The effects of habitat fragmentation on the demography and population genetic structure of Uromys Caudimaculatus

Habitat fragmentation can have an impact on a wide variety of biological processes including abundance, life history strategies, mating system, inbreeding and genetic diversity levels of individual species. Although fragmented populations have received much attention, ecological and genetic responses of species to fragmentation have still not been fully resolved. The current study investigated the ecological factors that may influence the demographic and genetic structure of the giant white-tailed rat (Uromys caudimaculatus) within fragmented tropical rainforests. It is the first study to examine relationships between food resources, vegetation attributes and Uromys demography in a quantitative manner. Giant white-tailed rat densities were strongly correlated with specific suites of food resources rather than forest structure or other factors linked to fragmentation (i.e. fragment size). Several demographic parameters including the density of resident adults and juvenile recruitment showed similar patterns. Although data were limited, high quality food resources appear to initiate breeding in female Uromys. Where data were sufficient, influx of juveniles was significantly related to the density of high quality food resources that had fallen in the previous three months. Thus, availability of high quality food resources appear to be more important than either vegetation structure or fragment size in influencing giant white-tailed rat demography. These results support the suggestion that a species’ response to fragmentation can be related to their specific habitat requirements and can vary in response to local ecological conditions. In contrast to demographic data, genetic data revealed a significant negative effect of habitat fragmentation on genetic diversity and effective population size in U. caudimaculatus. All three fragments showed lower levels of allelic richness, number of private alleles and expected heterozygosity compared with the unfragmented continuous rainforest site. Populations at all sites were significantly differentiated, suggesting restricted among population gene flow. The combined effects of reduced genetic diversity, lower effective population size and restricted gene flow suggest that long-term viability of small fragmented populations may be at risk, unless effective management is employed in the future. A diverse range of genetic reproductive behaviours and sex-biased dispersal patterns were evident within U. caudimaculatus populations. Genetic paternity analyses revealed that the major mating system in U. caudimaculatus appeared to be polygyny at sites P1, P3 and C1. Evidence of genetic monogamy, however, was also found in the three fragmented sites, and was the dominant mating system in the remaining low density, small fragment (P2). High variability in reproductive skew and reproductive success was also found but was less pronounced when only resident Uromys were considered. Male body condition predicted which males sired offspring, however, neither body condition nor heterozygosity levels were accurate predictors of the number of offspring assigned to individual males or females. Genetic spatial autocorrelation analyses provided evidence for increased philopatry among females at site P1, but increased philopatry among males at site P3. This suggests that male-biased dispersal occurs at site P1 and female-biased dispersal at site P3, implying that in addition to mating systems, Uromys may also be able to adjust their dispersal behaviour to suit local ecological conditions. This study highlights the importance of examining the mechanisms that underlie population-level responses to habitat fragmentation using a combined ecological and genetic approach. The ecological data suggested that habitat quality (i.e. high quality food resources) rather than habitat quantity (i.e. fragment size) was relatively more important in influencing giant white-tailed rat demographics, at least for the populations studied here . Conversely, genetic data showed strong evidence that Uromys populations were affected adversely by habitat fragmentation and that management of isolated populations may be required for long-term viability of populations within isolated rainforest fragments.

