction across sub-Saharan Africa brought the black rhinoceros (Diceros bicornis) close to extinction. Over the past few decades, however, one of four subspecies, D. b. bicornis, has experienced a significant population increase as a consequence of its protection within Etosha National Park (ENP), Namibia. We report here on the level and spatial distribution of black rhinoceros genetic diversity within ENP. Using nine microsatellite loci, genetic variation was assessed from 144 individuals. Our results are consistent with the observation of lower levels of genetic diversity in D. b. bicornis, when compared to D. b. michaeli, but greater diversity when compared to D. b. minor. We also showed that ENP’s black rhino genetic diversity is well represented in Waterberg National Park, originally founded with ENP individuals. We found no genetic signature of a recent bottleneck in ENP, however, suggesting that the genetic diversity within ENP has not been adversely affected by the recent severe population decline. Using Bayesian clustering methods, we observed no significant population structure within ENP, but positive spatial genetic correlation is observed at distances up to 25 km. This relationship exists in females but not males, suggesting reduced dispersal among females, the first evidence of limited female dispersal or philopatry in any species of rhinoceros.     

Do species population parameters and landscape characteristics affect the relationship between local population abundance and surrounding habitat amount?

In landscape ecology, correlational approaches are typically used to analyse links between local population abundance, and the surrounding habitat amount to estimate biologically-relevant landscape size (extent) for managing endangered or pest populations. The direction, strength, and spatial extent of the correlations are then sometimes interpreted in terms of species population parameters. Here we simulated the population dynamics of generalized species across spatially explicit landscapes that included two distinct habitat types. We investigated how characteristics of a landscape (structure), including the variation in habitat quality and spatial aggregation of the habitat, and the precise population-dynamic properties of the simulated species (dispersal and growth rates) affect the correlation between population abundance and amount of surrounding favourable habitat in the landscape. To evaluate these spatial extents of correlation, proportions of favourable habitat were calculated within several circles of increasing diameter centred on sampling patches of favourable habitat where population abundance was recorded.We found that the value of the correlation coefficients between population abundance and amount of surrounding favourable habitat depended on both population dynamic parameters and landscape characteristics. Coefficients of correlation increased with the variation in habitat quality and the aggregation of favourable habitat in the landscape, but decreased with the dispersal distance. The distance at which the correlation was maximized was sensitive to an interaction between the level of aggregation of the habitat and the dispersal distance; whereas the greatest distance at which a significant correlation occurred was more sensitive to the variation in habitat quality. Our results corroborate the view that correlational analyses do provide information on the local population dynamics of a species in a given habitat type and on its dispersal rate parameters. However, even in simplified, model frameworks, direct relationships are often difficult to disentangle and global landscape characteristics should be reported in any studies intended to derive population-dynamic parameters from correlations. Where possible, replicated landscapes should be examined in order to control for the interaction between population dynamics and landscape structure. Finally, we recommend using species-specific, population-dynamic modelling in order to interpret correctly the observed patterns of correlation in the landscape.     

Comparison of whole mitochondrial genome sequences of northern and southern white rhinoceroses (<Emphasis Type="Italic">Ceratotherium simum</Emphasis>): the conservation consequences of species definitions

The probable extinction of the last confirmed population of northern white rhinoceros (Ceratotherium simum cottoni) in the world has ignited debate regarding its species status compared to the southern white rhinoceros (Ceratotherium simum simum). Previous studies, based on partial mitochondrial sequences, have reported conflicting results regarding the species status of the northern white rhinoceros. We use whole mitochondrial genomes obtained using Next Generation Sequencing of four northern white rhinoceros and three southern white rhinoceros using novel primers in three overlapping fragments. Phylogenetic relationships were constructed, using Maximum Likelihood, and recovered monophyletic clades for northern white rhinoceros and southern white rhinoceros. The divergence time between the two mitochondrial DNA lineages was estimated to be between 0.46 and 0.97 million years ago using Bayesian inferences. Since there are currently only three surviving northern white rhinoceros individuals these results put into sharp focus the ongoing debate regarding the methods of species definition, and in particular the consequences of such definitions in conservation management of endangered species and subspecies. We conclude that the designation of sub-species status is more applicable to northern and southern white rhinoceros.     

