Small mammal assemblages in a disturbed tropical landscape at Pozuzo,Peru

Deforestation in the Peruvian pre-montane forests has increased alarmingly in recent years. Particularly, fragmentation in these forests takes place against a background of very extensive continuous forest. Until now, no studies have detailed how Peruvian small mammals responded to forest fragmentation. In this paper, we report the results of an assessment of nonvolant small mammal fauna in forest remnants (one large fragment and one small fragment) and cattle pasture in a disturbed landscape at Pozuzo, formerly a pre-montane tropical forest in central Peru. In each forest remnant, we quantify habitat parameters and identified small mammals to know if there is a relationship between forest remnant size or microhabitat structure and small mammal assemblages. In addition, we investigated the influence of habitat structure on the presence of small mammal species. We did not find any evidence that habitat structure in large fragment was greater than small fragment. Actually, both forest remnants presented habitat features typical of forests subject to higher levels of disturbance. We found that abundance of small trees (10 - 19 cm DBH) was positively associated with the presence of Marmosops noctivagus, but negatively with the presence of Oecomys bicolor. Likewise, the presence of the arboreal Rhipidomys leucodactylus was associated with the abundance of larger trees (DBH > 50 cm). We recorded 12 small mammal species in large fragment, 9 in small fragment and 3 in cattle pasture. No significant differences were found in species richness between both fragments, but marsupial abundance in the small fragment was greater than large fragment. At Pozuzo, we recorded 9 species restricted to forest habitat and 3 to forest and cattle pastures. However, it is necessary to assess more fragments to improve our knowledge about species survival in this landscape.     

Extinction and colonization processes in subpopulations of five neotropical small mammal species

We report the extinction and colonization rates of five sympatric small mammal species at a semiarid locality in north central Chile. We provide information based on 6 years of monitoring on how colonization and extinction rates change according to landscape features (slope aspect) and on their relationship to populations size, population variability, and body size. We found that: (1) for all species in the assemblage, extinction rates of subpopulations from equatorial-facing slopes were significantly lower than those in polar-facing slopes, (2) population size was the most important factor determining extinction rates, (3) colonization rates did not vary between slopes, and were affected by population size only in equatorial-facing slopes, and (4) most species had higher extinction than colonization rates. Persistence of the metapopulation system for all five small mammal species appears to be fueled by repeated colonization events.     

Environmental and vegetation variables have a greater influence than habitat fragmentation in structuring lizard communities in remnant urban bushland

Abstract The expansion of urban areas and adjacent farming land into natural landscapes modifies habitats and produces small isolated pockets of native vegetation. This fragmentation of the natural habitat subdivides animal communities, reduces population sizes and increases vulnerability to extinction. In this paper we investigate whether fragmentation decreases lizard species richness, composition, overall abundance and abundance at the species level. Urban remnants consisting of five small (< 10 ha) and four large (> 10 ha) fragments of natural bushland were paired with continuous bushland areas located near Hobart, Tasmania, Australia. These remnants were surveyed six times, using pitfall traps, from November 2001 to March 2002. Lizard species richness and abundance were not significantly influenced by habitat fragmentation or fragment size. Egernia whitii was the only lizard species significantly influenced by fragment size, and was only present in large fragments and continuous bush. Vegetation type and structure as well as environmental variables (geology and aspect) influenced the structure of reptile communities. Lizard species that were able to use a number of different habitat types were found to persist at most sites, irrespective of fragment size. Edge environment did not significantly influence lizard species richness or abundance in remnant areas. Lizard species richness was significantly lower in sites that had a high ratio of exotic to native plant species. Therefore, if remnants continue to be invaded by exotic plants, lizard species that require native plant communities will become increasingly vulnerable to local extinction. Our results suggest that lizard species requiring specialized habitats, such as E. whitii, may persist in large urban remnants rather than small urban remnants because large reserves are more likely to encompass rare habitats, such as rocky outcrops. Habitat heterogeneity, rather than size, may be the key to their persistence.     