Stochastic demography and population dynamics in the red kangaroo Macropus rufus

1. Many organisms inhabit strongly fluctuating environments but their demography and population dynamics are often analysed using deterministic models and elasticity analysis, where elasticity is defined as the proportional change in population growth rate caused by a proportional change in a vital rate. Deterministic analyses may not necessarily be informative because large variation in a vital rate with a small deterministic elasticity may affect the population growth rate more than a small change in a less variable vital rate having high deterministic elasticity. 2. We analyse a stochastic environment model of the red kangaroo (Macropus rufus), a species inhabiting an environment characterized by unpredictable and highly variable rainfall, and calculate the elasticity of the stochastic growth rate with respect to the mean and variability in vital rates. 3. Juvenile survival is the most variable vital rate but a proportional change in the mean adult survival rate has a much stronger effect on the stochastic growth rate. 4. Even if changes in average rainfall have a larger impact on population growth rate, increased variability in rainfall may still be important also in long-lived species. The elasticity with respect to the standard deviation of rainfall is comparable to the mean elasticities of all vital rates but the survival in age class 3 because increased variation in rainfall affects both the mean and variability of vital rates. 5. Red kangaroos are harvested and, under the current rainfall pattern, an annual harvest fraction of c. 20% would yield a stochastic growth rate about unity. However, if average rainfall drops by more than c. 10%, any level of harvesting may be unsustainable, emphasizing the need for integrating climate change predictions in population management and increase our understanding of how environmental stochasticity translates into population growth rate.

Demography of three perennial grasses in a central Queensland eucalypt woodland.

The population dynamics of the palatable, perennial grasses Bothriochloa ewartiana (Domin) C.E.Hubb. (desert Mitchell grass), Chrysopogon fallax S.T.Blake (golden beard grass) and Heteropogon contortus (L.) P.Beauv. ex Roem. & Schult. (black speargrass), were studied in an extensive grazing study conducted in a eucalypt woodland within the Aristida-Bothriochloa pasture community in central Queensland between 1994 and 2000. Treatments were three grazing pressures based on light, medium and heavy utilisation of forage available at the end of summer and two timber treatments (trees intact and trees killed). Seasonal rainfall throughout this study was generally favourable for plant growth with no severe drought periods. Grazing pressure had a greater overall impact on plant dynamics than timber treatment, which had minimal impact. Grazing pressure had a large impact on H. contortus dynamics, an intermediate impact on B. ewartiana and no impact on C. fallax. Fluctuations in plant density of both B. ewartiana and C. fallax were small because both species were long lived with low levels of seedling recruitment and plant death, whereas fluctuations in H. contortus density were relatively high because of its relatively short life span and higher levels of both recruitment and death. Heavy grazing pressure increased the recruitment of B. ewartiana and H. contortus in some years but had no impact on that of C. fallax. Heavy grazing pressure reduced the survival of the original plants of both B. ewartiana and H.contortus but not of C. fallax. For H. contortus, the size of the original plants was larger where trees were killed than where trees were left intact and plants of the 1995 seedling cohort were larger in 1998 at heavy compared with those at light and medium grazing pressure. Grazing had a minor negative impact on the soil seed bank of H. contortus. Populations of all three species remained stable throughout this study, although the favourable seasonal rainfall experienced and the short duration of this study relative to the life span of these species may have masked longer term, deleterious impacts of heavy grazing pressure.

Demography of feral camels in central Australia and its relevance to population control

Since their release over 100 years ago, camels have spread across central Australia and increased in number. Increasingly, they are being seen as a pest, with observed impacts from overgrazing and damage to infrastructure such as fences. Irregular aerial surveys since 1983 and an interview-based survey in 1966 suggest that camels have been increasing at close to their maximum rate. A comparison of three models of population growth fitted to these, albeit limited, data suggests that the Northern Territory population has indeed been growing at an annual exponential rate of r = 0.074, or 8% per year, with little evidence of a density-dependent brake. A stage-structured model using life history data from a central Australian camel population suggests that this rate approximates the theoretical maximum. Elasticity analysis indicates that adult survival is by far the biggest influence on rate of increase and that a 9% reduction in survival from 96% is needed to stop the population growing. In contrast, at least 70% of mature females need to be sterilised to have a similar effect. In a benign environment, a population of large mammals such as camels is expected to grow exponentially until close to carrying capacity. This will frustrate control programs, because an ever-increasing number of animals will need to be removed for zero growth the longer that culling or harvesting effort is delayed. A population projection for 2008 suggests ~10 500 animals need to be harvested across the Northern Territory. Current harvests are well short of this. The ability of commercial harvesting to control camel populations in central Australia will depend on the value of animals, access to animals and the presence of alternative species to harvest when camels are at low density.

