Marginal Habitats Research Articles

Fungi inhabiting fine roots of Pinus heldreichii in the Montenegrin montane forests

Pinus heldreichii H. Christ. is a tertiary relict and endemic to the western Balkans and southern part of Apennine peninsula. It is an Oro-Mediterranean species occurring at altitudes between 1200 and 2000 m and primarily on calcareous soils. P. heldreichii forests are of key importance for nature conservation, protection against gravitational natural hazards, landscape conservation and recreation. However, these forests are currently highly fragmented and require the elaboration of guidelines for sustainable management and conservation that should be based on scientific knowledge. Ectomycorrhizal (ECM) fungi are important for successful regeneration, establishment and growth of P. heldreichii. The aim of this study was to investigate ECM and other fungal communities associated with fine roots of P. heldreichii at two different sites in Kuci Mountains, south-eastern Montenegro. Roots and soil were sampled from 70 trees. Soil was subjected to chemical analyses, fine roots were morphotyped and selected root morphotypes were Sanger sequenced using ITS rDNA as a marker. Sequencing resulted in 431 high-quality sequences representing 147 different fungal species including a large number of ECM species. The most common species were ECM fungi Lactarius sanguifluus (5.1%), Wilcoxina rehmii (4.2%) and Amphinema sp. KK28 (3.2%). Climatic factors were similar between the sites, but site size, inclination, elevation, tree age (old growth versus young trees), and some soil characteristics differed. The results demonstrate relatively high fungal diversity and site-specific effects on abundance and composition of fungal communities in fine roots of P. heldreichii growing in high-altitude marginal habitats.

Re‐introduction of structurally complex wood jams promotes channel and habitat recovery from overwidening: Implications for river conservation

Abstract Large wood is a powerful geomorphic agent in rivers, providing important habitat functions for a range of aquatic organisms, but has been subject to a long history of removal. Internationally, approaches to river restoration are increasingly incorporating large wood features, but generally favour simple flow deflectors (e.g. single logs, stripped of branches and anchored in place) over more complex structures that more accurately mimic natural wood jams. This paper explores channel response to wood‐based restoration of an overwidened lowland chalk stream that incorporated whole felled trees. Hydraulics, sediment, topography and vegetation data were assessed for a 3 year period for two restored reaches: an upstream reach where pre‐restoration baseline data were obtained, and a downstream reach restored before data collection. Where pre‐restoration data were available, the introduction of wood jams generated sediment deposition within jams leading to the development of vegetated marginal ‘benches’ and bed scour in adjacent areas of flow convergence. Patterns were less clear in the downstream reach, where restoration design was less ambitious and outcomes may have been affected by subsequent restoration work upstream. The results indicate that reintroduction of large wood (whole trees), can promote channel and habitat recovery from overwidening in lowland rivers, creating important ecological benefits through the provision of structurally complex marginal habitat and associated food resources. Longer‐term assessments are required to establish whether the trajectories of change are persistent. The work emphasizes the effectiveness of restoration approaches that aim to ‘work with nature’. The ambitious design, incorporating structurally complex wood jams, was also low‐cost, using materials available from the river corridor (existing riparian trees). Furthermore, ecosystem engineering effects were amplified by the colonization of wood jams by aquatic vegetation. The approach should, therefore, be transferable to other lowland rivers, subject to wider catchment constraints.

Range expansion potential of two co-occurring invasive vines to marginal habitats in Turkey

