Marginal Habitats Research Articles

Ecologically differentiated stress-tolerant endosymbionts in the dinoflagellate genus Symbiodinium (Dinophyceae) Clade D are different species

:We used an integrative genetics approach using sequences of (1) nuclear ribosomal rDNA (internal transcribed spacers and partial large subunit rDNA), (2) single-copy microsatellite nuclear DNA, (3) chloroplast-encoded 23S rDNA, (4) mitochondrial cytochrome b, and (5) repeat variation at eight microsatellite markers, to test the hypothesis that the stress-tolerant, ‘morphologically cryptic’ Clade D Symbiodinium (Dinophyceae) was composed of more than one species. Concordant phylogenetic and population genetic evidence clearly differentiate separately evolving, reproductively isolated lineages. We describe Symbiodinium boreum sp. nov. and S. eurythalpos sp. nov., two symbionts known to occur in colonies of the zebra coral, Oulastrea crispata (Scleractinia), which lives in turbid, marginal habitats extending from equatorial Southeast Asia to the main islands of Japan in the temperate northwest Pacific Ocean. Symbiodinium boreum was associated with O. crispata in temperate latitudes and S. eurythalpos was common to colonies in the tropics. The geographical ranges of both symbiont species overlapped in the subtropics where they sometimes co-occurred in the same host colony. Symbiodinium trenchii sp. nov. is also described. As a host-generalist symbiont, it often occurs in symbiosis with various species of Scleractinia possessing open (horizontal) modes of symbiont acquisition and is common to reef coral communities thriving in warm turbid reef habitats in the western Pacific Ocean, Indian Ocean, Arabian/Persian Gulf, Red Sea and western Atlantic (Caribbean). As is typical for dinoflagellates, S. boreum and S. eurythalpos were haploid, but microsatellite loci from field-collected and cultured S. trenchii often possessed two alleles, implying that a genome-wide duplication occurred during the evolution of this species. The recognition that Clade D Symbiodinium contains species exhibiting marked differences in host specificity and geographical distribution will yield greater scientific clarity about how stress-tolerant symbionts function in the ecological response of coral–dinoflagellate symbioses to global climate change.

Self-compatibility and floral traits adapted for self-pollination allow homostylous Nymphoides geminata (Menyanthaceae) to persist in marginal habitats

Many stylar polymorphic plants are equipped with reciprocal herkogamy and incompatibility systems so that each floral morph within a population can only mate with the compatible reciprocal morph. In the ecological or geographical margins of the range of stylar polymorphic taxa, homostyly may evolve in response to a lack of mate and/or poor pollinator visitation. This prediction is tested here by comparing floral traits, seed production without pollinators (i.e. autonomously), pollinator activity and ploidy level between the homostylous Nymphoides geminata that colonise geographically marginal habitats and its stylar polymorphic and self-incompatible closely related species N. montana that typically occur in natural water bodies in south-eastern Australia. Autonomous self-fertilisation is detected in glasshouse. Homostyly in N. geminata is also associated with floral adaptation to autonomous self-pollen deposition. There is a reduction in size and attractiveness of homostylous floral organs: over 80 % shorter stigma–anther separations and lower pollen:ovule ratio than the polymorphic flowers. The SEM images reveal that homostylous flowers possess similar pollen and stigma papillae morphology to those of short styles and long styles of the polymorphic flowers, respectively. Nymphoides geminata may be an example of recombinant homostyly: combining the pollen compatibility features of one morph with the stylar compatibility features of the reciprocal morph. Results of floral visitor observations and open pollinations indicate the ability of homostylous plants to reproduce successfully despite pollinator scarcity. Somatic chromosome counts indicate a possible lack of relationship between ploidy level and sexual system. Homostylous condition should provide an opportunity for reproductive assurance in environments with unreliable pollinators.

Population and habitat assessment of the Critically Endangered hirola Beatragus hunteri in Tsavo East National Park, Kenya

AbstractThe Critically Endangered hirola Beatragus hunteri exists in two populations, a natural population on the Kenyan–Somali border and a translocated population in Tsavo East National Park, Kenya. The Tsavo population is becoming increasingly important for the survival of the hirola yet its status is unknown and little is understood about the factors that limit its growth. Here we report the size, distribution and demographic parameters of the Tsavo population of hirola and consider whether insufficient suitable habitat could be limiting its growth. The Tsavo population has not increased since 2000, when the last census was carried out, but neither has it significantly decreased. The importance of habitat as a limiting factor is dependent on the hirola's ability to utilize marginal habitat.