Non-volant small mammal community responses to fragmentation of kerangas forests in Brunei Darussalam

This paper summarises the effects of kerangas forest fragmentation on the non-volant small mammal community resident in the fragments based on a long term study conducted in Brunei Darussalam. These effects are shown as eight responses—a significant reduction in the species richness and diversity in relation to size of fragments, the disappearance of species with large home ranges and specific habitat requirements, invasion of generalists/opportunists into the fragments from degraded habitats, absence of predators, a possible reduction in the activity space of generalists in fragments, erosion in the trophic structure of the small mammal community, changes in the small mammal abundance in relation to fragment size and dominance of one species over other co-occurring species in an isolated fragment. These responses are briefly explained and their significance discussed.     

Trophic structure stability and extinction dynamics of beetles (Coleoptera) in tropical forest fragments

A first analysis of the stability of trophic structure following tropical forest fragmentation was performed in an experimentally fragmented tropical forest landscape in Central Amazonia. A taxonomically and trophically diverse assemblage of 993 species of beetles was sampled from 920 m² of leaf litter at 46 sites varying in distance from forest edge and fragment area. Beetle density increased significantly towards the forest edge and showed non-linear changes with fragment area, due to the influx of numerous disturbed-area species into 10 ha and 1 ha fragments. There was a marked change in species composition with both decreasing distance from forest edge and decreasing fragment area, but surprisingly this change in composition was not accompanied by a change in species richness. Rarefied species richness did not vary significantly across any of the sites, indicating that local extinctions of deep forest species were balanced by equivalent colonization rates of disturbed-area species. The change in species composition with fragmentation was non-random across trophic groups. Proportions of predator species and xylophage species changed significantly with distance from forest edge, but no area-dependent changes in proportions of species in trophic groups were observed. Trophic structure was also analysed with respect to proportions of abundance in six trophic groups. Proportions of abundance of all trophic groups except xylomycetophages changed markedly with respect to both distance from forest edge and fragment area. Local extinction probabilities calculated for individual beetle species supported theoretical predictions of the differential susceptibility of higher trophic levels to extinction, and of changes in trophic structure following forest fragmentation. To reduce random effects due to sampling error, only abundant species (n = 46) were analysed for extinction probabilities, as defined by absence from samples. Of these common species, 27% had significantly higher probabilities of local extinction following fragmentation. The majority of these species were predators; 42% of all abundant predator species were significantly more likely to be absent from samples in forest fragments than in undisturbed forest. These figures are regarded as minimum estimates for the entire beetle assemblage because rarer species will inevitably have higher extinction probabilities. Absolute loss of biodiversity will affect ecosystem process rates, but the differential loss of species from trophic groups will have an even greater destabilizing effect on food web structure and ecosystem function.     

Ground Beetles (Coleoptera, Carabidae) in Forest Fragments of the Manawatu, New Zealand: Collapsed Assemblages?

Botanically diverse and well-maintained, protected forest fragments in the Manawatu area of the North Island of New Zealand contained very species-poor carabid assemblages. In a nearby large forest tract, the potential source area, nine species were caught in pitfall traps, while the largest forest remnant had two species, and a well-managed suburban forest patch had three species but only one with a potentially reproducing population. Lack of grazing and high botanical diversity was insufficient to maintain the potential carabid assemblage in these fragments. Predation risk and a low dispersal power in endemic New Zealand ground beetles, combined with fragment size and degree of isolation could contribute to this collapse. Active management of ground-active invertebrate species seems necessary to protect them in isolated forest fragments in New Zealand.     

Declining populations in one of the last refuges for threatened mammal species in northern Australia

Australia has contributed a disproportionate number of the world's mammal extinctions over the past 200 years, with the greatest loss of species occurring through the continent's southern and central arid regions. Many taxonomically and ecologically similar species are now undergoing widespread decline across the northern Australian mainland, possibly driven by predation by feral cats and changed fire regimes. Here, we report marked recent declines of native mammal species in one of Australia's few remaining areas that support an intact mammal assemblage, Melville Island, the largest island off the northern Australian coast. We have previously reported a marked decline on Melville Island of the threatened brush‐tailed rabbit‐rat (Conilurus penicillatus) over the period 2000–2015, linked to predation by feral cats. We now report a 62% reduction in small mammal trap‐success and a 36% reduction in site‐level species richness over this period. There was a decrease in trap‐success of 90% for the northern brown bandicoot (Isoodon macrourus), 64% for the brush‐tailed rabbit‐rat and 63% for the black‐footed tree‐rat (Mesembriomys gouldii), but no decline for the common brushtail possum (Trichosurus vulpecula). These results suggest that populations of native mammals on Melville Island are exhibiting similar patterns of decline to those recorded in Kakadu National Park two decades earlier, and across the northern Australian mainland more generally. Without the implementation of effective management actions, these species are likely to be lost from one of their last remaining strongholds, threatening to increase Australia's already disproportionate contribution to global mammal extinctions.     