Niche distribution models accurately predict the potential distribution range of invasive plants into new habitats based on their climatic requirements in the native regions. However, these models usually ignore the marginal habitats which can limit the distribution of exotic plants. We therefore tested the seedling survival, growth and nutrient acquisition capabilities of two co-occurring invasive vines [Persicaria perfoliata (L.) H. Gross and Sicyos angulatus L.] in three different manipulative greenhouse experiments to infer their range expansion potential to marginal habitats in Turkey. First experiment included five different moisture availability regimes (100, 75, 50, 25 and 12.5% available water), second experiment consisted of four different salinity levels (0, 3, 6 and 12 dSm−1 soil salinity) and third experiment had four different soil textures (clay-1, clay-2, sandy loam and silt-clay-loam). Seedling mortality was only observed under extreme moisture deficiency in both plant species, while most of the transplanted seedlings of both species did not survive under 6 and 12 dSm−1 salinity levels. Soil textures had no effect on seedling survival. POLPE better tolerated low moisture availability and high salinity compared to SIYAN. Biomass production in both plant species was linearly reduced with increasing salinity and moisture deficiency. SIYAN invested more resources towards shoot, accumulated higher K and P, whereas POLPE maintained higher root-to-shoot ratio under all experimental conditions. Both plant species employed different strategies to cope with adverse environmental conditions, but failed to persist under high soil salinity and moisture deficiency. Our study suggest that both plant species have limited potential of range expansion to marginal habitats and will be limited to moist and humid areas only. Therefore, further research activities should be concentrated in these regions to develop effective management strategies against both species.

Suitability of Laurentian Great Lakes for invasive species based on global species distribution models and local habitat

AbstractEfficient management and prevention of species invasions requires accurate prediction of where species of concern can arrive and persist. Species distribution models provide one way to identify potentially suitable habitat by developing the relationship between climate variables and species occurrence data. However, these models when applied to freshwater invasions are complicated by two factors. The first is that the range expansions that typically occur as part of the invasion process violate standard species distribution model assumptions of data stationarity. Second, predicting potential range of freshwater aquatic species is complicated by the reliance on terrestrial climate measurements to develop occurrence relationships for species that occur in aquatic environments. To overcome these obstacles, we combined a recently developed algorithm for species distribution modeling—range bagging—with newly available aquatic habitat‐specific information from the North American Great Lakes region to predict suitable habitat for three potential invasive species: golden mussel, killer shrimp, and northern snakehead. Range bagging may more accurately predict relative suitability than other methods because it focuses on the limits of the species environmental tolerances rather than central tendency or “typical” cases. Overlaying the species distribution model output with aquatic habitat‐specific data then allowed for more specific predictions of areas with high suitability. Our results indicate there is suitable habitat for northern snakehead in the Great Lakes, particularly shallow coastal habitats in the lower four Great Lakes where literature suggests they will favor areas of wetland and submerged aquatic vegetation. These coastal areas also offer the highest suitability for golden mussel, but our models suggest they are marginal habitats. Globally, the Great Lakes provide the closest match to the currently invaded range of killer shrimp, but they appear to pose an intermediate risk to the region. Range bagging provided reliable predictions when assessed either by a standard test set or by tests for spatial transferability, with golden mussel being the most difficult to accurately predict. Our approach illustrates the strength of combining multiple sources of data, while reiterating the need for increased measurement of freshwater habitat at high spatial resolutions to improve the ability to predict potential invasive species.

Second members of the harpacticoid genera Pontopolites and Pseudoleptomesochra (Crustacea, Copepoda) are new species from Korean marine interstitial

Monotypic genera present a specific set of problems in phylogenetic analyses, and their familiar placements are often provisional. Our survey of Korean copepods in marginal habitats resulted in a discovery of two new species, both representing second members of their respective genera: Pontopolites duljjae sp. nov. and Pseudoleptomesochra mannada sp. nov. This gave us an opportunity to contribute a set of novel characters, including previously unused cuticular organs on somites, and to evaluate the most important synapomorphies in each genus. Pontopolites duljjae differs from the northern Atlantic Pontopolites typicus Scott T., 1894 in the segmentation of antennula and armature of several appendages. Especially intriguing is the sexual dimorphism in the segmentation and armature of the second to fourth leg endopods, but this is probably a plesiomorphic character state in a larger group of nannopodids. We provide an amended diagnosis of the genus Pontopolites Scott T., 1894. In contrast, morphological differences between our Psudoleptomesochra mannada and its congener from the northeast Pacific, Psudoleptomesochra typica Lang, 1965, are rather minute and contribute very little to the generic diagnosis. In both new species, cuticular organs show very little intraspecific variability and sexual dimorphism, which provides an invaluable tool for matching females and males in samples from different localities or sympatric congeners, as well as for delimitation of closely related species using cuticular organs as landmarks for geometric morphometric analyses. They can also be useful characters for reconstructing phylogenetic relationships, especially in interstitial groups with numerous appendage reductions, such as members of Nannopodidae and Ameiridae.