Refining the process of agent selection through understanding plant demography and plant response to herbivory

Understanding plant demography and plant response to herbivory is critical to the selection of effective weed biological control agents. We adopt the metaphor of 'filters' to suggest how agent prioritisation may be improved to narrow our choices down to those likely to be most effective in achieving the desired weed management outcome. Models can serve to capture our level of knowledge (or ignorance) about our study system and we illustrate how one type of modelling approach (matrix models) may be useful in identifying the weak link in a plant life cycle by using a hypothetical and an actual weed example (Parkinsonia aculeata). Once the vulnerable stage has been identified we propose that studying plant response to herbivory (simulated and/or actual) can help identify the guilds of herbivores to which a plant is most likely to succumb. Taking only potentially effective agents through the filter of host specificity may improve the chances of releasing safe and effective agents. The methods we outline may not always lead us definitively to the successful agent(s), but such an empirical, data-driven approach will make the basis for agent selection explicit and serve as testable hypotheses once agents are released.

Historical demography of Lantana camara L. reveals clues about the influence of land use and weather in the management of this widespread invasive species

Weather is a general stochastic influence on the life history of weeds. In contrast, anthropogenic disturbance (e.g. land use) is an important deterministic influence on weed demography. Our aim with this study was to investigate the relative contributions of land use and weather on the demography of Lantana camara (lantana), a weed of agricultural and natural habitats, based on the intensive monitoring of lantana populations under three land uses (viz. farm[pasture], and burnt and grazed forests) in subtropical Australia. Lantana populations were growing vigorously across all land uses (asymptotic population growth rate, lambda > 3). Examination of historical demography using retrospective perturbation analyses showed that weather was a strong influence on lantana demography with the transition from an El Nino (2008-09) to a La Nina (2009-10) year having a strong positive effect on population growth rate. This effect was most marked at the grazed site, and to a lesser extent at the burnt site, with seedling-to-juvenile and juvenile-to-adult transitions contributing most to these effects. This is likely the result of burning and grazing having eliminated/reduced interspecific competition at these sites. Prospective perturbation analyses revealed that lambda was most sensitive to proportionate changes in growth transitions, followed by fecundity and survival transitions. Examination of context-specific patterns in elasticity revealed that growth and fecundity transitions are likely to be the more critical vital rates to reduce lambda in wet years at the burnt and grazed forest sites, compared to the farm/pasture site. Management of lantana may need to limit the transition of juveniles into the adult stages, especially in sites where lantana is free from competition (e.g. in the presence of fire or grazing), and this particularly needs to be achieved in wet years. Collectively, these results shed light on aspects of spatial and temporal variation in the demography of lantana, and offer insights on its context-specific management.

Historical demography of Lantana camara L. reveals clues about the influence of land use and weather in the management of this widespread invasive species

Weather is a general stochastic influence on the life history of weeds. In contrast, anthropogenic disturbance (e.g. land use) is an important deterministic influence on weed demography. Our aim with this study was to investigate the relative contributions of land use and weather on the demography of Lantana camara (lantana), a weed of agricultural and natural habitats, based on the intensive monitoring of lantana populations under three land uses (viz. farm[pasture], and burnt and grazed forests) in subtropical Australia. Lantana populations were growing vigorously across all land uses (asymptotic population growth rate, λ > 3). Examination of historical demography using retrospective perturbation analyses showed that weather was a strong influence on lantana demography with the transition from an El Niño (2008–09) to a La Niña (2009–10) year having a strong positive effect on population growth rate. This effect was most marked at the grazed site, and to a lesser extent at the burnt site, with seedling-to-juvenile and juvenile-to-adult transitions contributing most to these effects. This is likely the result of burning and grazing having eliminated/reduced interspecific competition at these sites. Prospective perturbation analyses revealed that λ was most sensitive to proportionate changes in growth transitions, followed by fecundity and survival transitions. Examination of context-specific patterns in elasticity revealed that growth and fecundity transitions are likely to be the more critical vital rates to reduce λ in wet years at the burnt and grazed forest sites, compared to the farm/pasture site. Management of lantana may need to limit the transition of juveniles into the adult stages, especially in sites where lantana is free from competition (e.g. in the presence of fire or grazing), and this particularly needs to be achieved in wet years. Collectively, these results shed light on aspects of spatial and temporal variation in the demography of lantana, and offer insights on its context-specific management.