Nymphalid butterfly dispersal among forest fragments at Serra da Canastra National Park,Brazil

Organisms must possess good dispersal ability to persist in fragmented landscapes, as extinction in habitat patches is frequent and patches must be re-colonised to keep viable metapopulations. Thus, metapopulation maintenance is dependent on patch size and distance, although these affect species differently. In order to evaluate the ability of Nymphalid butterfly species to live in naturally fragmented small forest fragments we marked and released 3,415 butterflies in 16 of these areas separated in two networks at the Serra da Canastra National Park (PNSC), south-eastern Brazil. Subsequent recaptures in different forest fragments enabled us to assess the dispersal rates and distances for several Nymphalid species. Seventeen butterflies from 11 out of the 50 species captured were directly observed to disperse from 500 m to 870 m. Dispersal rates varied between 1 and 7% of the marked individuals and were directly correlated to the mean forewing length of each butterfly species population. The connectivity of the forest fragments through creeks appear to facilitate butterfly dispersal among fragments within micro-basins, as only one out of 50 dispersing individuals was observed to fly from one micro-basin to the other. Several species had viable populations in the small-fragment network. The distance between fragments is crucial as the coarser fragment network was unlikely to sustain viable populations of most of the species. The protection of large forest fragments located outside of the PNSC may be necessary to promote colonization of the smaller forest fragments inside the Park.     

Terrestrial Mammal Occupancy in the Context of Widespread Forest Loss and a Proposed Interoceanic Canal in Nicaragua's Decreasingly Remote South Caribbean Region

Central America is experiencing rapid forest loss and habitat degradation both inside and outside of protected areas. Despite increasing deforestation, the Caribbean region of Nicaragua plays an important role in the survival or extinction of large mammal populations in Central America given that it still retains core areas of habitat for large mammal species. The proposed interoceanic canal project that would bisect the southern half of this Caribbean region represents a new threat that, combined with an advancing agricultural frontier, could affect populations of large mammal species such as jaguars, white-lipped peccaries, and Baird’s tapirs. We used occupancy models to examine the relative occupancy probabilities for an assemblage of terrestrial mammals in the south Caribbean region of Nicaragua to identify current core areas for our study species and conduct a preliminary evaluation of the potential impacts of the proposed interoceanic canal. We modeled a community level distribution of eight species with varying levels of sensitivity to human encroachment and a range of habitat associations. Our model results reveal three priority areas for terrestrial mammal conservation in our study area. The mapped predictions show that the only remaining area of suitable habitat for large mammals in the path of the proposed interoceanic canal is a relatively thin strip of forest that runs along the Caribbean Coast. In light of these findings, we propose five recommendations that will help ensure the conservation of this area of the proposed canal route as suitable habitat for our study species.     

Viability of ecological processes in small Afromontane forest patches in South Africa

Abstract The conservation of biodiversity is dependent on protecting ecosystem‐level processes. We investigated the effects of fragment size and habitat edge on the relative functioning of three ecological processes – decomposition, predation and regeneration of trees – in small Afromontane forests in KwaZulu‐Natal, South Africa. Ten sampling stations were placed in each of four forest categories: the interior of three large indigenous forest fragments (100 m from the edge), the edges of these large fragments, 10 small indigenous fragments (<1 ha) and 10 small exotic woodlands (<0.5 ha). Fragment size and edge effects did not affect the abundance of the amphipod Talitriator africana, a litter decomposer, and overall dung beetle abundance and species richness significantly. Bird egg predation was marginally greater at large patch edges compared with the other forest categories, while seed predation did not differ among forest categories. Tree seedling assemblage composition did not differ significantly among large patch interiors and edges, and small indigenous fragments. Sapling and canopy assemblage composition each differed significantly among these three indigenous forest categories. Thus, while tree recruitment was not negatively affected by patch size or distance from the edge, conditions in small fragments and at edges appear to affect the composition of advanced tree regeneration. These ecological processes in Afromontane forests appear to be resilient to fragmentation effects. We speculate that this is because the organisms in these forests have evolved under fragmented conditions. Repeated extreme changes in climate and vegetation over the Pleistocene have acted as significant distribution and ecological extinction filters on these southern hemisphere forest biota, resulting in fauna and flora that are potentially resilient to contemporary fragmentation effects. We argue that because small patches and habitat edges appear to be ecologically viable they should be included in future conservation decisions.     