Dissecting a biodiversity hotspot: The importance of environmentally marginal habitats in the Atlantic Forest Domain of South America

AbstractAimWe aimed to assess the contribution of marginal habitats to the tree species richness of the Mata Atlântica (Atlantic Forest) biodiversity hotspot. In addition, we aimed to determine which environmental factors drive the occurrence and distribution of these marginal habitats.LocationThe whole extension of the South American Atlantic Forest Domain plus forest intrusions into the neighbouring Cerrado and Pampa Domains, which comprises rain forests (“core” habitat) and five marginal habitats, namely high elevation forests, rock outcrop dwarf‐forests, riverine forests, semideciduous forests and restinga (coastal white‐sand woodlands).MethodsWe compiled a dataset containing 366,875 occurrence records of 4,431 tree species from 1,753 site‐checklists, which were a priori classified into 10 main vegetation types. We then performed ordination analyses of the species‐by‐site matrix to assess the floristic consistency of this classification. In order to assess the relative contribution of environmental predictors to the community turnover, we produced models using 26 climate and substrate‐related variables as environmental predictors.ResultsOrdination diagrams supported the floristic segregation of vegetation types, with those considered as marginal habitats placed at the extremes of ordination axes. These marginal habitats are associated with the harshest extremes of five limiting factors: temperature seasonality (high elevation and subtropical riverine forests), flammability (rock outcrop dwarf‐forests), high salinity (restinga), water deficit severity (semideciduous forests) and waterlogged soils (tropical riverine forests). Importantly, 45% of all species endemic to the Atlantic Domain only occur in marginal habitats.Main conclusionsOur results showed the key role of the poorly protected marginal habitats in contributing to the high species richness of the Atlantic Domain. Various types of environmental harshness operate as environmental filters determining the distribution of the Atlantic Domain habitats. Our findings also stressed the importance of fire, a previously neglected environmental factor.

The origin, diversification and adaptation of a major mangrove clade (Rhizophoreae) revealed by whole-genome sequencing.

Mangroves invade some very marginal habitats for woody plants-at the interface between land and sea. Since mangroves anchor tropical coastal communities globally, their origin, diversification and adaptation are of scientific significance, particularly at a time of global climate change. In this study, a combination of single-molecule long reads and the more conventional short reads are generated from Rhizophora apiculata for the de novo assembly of its genome to a near chromosome level. The longest scaffold, N50 and N90 for the R. apiculata genome, are 13.3 Mb, 5.4 Mb and 1.0 Mb, respectively. Short reads for the genomes and transcriptomes of eight related species are also generated. We find that the ancestor of Rhizophoreae experienced a whole-genome duplication ~70 Myrs ago, which is followed rather quickly by colonization and species diversification. Mangroves exhibit pan-exome modifications of amino acid (AA) usage as well as unusual AA substitutions among closely related species. The usage and substitution of AAs, unique among plants surveyed, is correlated with the rapid evolution of proteins in mangroves. A small subset of these substitutions is associated with mangroves' highly specialized traits (vivipary and red bark) thought to be adaptive in the intertidal habitats. Despite the many adaptive features, mangroves are among the least genetically diverse plants, likely the result of continual habitat turnovers caused by repeated rises and falls of sea level in the geologically recent past. Mangrove genomes thus inform about their past evolutionary success as well as portend a possibly difficult future.

A groundwater-fed coastal inlet as habitat for the Caribbean queen conch Lobatus gigas—an acoustic telemetry and space use analysis