Reduced parasite diversity and abundance in a marine whelk in its expanded geographical range

AbstractAimTo improve our understanding of how parasitism interacts with geographical range expansions by quantifying diversity and abundance of parasites in 25 populations of a large marine snail, Kellet's whelk (Kelletia kelletii), throughout its historical and recently expanded range, which are separated by a well‐known biogeographical boundary.LocationCalifornia coast (western North America).MethodsParasitological examinations were conducted on 199 whelks from 25 subtidal reefs throughout its expanded and historical ranges. We calculated infection risk, parasite intensity, and parasite species diversity. Abiotic (temperature, latitude, distance from range limit) and biotic (host density) variables were analysed as potential drivers of differential parasitism between expanded‐ and historical‐range populations.ResultsCompared with historical‐range whelks, expanded‐range whelks were 20% as likely to be infected by parasites, and those that were infected had 6% the number of individual parasites. On average, expanded‐range whelks had 14% the number of parasite species than the historical‐range whelks. The marked decrease in species richness of parasites infecting expanded‐range whelks was only partly explained by the low numbers of parasites. The reduced parasite abundance and diversity in the expanded‐range whelks was not explained by the examined abiotic factors or by whelk density.Main conclusionsExpanded‐range populations of Kellet's whelk experience substantially lower parasite abundance and diversity than the historical‐range populations, despite relatively poor demographic performance. The reduced parasitism observed resembles the enemy escape typically characterizing invasive species. A possible explanation for the observed ‘parasite escape’ is that the biogeographical boundary limits the movements or drives the low abundance of other host species (elasmobranchs) required to complete the life cycles of the ‘missing’ parasites. We suggest that parasite escape may generally characterize range‐margin expansions and be important in permitting expansions into what may otherwise be marginal habitats. This parasite escape may directly counter the spread of infectious diseases associated with global warming‐induced range shifts.

Livestock and food security: vulnerability to population growth and climate change

Livestock production is an important contributor to sustainable food security for many nations, particularly in low-income areas and marginal habitats that are unsuitable for crop production. Animal products account for approximately one-third of global human protein consumption. Here, a range of indicators, derived from FAOSTAT and World Bank statistics, are used to model the relative vulnerability of nations at the global scale to predicted climate and population changes, which are likely to impact on their use of grazing livestock for food. Vulnerability analysis has been widely used in global change science to predict impacts on food security and famine. It is a tool that is useful to inform policy decision making and direct the targeting of interventions. The model developed shows that nations within sub-Saharan Africa, particularly in the Sahel region, and some Asian nations are likely to be the most vulnerable. Livestock-based food security is already compromised in many areas on these continents and suffers constraints from current climate in addition to the lack of economic and technical support allowing mitigation of predicted climate change impacts. Governance is shown to be a highly influential factor and, paradoxically, it is suggested that current self-sufficiency may increase future potential vulnerability because trade networks are poorly developed. This may be relieved through freer trade of food products, which is also associated with improved governance. Policy decisions, support and interventions will need to be targeted at the most vulnerable nations, but given the strong influence of governance, to be effective, any implementation will require considerable care in the management of underlying structural reform.

A Comparison of Bromus tectorum Growth and Mycorrhizal Colonization in Salt Desert vs. Sagebrush Habitats

Cheatgrass (Bromus tectorum) has recently invaded marginal low-elevation salt desert habitats across the Great Basin. We tested the hypothesis that cheatgrass seed produced in populations from the more stressful salt desert vs. upland sagebrush habitats should grow differently in salt desert soils compared to adjacent upland sagebrush soil, and vice versa. We evaluated growth, incidence of flowering, and arbuscular mycorrhizal fungi (AMF) colonization of plants grown in the soils from which their seeds were collected vs. in the reciprocal soils from the nearest sagebrush or salt desert site in three large basins in northern Nevada. Simultaneously we measured nutrient cations, available nitrogen and phosphorus, percent carbon and nitrogen, texture, and dry-down characteristics in all soils. We found that salt desert soils were generally more nutrient poor and more saline than their upland (sagebrush) counterparts; salt desert soils also generally had a higher percentage of sand compared to their upland counterparts and were consistently drier. The most dramatic plant responses to soil and seed source were 1) lower aboveground biomass of mature plants in most salt desert soils compared to sagebrush soils, or lower biomass in plants grown from salt desert seed; 2) lower root:shoot ratios in plants grown in salt desert soil across two of three basins, irrespective of seed source; 3) a higher percentage of flowering individuals from salt desert seed sources at harvest, irrespective of soil source; 4) depressed AMF colonization of plants in salt desert soils; and 5) strong influence exerted by seed source on AMF, whereby sagebrush-originating plants grown in sagebrush soils had greater AMF colonization compared to salt desert soils but salt desert-originating seedlings had very low AMF colonization rates irrespective of soil source. These results suggest that both population level and soil-based controls are important as this widespread weed moves into marginal habitat.