Small and large mammals from the Ciota Ciara cave (Borgosesia,Vercelli, Italy): An Isotope Stage 5 assemblage

The Ciota Ciara cave is located within the Monte Fenera karst system (Borgosesia, Vercelli, Italy) at 670 m a.s.l. The cave entrance presents a deposit with Mousterian quartz and flint industry. The faunal remains from Stratigraphic Units 13, 14 and 103 are the subject of this work and are presented here as a whole for the first time. The large mammal assemblage is dominated by Ursus spelaeus. In addition, a few remains of carnivores such as Panthera leo spelaea, Panthera pardus, Meles meles have been found together with ungulates such as Cervus elaphus and Rupicapra rupicapra. The small mammal assemblage is characterized by a high biodiversity, especially in bats, by the dominance of Clethrionomys glareolus and by a relatively large number of Pliomys coronensis, a species that seems to disappear from the Italian Peninsula at the end of Marine Isotope Stage 5 or during the beginning of MIS 4. The changes in frequency of the small and large mammals between the two S.U. suggest a change from a relatively cold-humid (S.U. 14) to a warmer-still humid climate (S.U. 13). Although no radiometric dates are available yet, the small and large mammal assemblages, the presence of Pliomys coronensis and the climate change inferred by the variation of the small mammal frequencies allow us to correlate these two units of the Ciota Ciara cave to a relatively warm moment of Marine Isotope Stage 5.     

Using population viability analysis to predict the effect of fire on the extinction risk of an endangered shrub <Emphasis Type="Italic">Verticordia fimbrilepis</Emphasis> subsp. <Emphasis Type="Italic">fimbrilepis</Emphasis> in a fragmented landscape

The fragmentation of mediterranean climate landscapes where fire is an important landscape process may lead to unsuitable fire regimes for many species, particularly rare species that occur as small isolated populations. We investigate the influence of fire interval on the persistence of population fragments of the endangered shrub Verticordia fimbrilepis Turcz. subsp. fimbrilepis in mediterranean climate south-west of Western Australia. We studied the population biology of the species over 5 years. While the species does recruit sporadically without fire this occurs only in years with above average rainfall, so fire seems to be the main environmental factor producing extensive recruitment. Transition matrix models were constructed to describe the shrub’s population dynamics. As the species is killed by fire and relies on a seed bank stimulated to germinate by smoke, stochastic simulations to compare different fire frequencies on population viability were completed. Extinction risk increased with increasing average fire interval. Initial population size was also important, with the lowest extinction risk in the largest population. For populations in small reserves where fire is generally excluded, inevitable plant senescence will lead to local extirpation unless fires of suitable frequency can be used to stimulate regeneration. While a suitable fire regime reduces extinction risk small populations are still prone to extinction due to stochastic influences, and this will be exacerbated by a projected drying climate increasing rates of adult mortality and also seedling mortality in the post-fire environment.     

First human-caused extinction of a cetacean species?

The Yangtze River dolphin or baiji (Lipotes vexillifer), an obligate freshwater odontocete known only from the middle-lower Yangtze River system and neighbouring Qiantang River in eastern China, has long been recognized as one of the world's rarest and most threatened mammal species. The status of the baiji has not been investigated since the late 1990s, when the surviving population was estimated to be as low as 13 individuals. An intensive six-week multi-vessel visual and acoustic survey carried out in November-December 2006, covering the entire historical range of the baiji in the main Yangtze channel, failed to find any evidence that the species survives. We are forced to conclude that the baiji is now likely to be extinct, probably due to unsustainable by-catch in local fisheries. This represents the first global extinction of a large vertebrate for over 50 years, only the fourth disappearance of an entire mammal family since AD 1500, and the first cetacean species to be driven to extinction by human activity. Immediate and extreme measures may be necessary to prevent the extinction of other endangered cetaceans, including the sympatric Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis).     