Studies on Plant Demography: Ranunculus Repens L., R. Bulbosus L. and R. Acris L.: III. A Mathematical Model Incorporating Multiple Modes of Reproduction

This paper presents a comparative population dynamics study of three closely related species of buttercups (Ranunculus repens, R. acris, and R. bulbosus). The study is based on an investigation of the behaviour of the seeds in soil under field conditions and a continuous monitoring of survival and reproduction of some 9000 individual plants over a period of 21/2 years in a coastal grassland in North Wales. The data were analysed with the help of an extension of Leslie's matrix method which makes possible an simultaneous treatment of vegetative and sexual reproduction. It was found that R. repens (a) depends more heavily on vegetative as compared with sexual reproduction, (b) shows indications of negatively density-dependent population regulation, and (c) exhibits little variation in population growth rates from site to site and from one year to the next. In contrast, R. bulbosus (a) depends exclusively on sexual reproduction, (b) shows indications of a positively density-dependent population behaviour, and (c) exhibits great variation in population growth rates from site to site and from one year to the next. R. acris exhibits an intermediate behaviour in all these respects. It is suggested that the attributes of R. repens are those expected of a species inhabiting a stable environment, while R. bulbosus exhibits some of the characteristics of a fugitive species.

Distribution, Demography, and Conservation of Lion-tailed Macaques (Macaca silenus) in the Anamalai Hills Landscape, Western Ghats, India

The status of the endemic and endangered lion-tailed macaque (Macaca silenus) has not been properly assessed in several regions of the Western Ghats of southern India. We conducted a study in Parambikulam Forest Reserve in the state of Kerala to determine the distribution, demography, and status of lion-tailed macaques. We laid 5km(2) grid cells on the map of the study area (644km(2)) and made four replicated walks in each grid cell using GPS. We gathered data on lion-tailed macaque group locations, demography, and site covariates including trail length, duration of walk, proportion of evergreen forest, height of tallest trees, and human disturbance index. We also performed occupancy modeling using PRESENCE ver. 3.0. We estimated a minimum of 17 groups of macaques in these hills. Low detection and occupancy probabilities indicated a low density of lion-tailed macaques in the study area. Height of the tallest trees correlated positively whereas human disturbance and proportion of evergreen forest correlated negatively with occupancy in grid cells. We also used data from earlier studies carried out in the surrounding Anamalai Tiger Reserve and Nelliyampathy Hills to discuss the conservation status in the large Anamalai Hills Landscape. This landscape harbors an estimated population of 1108 individuals of lion-tailed macaques, which is about one third of the entire estimated wild population of this species. A conservation plan for this landscape could be used as a model for conservation in other regions of the Western Ghats.

Intracellular demography and the dynamics of Salmonella enterica infections.

An understanding of within-host dynamics of pathogen interactions with eukaryotic cells can shape the development of effective preventive measures and drug regimes. Such investigations have been hampered by the difficulty of identifying and observing directly, within live tissues, the multiple key variables that underlay infection processes. Fluorescence microscopy data on intracellular distributions of Salmonella enterica serovar Typhimurium (S. Typhimurium) show that, while the number of infected cells increases with time, the distribution of bacteria between cells is stationary (though highly skewed). Here, we report a simple model framework for the intensity of intracellular infection that links the quasi-stationary distribution of bacteria to bacterial and cellular demography. This enables us to reject the hypothesis that the skewed distribution is generated by intrinsic cellular heterogeneities, and to derive specific predictions on the within-cell dynamics of Salmonella division and host-cell lysis. For within-cell pathogens in general, we show that within-cell dynamics have implications across pathogen dynamics, evolution, and control, and we develop novel generic guidelines for the design of antibacterial combination therapies and the management of antibiotic resistance.