Effects of two types of herbivores on the population dynamics of a perennial herb

The joint effects of multiple herbivores on their shared host plant have received increasing interest recently. The influence of herbivores on population dynamics of their host plants, especially the relative roles of different types of damage, is, however, still poorly understood. Here, we present a modelling approach, including both deterministic and stochastic matrix modelling, to be used in estimating fitness effects of multiple herbivores on perennial plants. We examined the effects and relative roles of two specialist herbivores, a pre-dispersal seed predator, Euphranta connexa, and a leaf-feeding moth, Abrostola asclepiadis, on the population dynamics and long-term fitness of their shared host plant, a long-lived perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). We collected demographic data during 3 years and combined these data with the effects of natural levels of herbivory measured from the same individuals. We found that both seed predation and leaf herbivory reduced population growth of V. hirundinaria, but only very high damage levels changed the growth trend of the vigorously growing study populations from positive to negative. Demographic modelling indicated that seed predation had a greater impact on plant population growth than leaf herbivory. The effect of leaf herbivory was weaker and diminished with increasing level of seed predation. Evaluation of individual fitness components, however, suggested that leaf herbivory contributed more strongly to host plant fitness than seed predation. Our results emphasize that understanding the effects of a particular herbivore on plant population dynamics requires also knowledge on other herbivores present in the system, because the effect of a particular type of herbivory on plant population dynamics is likely to vary according to the intensity of other types of herbivory. Furthermore, evaluating herbivore impact from using individual fitness components does not necessarily reflect the long-term effects on total plant fitness.     

Demographic heterogeneity impacts density-dependent population dynamics

Among-individual variation in vital parameters such as birth and death rates that is unrelated to age, stage, sex, or environmental fluctuations is referred to as demographic heterogeneity. This kind of heterogeneity is prevalent in ecological populations, but is almost always left out of models. Demographic heterogeneity has been shown to affect demographic stochasticity in small populations and to increase growth rates for density-independent populations. The latter is due to ??cohort selection,?? where the most frail individuals die out first, lowering the cohort??s average mortality as it ages. The importance of cohort selection to population dynamics has only recently been recognized. We use a continuous-time model with density dependence, based on the logistic equation, to study the effects of demographic heterogeneity in mortality and reproduction. Reproductive heterogeneity is introduced in three ways: parent fertility, offspring viability, and parent?Coffspring correlation. We find that both the low-density growth rate and the equilibrium population size increase as the magnitude of mortality heterogeneity increases or as parent?Coffspring phenotypic correlation increases. Population dynamics are affected by complex interactions among the different types of heterogeneity, and trade-off scenarios are examined which can sometimes reverse the effect of increased heterogeneity. We show that there are a number of different homogeneous approximations to heterogeneous models, but all fail to capture important parts of the dynamics of the full model.     

Important population viability analysis parameters for giant pandas (Aliuropoda melanoleuca)

Population viability analysis(PVA) is a tool to evaluate the risk of extinction for endangered species and aid conservation decision-making.The quality of PVA output is dependent on parameters related to population dynamics and life-history;however,it has been difficult to collect this information for the giant panda(Aliuropoda melanoleuca),a rare and endangered mammal native to China,confined to some 30 fragmented habitat patches.Since giant pandas are long-lived,mature late,have lower reproductive rates,and show little sexual dimorphism,obtaining data to perform adequate PVA has been difficult.Here,we develop a parameter sensitivity index by modeling the dynamics of six giant panda populations in the Minshan Mountains,in order to determine the parameters most influential to giant panda populations.Our data shows that the giant panda populations are most sensitive to changes in four female parameters:initial breeding age,reproductive rate,mortality rate between age 0 and 1,and mortality rate of adults.The parameter sensitivity index strongly correlated with initial population size,as smaller populations were more sensitive to changes in these four variables.This model suggests that demographic parameters of females have more influence on the results of PVA,indicating that females may play a more important role in giant panda population dynamics than males.Consequently,reintroduction of female individuals to a small giant panda population should be a high priority for conservation efforts.Our findings form a technical basis for the coming program of giant panda reintroduction,and inform which parameters are crucial to successfully and feasibly monitoring wild giant panda populations.     

Demography of cape petrels in response to environmental changes

Predicting the responses of populations in changing environments is an important task for ecologists. Understanding the population dynamics of high-latitude breeding species is critical given the particularly rapid environmental changes that occur in these regions. Using long-term mark–resighting data acquired over 53-years in Pointe Géologie, Terre Adélie, Antarctica, we estimated age-specific demographic parameters and evaluated the effect of the environment on survival of a poorly known species, the cape petrel Daption capense. We then modeled the dynamics of this population using a life-history model and performed prospective and retrospective analyses to estimate the sensitivity of the population growth rate to demographic parameters, and to quantify their relative contribution. Survival of cape petrel increased with age, being 0.610 (±0.193) for juveniles, 0.739 (±0.158) for individuals from 2 to 4, and 0.920 (±0.031) for older individuals. Minimum age at first reproduction was 3 years old, the age at which all birds were recruited was 14 years, and mean age at first reproduction was 9.05 (±2.06) years. Adult survival increased over time and was positively correlated with the southern annular mode (SAM). The stochastic population growth rate was estimated at 1.019, and adult survival over age 5 made the largest contribution to variance of the population growth rate. Sensitivity analyses revealed that population regulation was mainly driven by the SAM. Our results suggest that despite the decrease in breeding success, the population of cape petrels at Pointe Géologie increased due to the increase in immature and adult survival.