Relative influence of size,connectivity and disturbance history on plant species richness and assemblages in fragmented grasslands

Questions: What is the relative influence of size, connectivity and disturbance history on plant species richness and assemblages of fragmented grasslands? What is the contribution of small fragments to the conservation of native species pool of the region? Location: Tandilia's Range, Southern Pampa, Argentina. Methods: Cover of plants was registered within 24 fragments of tall‐tussock grassland remnants within an agricultural landscape using modified Whittaker nested sampling. We analysed the influence of site variables related to disturbance history (canopy height, litter thickness) and fragment variables (size, connectivity) on species richness (asymptotic species richness, slope of the species–area curve) as well as on species assemblages by multiple regressions analysis and canonical correspondence analyses, respectively. Cumulative area was used for analysing whether small fragments or large fragments are more important to species diversity in the landscape. Results: Asymptotic species richness was significantly influenced by site variables, in particular by Paspalum quadrifarium's canopy height, but not by fragment variables. Species assemblages were also affected by site variables (12.2% of total variation), but no additional portion of the species assemblage variability was significantly explained by fragment size and connectivity. Sampling of several small fragments rendered more exotic and native species than sampling of few large fragments of the same total area. Conclusions: Our results agree with previous studies reporting low sensitivity of species diversity to size and isolation of grassland fragments in fragmented landscapes and high sensitivity of species diversity to local variables. The higher capture of regional native species pool by small grassland fragments than by few larger ones of equivalent accumulated area highlights the value of small fragments for conservation.     

Habitat patch and matrix effects on small-mammal persistence in Amazonian forest fragments

Tropical forest mammal assemblages are widely affected by the twin effects of habitat loss and habitat fragmentation. We evaluated the effects of forest patch metrics, habitat structure, age of patch isolation, and landscape metrics on the species richness, abundance and composition of small mammals at 23 forest fragments (ranging in size from 43 to 7,035 ha) in a highly deforested 3,609-km² landscape of southwestern Brazilian Amazonia. Using pitfall traps and both terrestrial and arboreal traplines of Sherman, Tomahawk and snap traps, we captured a total of 844 individuals over 34,900 trap-nights representing 26 species and 20 genera of small-mammals, including 13 rodent and 13 marsupial species. We also consider the effects of distance from forest edges on species occupancy and abundance. Overall small mammal abundance, species richness and species composition were primarily affected by the quality of the open-habitat matrix of cattle pastures, rather than by patch metrics such as fragment size. Ultimately, small mammal community structure was determined by a combination of both landscape- and patch-scale variables. Knowledge of the anthropogenic factors that govern small mammal community structure is of critical importance for managing the persistence of forest vertebrates in increasingly fragmented neotropical forest landscapes.     

Small-scale experimental habitat fragmentation reduces colonization rates in species-rich grasslands

Habitat fragmentation is one of the most important threats to biodiversity. Decreasing patch size may lead to a reduction in the size of populations and to an increased extinction risk of remnant populations. Furthermore, colonization rates may be reduced in isolated patches. To investigate the effects of isolation and patch size on extinction and colonization rates of plant species, calcareous grasslands at three sites in the Swiss Jura Mountains were experimentally fragmented into patches of 0.25, 2.25, and 20.25 m² by frequent mowing of the surrounding area from 1993 to 1999. Species richness in the fragment plots and adjacent control plots of the same sizes was recorded during these 7 years. In agreement with the theory of island biogeography, colonization rate was reduced by 30% in fragments versus non-isolated controls, and extinction increased in small versus large plots. Habitat specialists, in contrast to generalists, were less likely to invade fragments. In the last 4 years of the experiment, extinction rates tended to be higher in fragment than in control plots at two of the three sites. Despite reduced colonization rates and a tendency of increased extinction rates in fragments, fragmented plots had only marginally fewer species than control plots after 7 years. Hence, rates were a more sensitive measure for community change than changes in species richness per se. From a conservation point of view, the detected reduced colonization rates are particularly problematic in small fragments, which are more likely to suffer from high extinction rates in the long run.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

The conservation value of small, isolated fragments of lowland tropical rain forest

Deforestation is occurring at an alarming rate in the lowland tropics. In many tropical regions, rain forest is restricted to small (<100 ha), isolated fragments. While only the preservation of large areas of tropical rain forest can safeguard the complete biota, recent research has shown that a substantial number of forest species can persist for decades in fragmented forest, though large vertebrates are susceptible to habitat fragmentation. Inevitably, small fragments will become the last refuges of many rainforest species that are on the brink of extinction. In areas with little rain forest remaining, fragments can be the 'seeds' from which to re-establish extensive forest.     