Intracellular demography and the dynamics of Salmonella enterica infections.

An understanding of within-host dynamics of pathogen interactions with eukaryotic cells can shape the development of effective preventive measures and drug regimes. Such investigations have been hampered by the difficulty of identifying and observing directly, within live tissues, the multiple key variables that underlay infection processes. Fluorescence microscopy data on intracellular distributions of Salmonella enterica serovar Typhimurium (S. Typhimurium) show that, while the number of infected cells increases with time, the distribution of bacteria between cells is stationary (though highly skewed). Here, we report a simple model framework for the intensity of intracellular infection that links the quasi-stationary distribution of bacteria to bacterial and cellular demography. This enables us to reject the hypothesis that the skewed distribution is generated by intrinsic cellular heterogeneities, and to derive specific predictions on the within-cell dynamics of Salmonella division and host-cell lysis. For within-cell pathogens in general, we show that within-cell dynamics have implications across pathogen dynamics, evolution, and control, and we develop novel generic guidelines for the design of antibacterial combination therapies and the management of antibiotic resistance.

James's rule and causes and consequences of a latitudinal cline in the demography of John's Snapper (Lutjanus johnii) in coastal waters of Australia

Demographic parameters were derived from sectioned otoliths of John’s Snapper (Lutjanus johnii) from 4 regions across 9° of latitude and 23° of longitude in northern Australia. Latitudinal variation in size and growth rates of this species greatly exceeded longitudinal variation. Populations of John’s Snapper farthest from the equator had the largest body sizes, in line with James’s rule, and the fastest growth rates, contrary to the temperature-size rule for ectotherms. A maximum age of 28.6 years, nearly 3 times previous estimates, was recorded and the largest individual was 990 mm in fork length. Females grew to a larger mean asymptotic fork length (L∞) than did males, a finding consistent with functional gonochorism. Otolith weight at age and gonad weight at length followed the same latitudinal trends seen in length at age. Length at maturity was ~72–87% of L∞ and varied by ~23% across the full latitudinal gradient, but age at first maturity was consistently in the range of 6–10 years, indicating that basic growth trajectories were similar across vastly different environments. We discuss both the need for complementary reproductive data in age-based studies and the insights gained from experiments where the concept of oxygen- and capacity-limited thermal tolerance is applied to explain the mechanistic causes of James’s rule in tropical fish species.

Scales of spatial variation in demography of a large coral-reef fish: an exception to the typical model?

Spatial variation in demographic parameters of the red throat emperor (Lethrinus miniatus) was examined among 12 coral reefs in three geographic regions (Townsville, Mackay, and Storm Cay) spanning over 3° of latitude of the Great Barrier Reef, Australia. Estimates of demographic parameters were based on age estimates from counts of annuli in whole otoliths because there was no significant difference in age estimates between whole and sectioned otoliths. There were significant regional differences in age structures, rates of somatic and otolith growth, and total mortality. The Townsville region was characterized by the greatest proportion of older fish, the smallest maximum size, and the lowest rates of otolith growth and total mortality. In contrast the Mackay region was characterized by the highest proportion of younger fish, the largest maximum size, and the highest rates of otolith growth and total mortality. Demographic parameters for the Storm Cay region were intermediate between the other two regions. Historic differences in fishing pressure and regional differences in productivity are two alternative hypotheses given to explain the regional patterns in demographic parameters. All demographic parameters were similar among the four reefs within each region. Thus, subpopulations with relatively homogeneous demographic parameters occurred on scales of reef clusters. Previous studies, by contrast, have found substantial between-reef variation in demographic parameters within regions. Thus spatial variation in demographic parameters for L. miniatus may differ from what is assumed typical for a coral-reef fish metapopulation.