La diversidad floral del borde afecta la riqueza y abundancia de visitantes florales nativos en cultivos de girasol

Ecología Austral, 24:94-102 (2014)Pollinator decline has been related to different types of anthropogenic disturbances, a topic largely debated in the context of global change. Within this debate, it has been stressed the importance of natural or semi-natural habitat remnants as pollinator sources for nearby agricultural fields. The SW of Buenos Aires Province is a highly-impacted and intensively-used agricultural region. However, field margins associated with country roads can provide diverse floristic resources and nesting sites for a variety of pollinators, which can eventually improve pollination and productivity of neighboring pollinator-dependent crops. Here, we evaluated the effect of floral diversity of field margins on the richness and abundance of floral visitors across 17 sunflower lots in the districts of Balcarce and Lobería. Our results indicate that floral diversity of field margins had a positive effect on the richness and abundance of native floral visitors in the adjacent crop field. As a consequence, pollination services in these agroecosystems would be promoted by increasing the amount and diversity of these marginal habitats.

La diversidad floral del borde afecta la riqueza y abundancia de visitantes florales nativos en cultivos de girasol

Ecología Austral, 24:94-102 (2014)Pollinator decline has been related to different types of anthropogenic disturbances, a topic largely debated in the context of global change. Within this debate, it has been stressed the importance of natural or semi-natural habitat remnants as pollinator sources for nearby agricultural fields. The SW of Buenos Aires Province is a highly-impacted and intensively-used agricultural region. However, field margins associated with country roads can provide diverse floristic resources and nesting sites for a variety of pollinators, which can eventually improve pollination and productivity of neighboring pollinator-dependent crops. Here, we evaluated the effect of floral diversity of field margins on the richness and abundance of floral visitors across 17 sunflower lots in the districts of Balcarce and Lobería. Our results indicate that floral diversity of field margins had a positive effect on the richness and abundance of native floral visitors in the adjacent crop field. As a consequence, pollination services in these agroecosystems would be promoted by increasing the amount and diversity of these marginal habitats.

Adaptation in a Peripheral Habitat: Intraspecific Variation in Osmoregulatory Traits in an Estuarine Hermit Crab (Pagurus minutus)

We investigated osmoregulatory ability of Pagurus minutus from three sites on the Waka River estuary (upper reach, river mouth, and middle). The sites have different salinity regimes and we tested whether crabs were adapted to their site locally or whether they showed phenotypic plasticity. The upper reach had a low salinity (∼16) during low tide, unlike the other two sites (∼28–31). The exposure of crabs from each site to seven different salinity levels (0–52.5) showed that P. minutus is a hyper-hypo-osmoregulator at all sites and is able to survive in 8.75–52.5 salinity levels for 24 h. Crabs from the upper reach, however, maintained greater hemolymph osmolarity in diluted media, and most of them were able to survive in even 0 salinity medium for 24 h. Crabs from the other two sites were not able to tolerate this condition and all died. Another experiment, involving a 1-wk acclimation to 5 decreasing salinity media (35 to 0), showed that the survival was about 1.3 times greater for crabs from the upper reach than from the other two sites. Because the crab density of the upper reach was significantly less than that of the other two sites, the upper reach is a marginal habitat established by asymmetrical larval dispersal from central saline habitats. Nevertheless, no crab from the saline sites survived exposure to the salinity of zero medium. These results suggest that the high osmoregulatory ability of crabs from the upper reach of the estuary does not represent a local adaptation; crabs acquire it after dispersal through developmental plasticity.