The queen conch Lobatus (Strombus) gigas, a marine snail, is among the most important fisheries resources of the Caribbean region. To provide effective protection in marine reserves, a good understanding of its habitat usage is essential. Queen conches commonly inhabit marine habitats. In this study, its activity space in a marginal estuarine-like habitat, the groundwater-fed inlet of Xel-Ha (Mexico) was determined using high-resolution acoustic telemetry (VEMCO Positioning System). Thirty-eight animals with syphonal lengths ranging from 80 to 200 mm were tagged, 1 of them with an accelerometer tag. Their trajectories were recorded for 20 mo at 5 m resolution in a closely spaced array of 12 receivers. Space–time kernel home ranges ranged from 1000 to 18500 m2 with an ontogenetically increasing trend. Juveniles, subadults and most adults displayed continuous, non-patchy home ranges consistent with the typical intensive feeding activity by this fast-growing gastropod. In some adults, Levy flight-like fragmentation of home ranges was observed that may be related to feeding range expansion or other ecological drivers such as the breeding cycle. The observed small home ranges indicate that the space use of queen conch in this estuarine-like habitat is not conditioned by food availability, and despite environmental stress due to daily low-oxygen conditions, space use is comparable to that observed in more typical marine habitats. In a marine reserve context, the groundwater-fed inlet provides adequate protection of this inshore queen conch population. Such marginal habitats may play an increasingly important role in conservation management as pressure on populations increase.

Reciprocal transplants support a plasticity-first scenario during colonisation of a large hyposaline basin by a marine macro alga

BackgroundEstablishing populations in ecologically marginal habitats may require substantial phenotypic changes that come about through phenotypic plasticity, local adaptation, or both. West-Eberhard’s “plasticity-first” model suggests that plasticity allows for rapid colonisation of a new environment, followed by directional selection that develops local adaptation. Two predictions from this model are that (i) individuals of the original population have high enough plasticity to survive and reproduce in the marginal environment, and (ii) individuals of the marginal population show evidence of local adaptation. Individuals of the macroalga Fucus vesiculosus from the North Sea colonised the hyposaline (≥2–3‰) Baltic Sea less than 8000 years ago. The colonisation involved a switch from fully sexual to facultative asexual recruitment with release of adventitious branches that grow rhizoids and attach to the substratum. To test the predictions from the plasticity-first model we reciprocally transplanted F. vesiculosus from the original population (ambient salinity 24‰) and from the marginal population inside the Baltic Sea (ambient salinity 4‰). We also transplanted individuals of the Baltic endemic sister species F. radicans from 4 to 24‰. We assessed the degree of plasticity and local adaptation in growth and reproductive traits after 6 months by comparing the performance of individuals in 4 and 24‰.ResultsBranches of all individuals survived the 6 months period in both salinities, but grew better in their native salinity. Baltic Sea individuals more frequently developed asexual traits while North Sea individuals initiated formation of receptacles for sexual reproduction.ConclusionsMarine individuals of F. vesiculosus are highly plastic with respect to salinity and North Sea populations can survive the extreme hyposaline conditions of the Baltic Sea without selective mortality. Plasticity alone would thus allow for an initial establishment of this species inside the postglacial Baltic Sea at salinities where reproduction remains functional. Since establishment, the Baltic Sea populations have evolved adaptations to extreme hyposaline waters and have in addition evolved asexual recruitment that, however, tends to impede local adaptation. Overall, our results support the “plasticity-first” model for the initial colonisation of the Baltic Sea by Fucus vesiculosus.

Competition drives trait evolution and character displacement between Mimulus species along an environmental gradient.

Closely related species may evolve to coexist stably in sympatry through niche differentiation driven by in situ competition, a process termed character displacement. Alternatively, past evolution in allopatry may have already sufficiently reduced niche overlap to permit establishment in sympatry, a process called ecological sorting. The relative importance of each process to niche differentiation is contentious even though they are not mutually exclusive and are both mediated via multivariate trait evolution. We explore how competition has impacted niche differentiation in two monkeyflowers, Mimulus alsinoides and M. guttatus, which often co-occur. Through field observations, common gardens, and competition experiments, we demonstrate that M. alsinoides is restricted to marginal habitats in sympatry and that the impacts of character displacement on niche differentiation are complex. Competition with M. guttatus alters selection gradients and has favored taller M. alsinoides with earlier seasonal flowering at low elevation and floral shape divergence at high elevation. However, no trait exhibits the pattern typically associated with character displacement, higher divergence between species in sympatry than allopatry. Thus, although character displacement was unlikely the process driving initial divergence along niche axes necessary for coexistence, we conclude that competition in sympatry has likely driven trait evolution along additional niche axes.

Habitat degradation affects the summer activity of polar bears.

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species' distributions, few studies have examined associated changes in behavior. Polar bear (Ursus maritimus) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50-75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.