Effects of microhabitat on palm seed predation in two forest fragments in southeast Brazil

The establishment of plants depends crucially on where seeds are deposited in the environment. Some authors suggest that in forest understory seed predation is lower than in gaps, and higher than at the forest edge. However, most studies have been carried out in large forest patches and very little is known about the effects of microhabitat conditions on seed predation in forest fragments. We evaluated the effects of three microhabitats (gaps, forest edge, and understory) on seed predation of two palm species (Euterpe edulis and Syagrus romanzoffiana) in two semi-deciduous forest fragments (230 and 2100 ha) in southeast Brazil. Our objective was to test two hypotheses: (1) Low rodent abundance in small fragments as a result of meso-predator action levels leads to lower seed predation in small fragments. (2) Most mammal species in small fragments are generalists with respect to diet and habitat, so that seed predation is similar in different microhabitats (gaps, forest edge and understory) in the small fragment, but not in the larger one. The study community of small fragments is usually composed of generalist species (in diet and habitat aspects), so we expected the same rate of seed predation among microhabitats (gaps, forest edge and understory) in the tested smaller fragment. The experiment was carried out in the dry season (for E. edulis) and in the wet season (for S. romanzoffiana) in 1999. We conclude that post-dispersal seed predation in forest fragments can be directly connected with mammal communities, reflecting their historical and ecological aspects.     

Robustness of a meta-network to alternative habitat loss scenarios

Studying how habitat loss affects the tolerance of ecological networks to species extinction (i.e. their robustness) is key for our understanding of the influence of human activities on natural ecosystems. With networks typically occurring as local interaction networks interconnected in space (a meta-network), we may ask how the loss of specific habitat fragments affects the overall robustness of the meta-network. To address this question, for an empirical meta-network of plants, herbivores and natural enemies we simulated the removal of habitat fragments in increasing and decreasing order of area, age and connectivity for plant extinction and the secondary extinction of herbivores, natural enemies and their interactions. Meta-network robustness was characterized as the area under the curve of remnant species or interactions at the end of a fragment removal sequence. To pinpoint the effects of fragment area, age and connectivity, respectively, we compared the observed robustness for each removal scenario against that of a random sequence. The meta-network was more robust to the loss of old (i.e. long-fragmented), large, connected fragments than of young (i.e. recently fragmented), small, isolated fragments. Thus, young, small, isolated fragments may be particularly important to the conservation of species and interactions, while contrary to our expectations larger, more connected fragments contribute little to meta-network robustness. Our findings highlight the importance of young, small, isolated fragments as sources of species and interactions unique to the regional level. These effects may largely result from an unpaid extinction debt, whereby younger fragments are likely to lose species over time. Yet, there may also be more long-lasting effects from cultivated lands (e.g. water, fertilizers and restricted cattle grazing) and network complexity in small, isolated fragments. Such fragments may sustain important biological diversity in fragmented landscapes, but maintaining their conservation value may depend on adequate restoration strategies.     

High RAPD but no cpDNA sequence variation in the endemic and endangered plant, Heptacodium miconioides Rehd. (Caprifoliaceae)

Heptacodium miconioides Rehd. is an endangered species endemic to China and has suffered rapid decrease of distribution range and population size. This species has been disappeared in central China where the modal specimen was collected. We analyzed the genetic variation of the remaining populations to reveal whether the genetic diversity also suffered decrease and to provide some suggestions for conservation. All the nine known remaining populations were sampled. Genetic variation was analyzed based on RAPD markers and two fragments of cpDNA sequence, intergenic spacers of petG-trnP and trnS-trnG. No variation was observed in the two fragments of cpDNA sequence. However, the species exhibited high level of RAPD variation compared to other threatened or rare plants. Measures of genetic diversity within populations were strongly related to the log of estimated population size, indicating that large populations usually have more genetic diversity than that of small ones. About 25% of the variation was partitioned among populations. Significant relationship was observed between differentiation and geographical distance, indicating a pattern of isolation-by-distance. Given for few populations remaining, all the populations should be protected and urgent efforts be paid on the small populations to avoid their local extinction.