Testing Aleppo pine seed sources response to climate change by using trial sites reflecting future conditions

Large-scale biogeographical shifts in forest tree distributions are predicted in response to the altered precipitation and temperature regimes associated with climate change. Adaptive forest management to climate change experienced in either stable or rapidly changing environments must consider this fact when carrying out reforestation programs or specifically assisted population migration for conservation purposes. The aim of this study was to compare field performance of eleven seed sources of Aleppo pine outplanted in core and marginal habitats and to assess their phenotypic plasticity for further screening under specific conditions in particular reforestation areas. We hypothesize that current marginal habitat due to low temperature is shifting toward conditions found on the core habitat and that current core habitat will shift toward warmer and drier marginal habitat. Our study reproduced real conditions of reforestation in potential future climatic conditions. Results suggest that it is difficult to predict Aleppo pine provenances’ performance in different natural sites from their performance at a single location, even though ‘Levante interior’ and ‘La Mancha’ seed sources showed the best overall response among sites. On a site basis, provenances were matched in groups according to their survival and growth responses. Seedlings grown from local seed sources or seed orchards performed better on the core habitat. However, as conditions shifted to marginal habitats, seedlings from climatically similar regions performed better than local sources at least in the short term; our findings suggest that new plantations in areas already affected by global change could be better adapted if they use alternative seed sources.

Ontogenetic shifts in plant–plant interactions in a rare cycad within angiosperm communities

Gymnosperms and angiosperms can co-occur within the same habitats but key plant traits are thought to give angiosperms an evolutionary competitive advantage in many ecological settings. We studied ontogenetic changes in competitive and facilitative interactions between a rare gymnosperm (Dioon sonorense, our target species) and different plant and abiotic neighbours (conspecific-cycads, heterospecific-angiosperms, or abiotic-rocks) from 2007 to 2010 in an arid environment of northwestern Mexico. We monitored survival and growth of seedlings, juveniles, and adults of the cycad Dioon sonorense to evaluate how cycad survival and relative height growth rate (RHGR) responded to intra- and interspecific competition, canopy openness, and nearest neighbour. We tested spatial associations among D. sonorense life stages and angiosperm species and measured ontogenetic shifts in cycad shade tolerance. Canopy openness decreased cycad survival while intraspecific competition decreased survival and RHGR during early ontogeny. Seedling survival was higher in association with rocks and heterospecific neighbours where intraspecific competition was lower. Shade tolerance decreased with cycad ontogeny reflecting the spatial association of advanced stages with more open canopies. Interspecific facilitation during early ontogeny of our target species may promote its persistence in spite of increasing interspecific competition in later stages. We provide empirical support to the long-standing assumption that marginal rocky habitats serve as refugia from angiosperm competition for slow-growing gymnosperms such as cycads. The lack of knowledge of plant-plant interactions in rare or endangered species may hinder developing efficient conservation strategies (e.g. managing for sustained canopy cover), especially under the ongoing land use and climatic changes.

Unrecognized coral species diversity masks differences in functional ecology

Porites corals are foundation species on Pacific reefs but a confused taxonomy hinders understanding of their ecosystem function and responses to climate change. Here, we show that what has been considered a single species in the eastern tropical Pacific, Porites lobata, includes a morphologically similar yet ecologically distinct species, Porites evermanni. While P. lobata reproduces mainly sexually, P. evermanni dominates in areas where triggerfish prey on bioeroding mussels living within the coral skeleton, thereby generating asexual coral fragments. These fragments proliferate in marginal habitat not colonized by P. lobata. The two Porites species also show a differential bleaching response despite hosting the same dominant symbiont subclade. Thus, hidden diversity within these reef-builders has until now obscured differences in trophic interactions, reproductive dynamics and bleaching susceptibility, indicative of differential responses when confronted with future climate change.