Assessing common birds' ecological requirements to address nature conservation in permanent crops: Lessons from Italian vineyards

Viticulture has contributed to shaping cultural landscapes in several regions across all continents. Recent farming intensification is causing landscape homogenization and biodiversity loss in several of those areas, but knowledge about the impacts on biodiversity in vineyards is still scarce.Simplified agro-ecosystems resulting from intensification host mainly generalist and common species, which still play a key role in the regulation of ecosystems and in the provision of ecosystem services.We assessed the abundance of 11 common bird species at 47 linear transects in a vineyard-dominated landscape in Trentino (NE Italy), in both spring and winter, and analysed abundance variation in relation to three independent groups of predictors: landscape, management, and topographic-climatic variables.In the majority of species (7), abundance was primarily or considerably affected by landscape attributes. However, an additional 5 species were largely affected by management practices, often with conspicuous seasonal differences. Overall, landscape and management heterogeneity positively affected the abundance of 6 species.Vineyard cover (and in particular the new spalliera trellising system) was negatively related with the abundance of 6 species, with the strongest impacts occurring in winter. On the contrary, the cover of marginal habitats had major positive effects over 8 species.Hedgerows, tree rows, and dry stone walls, as well as traditional pergola vineyards and landscape and management heterogeneity should be conserved or restored in viticultural landscapes to promote the abundance of common bird species. This strategy would ensure the maintenance of the ecosystem services they provide, while promoting the general sustainability of the agroecosystem.

Loss of habitat and connectivity erodes species diversity, ecosystem functioning, and stability in metacommunity networks

Habitat loss fragments metacommunities, altering the movement of species between previously connected habitat patches. The consequences of habitat loss for ecosystem functioning depend, in part, on how these changes in connectivity alter the spatial insurance effects of biodiversity. Spatial insurance is the maintenance of biodiversity and stable ecosystem functioning in changing environments that occurs when species are able to move between local habitat patches in order to track conditions to which they are adapted. Spatial insurance requires a combination of species sorting dynamics, which allow species to disperse to habitats where they are productive, and mass effect dynamics, where dispersal allows species to persist in marginal habitats where environmental conditions do not support growth. Here we use a spatially explicit metacommunity model to show that the relative contribution of species sorting and mass effects to spatial insurance changes with the rate of dispersal. We then simulate different sequences of habitat loss by removing habitat patches based on their betweenness centrality (the degree to which a patch serves as a connection between other patches in the metacommunity). We demonstrate that the sequence of habitat loss has a large, non‐linear impact on diversity, ecosystem functioning and stability. Spatial insurance is lost because habitat fragmentation impedes species sorting, while promoting mass effects and dispersal limitation. We find that species sorting dynamics, and thus spatial insurance, are most robust to the removal of habitat patches with low betweenness centrality. These findings advance our understanding of how habitat connectivity facilitates the maintenance of biodiversity and ecosystem functioning, and may prove useful for the design of habitat networks.

Evidence of genotypic adaptation to the exposure to volcanic risk at the dopamine receptor DRD4 locus.

Humans have colonized and adapted to extremely diverse environments, and the genetic basis of some such adaptations, for example to high altitude, is understood. In some cases, local or regional variation in selection pressure could also cause behavioural adaptations. Numerous genes influence behaviour, such as alleles at the dopamine receptor locus D4 (DRD4), which are associated with attitude toward risk in experimental settings. We demonstrate genetic differentiation for this gene, but not for five unlinked microsatellite loci, between high- and low risk environments around Mount Merapi, an active volcano in Java, Indonesia. Using a behavioural experiment, we further show that people inhabiting the high risk environment are significantly more risk averse. We provide evidence of a genetic basis for this difference, showing that heterozygotes at the DRD4 locus are more risk averse than either homozygotes. In the high risk environment, allele frequencies are equilibrated, generating a high frequency of heterozygotes. Thus it appears that overdominance (i.e. selective advantage of heterozygotes) generates negative frequency dependent selection, favouring the rarer allele at this locus. Our results therefore provide evidence for adaptation to a marginal habitat through the selection of a neurocognitive trait with a genetic basis.

Ancient Colonization of Marginal Habitats: A Comparative Analysis of Case Studies from the Old World

The present contribution deals with the concepts of marginal habitats in selected regions of the ancient world, ranging from modern Spain to the Jordanian desert and from Turkey to the Ethiopian highlands. Central to this research is the hypothesis that the occupation of areas beyond the ‘normal’ settlement patterns corresponds to colonization processes which reflect specific social strategies and may have stimulated the development of new technological skills. A review of ‘marginality’ research in various disciplines indicates that there is no comprehensive definition of the concept, which can be approached from a multitude of perspectives and with manifold objectives. A survey of the eight case studies and two more in-depth discussions of the sites of Musawwarat (Sudan) and Ayamonte (Spain) highlight the potentials as well as the limits of the archaeological investigation into past marginalities. Patterns of spatial marginalization are the easiest to detect. The studies also show that we must not limit our analysis to the adverse factors connected to different kinds of marginalities. Instead, our analyses suggest that spatially marginal areas were deliberately chosen for settlement – an integration with core-periphery approaches may help us to understand these scenarios, which have received little attention in ‘marginality’ research in archaeology or elsewhere so far.

Small creeks in a big lagoon: The importance of marginal habitats for fish populations

Temperate transitional water systems, as in the case of the Venice Lagoon, are characterised by many different shallow-water habitats. The availability of trophic resources and the low predator pressure make salt marshes one of the most important habitats for many fish species, both resident and marine migrant, but several anthropogenic pressures, erosion and relative sea level rise in particular, are causing a significant loss of this habitat. A part from natural habitats, in many small islands of the Venice lagoon, artificial creeks of different size and morphology are present, once used in traditional aquaculture activities or built up as defence lines. Aims of this study is to analyse and compare the structure and composition of fish communities inhabiting small-sized creeks, considering both the natural and artificial ones, in order to evaluate the ecological importance of these marginal habitats for fish populations. A particular attention was given to artificial sites, assessing their ecological value as alternative refuge habitats to natural salt marsh creeks. One year samplings conducted in four sites (two natural salt marshes and two artificial creeks) allowed to describe the local fish communities, which comprised 20 species overall. The analysis of how water parameters and habitat structure influenced the fish communities showed the importance of the connection between small creeks and the open lagoon, but also the refuge function offered by confined systems. High abundances of resident fish species listed in the Annex II of the Habitat Directive were observed and juveniles of eight species of marine migrant fish were found, some of which are of economic importance. This study underlined the presence of significant densities of juvenile marine migrant and lagoon resident fish species in the two artificial habitats. Actually, species richness and density resulted to be, in some cases, higher than in natural salt marsh systems. Results of this study emphasize the ecological importance of these marginal habitats for many fish species, of both conservation and economic importance Thus a proper management and restoration strategy of these sites is needed to maintain their functionality and to buffer the disappearance of natural salt marshes.

Niche partitioning among sexual and unisexual Ambystoma salamanders

AbstractOrganisms that have ecologically similar sexual and asexual forms present an evolutionary puzzle, as theory predicts that eventually one form should eliminate the other. However, both forms may persist if there is niche partitioning between them. Geographical parthenogenesis is a hypothesis that predicts that in terms of niche use the asexual form in such pairs should be more ecologically successful in marginal habitats. This model has rarely been considered for gynogenetic taxa, as they are sperm dependent and thus constrained to the geographical range of sexual “sperm donor” species. Unisexual Ambystoma salamanders present a unique opportunity to look for evidence of niche partitioning and geographical parthenogenesis in a gynogenetic lineage, as there are multiple possible sperm donor species and so the geographical distributions of unisexuals are less constrained. We used broad sampling, comparative ecological niche models, and a model selection approach to determine (1) whether niche partitioning occurs among sexual and unisexual salamanders and (2) whether unisexual lineages indeed occupy marginal habitat relative to their most similar sexual species. Nearly all unisexual and sexual types showed significant niche differentiation. Predictions consistent with geographical parthenogenesis were upheld for one sexual–unisexual pair (A. jeffersonianum and LJJ unisexuals), but not for a second pair (A. texanum and LTT unisexuals). Our study provides evidence that different biotypes within the unisexual lineage have distinct ecological interactions with sexual taxa, supporting a role for these differences as a mechanism promoting coexistence between some sexual and unisexual forms. However, geographical parthenogenesis is only a partial explanation for sexual–unisexual coexistence, suggesting that other ecological and genetic mechanisms also play important roles in mediating coexistence among these salamanders.