Thermal Tolerance Limits of Diamondback Moth in Ramping and Plunging Assays

Thermal sensitivity is a crucial determinant of insect abundance and distribution. The way it is measured can have a critical influence on the conclusions made. Diamondback moth (DBM), Plutella xylostella (L.) (Lepidoptera: Plutellidae) is an important insect pest of cruciferous crops around the world and the thermal responses of polyphagous species are critical to understand the influences of a rapidly changing climate on their distribution and abundance. Experiments were carried out to the lethal temperature limits (ULT0 and LLT0: temperatures where there is no survival) as well as Upper and Lower Lethal Temperature (ULT25 and LLT25) (temperature where 25% DBM survived) of lab-reared adult DBM population to extreme temperatures attained by either two-way ramping (ramping temperatures from baseline to LT25 and ramping back again) or sudden plunging method. In this study the ULT0 for DBM was recorded as 42.6°C and LLT0 was recorded as −16.5°C. DBM had an ULT25 of 41.8°C and LLT25 of −15.2°C. The duration of exposure to extreme temperatures had significant impacts on survival of DBM, with extreme temperatures and/or longer durations contributing to higher lethality. Comparing the two-way ramping temperature treatment to that of direct plunging temperature treatment, our study clearly demonstrated that DBM was more tolerant to temperature in the two-way ramping assay than that of the plunging assay for cold temperatures, but at warmer temperatures survival exhibited no differences between ramping and plunging. These results suggest that DBM will not be put under physiological stress from a rapidly changing climate, rather access to host plants in marginal habitats has enabled them to expand their distribution. Two-way temperature ramping enhances survival of DBM at cold temperatures, and this needs to be examined across a range of taxa and life stages to determine if enhanced survival is widespread incorporating a ramping recovery method.

Lack of Evidence for Energetic Costs of Mate-Guarding in Wild Male Assamese Macaques (Macaca assamensis)

Mate-guarding is a widespread and efficient male strategy for increasing paternity success. The inability to guard entire female receptive phases or complete lack of mate-guarding has been explained by energetic constraints posed on males. The energetic costs per unit time a male can afford to suffer are thought to be lowest in year-round breeding species in marginal habitats and highest in seasonally breeding species in rich habitats. Here we test the prediction that mate-guarding is energetically costly in seasonal breeders in marginal habitats. We observed all males in one group of wild Assamese macaques (Macaca assamensis) for two 4-mo mating seasons and recorded activity and travel paths via focal animal sampling, physical condition via visual inspection, and collected feces for analysis of glucocorticoid levels. Generalized linear mixed model (GLMM) analyses did not reveal an effect of mate-guarding on time spent feeding or moving or on distance traveled, travel speed, directedness of travel, or glucocorticoid levels. This lack of mate-guarding costs was consistent with the lack of a relationship between individual time spent mate-guarding and change in physical condition over the mating season. Within the limitations of our study the results do not support the idea that seasonally breeding males in marginal habitats face energetic costs of mate-guarding. With others we suggest instead that though these costs may be found in species where strength, size, and mass predict rank, mating, and reproductive success, males may not engage in costly mate-guarding in species such as Assamese macaques where also smaller, weaker males may attain high social status via political coalitions or other routes alternative to direct contest over guarded females.

Marginal habitats as important refugia for riparian birds during flood years

Capsule During flood years (2003, and 2010, during a study 2003-2011) riparian birds nested more numerously on arable land than in their preferred habitats of the meadows and gravel pits in the Lower Odra Valley, NW Poland. The study showed the importance of even marginal habitats during stochastic disturbances and that they should be taken into consideration in conservation management.

Small islands, valuable insights: systems of customary resource use and resilience to climate change in the Pacific

Understanding how social-ecological systems are and can be resilient to climate change is one of the world's most crucial problems today. It requires knowledge at local and global scales, the integration of natural and social sciences, and a focus on biocultural diversity. Small Pacific Islands and the knowledge-practice-belief systems of their peoples have a long history of resilience to environmental variability and unpredictability, including in areas with marginal habitats and with periodic, severe disturbance (e.g., drought, flood, storms, and tsunami). We review the state of research on these knowledge systems as it pertains to resilience and adaptation, and we highlight critical research needs to address the interrelated areas of: (1) local-scale expertise and observations of change with regard to weather, life-history cycles, and ecological processes; (2) customary resource management institutions and practices (i.e., with agroforests and the nearshore marine environment); and (3) the roles of leaders, social institutions, and social networks in the context of disturbance and change. We conclude that these knowledge systems can contribute high-resolution observations, benchmark data, and insights into practices that enhance resilience and adaptive capacity in integrated terrestrial and marine systems. Community-based and participatory approaches can complement and ground-truth climate models and direct culturally appropriate resource management, research, and adaptation measures. Although most islands in the Pacific are small, their knowledge systems include valuable insights on seasonal cycles, ecological processes, and the management of biocultural diversity that are relevant at a broad scale for understanding resilience and adaptability to the social-ecological effects of climate change.