Recognition and management of ecological refugees: A case study of the Cape mountain zebra

Anthropogenic activities have led to long-term range contraction in many species, creating isolated populations in ecologically marginal and suboptimal habitats. ‘Refugee’ species have a current distribution completely restricted to suboptimal habitat. However, it is likely that many species are partial refugees, where one or more populations are managed in ecologically unsuitable habitat. Here, we develop a framework to assess potential refugee populations in marginal habitats using a model species: the Cape mountain zebra. We assessed habitat quality by the abundance and palatability of grass and diet quality using proximate nutrient and element analysis. High grass abundance was associated with higher population growth rates and zebra density and less skewed adult sex ratios. Furthermore, faecal nutrient and dietary element quality was also positively associated with grass abundance. Our results show that poorly performing populations were characterised by suboptimal habitat, supporting the hypothesis that the Cape mountain zebra has refugee populations. In addition, we found more variance in sex ratio and population growth rates in smaller populations suggesting they may be more at risk for random stochastic effects, such as a biased sex ratio, compounding poor performance. We show how the ‘refugee’ concept can be applied more generally when managing species with fragmented populations occurring across marginal habitats. More broadly, the results presented herein highlight the importance of recognizing the range of habitats historically occupied by a species when assessing ecological suitability. Identifying and mitigating against refugee, relict and gap populations is especially critical in the face of on-going environmental change.

Demographic and reproductive plasticity across the depth distribution of a coral reef fish.

As humans expand into natural environments, populations of wild organisms may become relegated to marginal habitats at the boundaries of their distributions. In the ocean, mesophotic coral ecosystems (30–150 m) at the depth limit of photosynthetic reefs are hypothesized to act as refuges that are buffered from anthropogenic and natural disturbances, yet the viability and persistence of subpopulations in these peripheral habitats remains poorly understood. To assess the potential for mesophotic reefs to support robust coral reef fish populations, we compared population density and structure, growth, size, and reproductive output of the bicolor damselfish (Stegastes partitus) from shallow (<10 m), deep shelf (20–30 m), and mesophotic reefs (60–70 m) across the Florida Platform. Population densities decreased and size and age distributions shifted toward older and larger individuals in deeper habitats. Otolith-derived ages indicated that S. partitus found on mesophotic reefs reach larger asymptotic sizes and have longer lifespans than fish in shallower habitats. Based on measurements of oocyte area and batch fecundity, mesophotic fish also have higher reproductive investment. These demographic patterns indicate that mesophotic fish populations composed of large, fecund individuals produce high condition larvae and rely on longevity of individuals for population persistence and viability.

Effects of traditional coppice practices and microsite conditions on tree health in a European beech forest at its southernmost range

Abstract: European beech (Fagus sylvatica) grows at the southern limit of its range in the mountain-Mediterranean vegetation belt up to the timberline. The southernmost beech forests of Sicily (southern Italy) show peculiar ecological, structural and silvicultural characteristics, growing in fragmented and isolated stands near the timberline and in topographically marginal unfavorable habitats. Past silvicultural practices increased the heterogeneity of stand structure at these sites. We compared stand structural characteristics and tree health in coppice-cut and control beech stands with respect to the local topographic gradient (bottom, slope and ridge) and canopy cover (clearing/border vs. interior trees). Our results clearly showed a correlation between declining tree health (crown and bark damage, higher percentage of dead trees and lower seedling density) and recent coppice-cuts, poor (marginal) site quality (on ridges and slopes) and reduced canopy cover (in clearing/border trees). The decrease of tree health indicate an increasing threat to the long-term viability of beech stands facing multiple environmental stress factors (such as those related to southern latitude and topographic position). Declining tree health in the control plots also supports this hypothesis. We concluded that traditional forest management practices, such as coppice-cuts applied regardless to the specific microenvironmental conditions, may pose a risk to beech forest health at the southernmost edge of the species’ range.