Habitat Suitability Modeling of Asian-Moon Scallop (<i>Amusium pleuronectes</i>) in Brebes District Waters, Central Java, Indonesia

The purpose of this study was to analyze the characteristics of the environment that determines the bottom habitat of the scallop from Brebes District and to map the potential habitat of scallop using spatial modeling. Research areas include the waters covering 806.03 km 2 of Brebes District extending 12 miles outward the sea. Habitat suitability mapping was conducted using Ecological Niche Factor Analysis (ENFA). Kolmogorov-Smirnov test was used to test differences in each environment occupied by scallop with the overall study area. ENFA results showed that the scallop required habitats with very different conditions from the mean habitat conditions existing in all study areas (p 5 km). In addition, the scallop also required bottom waters habitat with current velocity and high salinity (> 0.06 m/sec; 31.85-32.65 ‰) and low temperature of the bottom waters (29.92-30.06 °C), while towards the pH level of the bottom waters, scallop were relatively more tolerant (7.3-7.4). Based on spatial modeling of habitat suitability in the study area, there were three categories obtained: 1) “suitable habitat” area of ​​ 4,629.01 ha (5.8%), 2) “marginal habitat” area of 9,291.09 ha (11.5%), and 3) “unsuitable habitat” for scallops covering an area of ​​ 66,682.68 ha (82.7%).

Hunter–gatherers have less famine than agriculturalists

The idea that hunter-gatherer societies experience more frequent famine than societies with other modes of subsistence is pervasive in the literature on human evolution. This idea underpins, for example, the 'thrifty genotype hypothesis'. This hypothesis proposes that our hunter-gatherer ancestors were adapted to frequent famines, and that these once adaptive 'thrifty genotypes' are now responsible for the current obesity epidemic. The suggestion that hunter-gatherers are more prone to famine also underlies the widespread assumption that these societies live in marginal habitats. Despite the ubiquity of references to 'feast and famine' in the literature describing our hunter-gatherer ancestors, it has rarely been tested whether hunter-gatherers suffer from more famine than other societies. Here, we analyse famine frequency and severity in a large cross-cultural database, in order to explore relationships between subsistence and famine risk. This is the first study to report that, if we control for habitat quality, hunter-gatherers actually had significantly less--not more--famine than other subsistence modes. This finding challenges some of the assumptions underlying for models of the evolution of the human diet, as well as our understanding of the recent epidemic of obesity and type 2 diabetes mellitus.

Model Distribution of Silver Chub (Macrhybopsis storeriana) in Western Lake Erie

Silver Chub (Macrhybopsis storeriana) was once a common forage fish in Lake Erie but has declined greatly since the 1950s. Identification of optimal and marginal habitats would help conserve and manage this species. We developed neural networks to use broad-scale habitat variables to predict abundance classes of Silver Chub in western Lake Erie, where its largest remaining population exists. Model performance was good, particularly for predicting locations of habitat with the potential to support the highest and lowest abundances of this species. Highest abundances are expected in waters >5 m deep; water depth and distance to coastal habitats were important model features. These models provide initial tools to help conserve this species, but their resolution can be improved with additional data and consideration of other ecological factors.

Habitat differences influence genetic impacts of human land use on the American beech (Fagus grandifolia).

Natural reforestation after regional forest clearance is a globally common land-use sequence. The genetic recovery of tree populations in these recolonized forests may depend on the biogeographic setting of the landscape, for instance whether they are in the core or in the marginal part of the species' range. Using data from 501 individuals genotyped across 7 microsatellites, we investigated whether regional differences in habitat quality affected the recovery of genetic variation in a wind-pollinated tree species, American beech (Fagus grandifolia) in Massachusetts. We compared populations in forests that were recolonized following agricultural abandonment to those in remnant forests that have only been logged in both central inland and marginal coastal regions. Across all populations in our entire study region, recolonized forests showed limited reduction of genetic diversity as only observed heterozygosity was significantly reduced in these forests (H(O) = 0.520 and 0.590, respectively). Within inland region, this pattern was observed, whereas in the coast, recolonized populations exhibited no reduction in all genetic diversity estimates. However, genetic differentiation among recolonized populations in marginal coastal habitat increased (F(st) logged = 0.072; F(st) secondary = 0.249), with populations showing strong genetic structure, in contrast to inland region. These results indicate that the magnitude of recovery of genetic variation in recolonized populations can vary at different habitats.