Bryophytes in a changing landscape: The hierarchical effects of habitat fragmentation on ecological and evolutionary processes

Testing the myriad predictions associated with the community, demographic and genetic impacts of habitat fragmentation remains a high conservation priority. Many bryophyte taxa are ideal model systems for experimentally testing such metapopulation-based and population genetic predictions due to their relatively fast colonisation-extinction rates, high substrate specificity, dominant haploid condition, and diminutive size. Herein, we review the community, demographic and population genetic impacts of habitat fragmentation on bryophytes, highlight the present knowledge gaps, and offer ideas on how experimental studies utilizing bryophytes may be used to address the broader conservation implications associated with fragmented ecosystems. Previous research suggests that dispersal limitation best explains observed patterns of abundance and distribution of bryophytes in some fragmented habitats. However, edge effects influence bryophyte community structure of border habitats especially where abrupt differences in micro-climatic conditions between the matrix and the forest remnant exist, or where the species pool contains members with inherently restricted ecological amplitudes. Existing studies do not agree on the relationship between basic attributes of bryophyte community structure (i.e., species richness and local density), and habitat area and degree of spatial-isolation. Demographic studies are a critical step in structuring conservation strategies, however surprisingly little empirical information exists as to the impacts of habitat fragmentation on plant population dynamics. We propose that bryophytes offer great potential for testing predictions with respect to plant population persistence in spatially-structured landscapes.     

Microsatellite analysis of genetic population structure of the endangered cyprinid Anaecypris hispanica in Portugal: implications for conservation

The endangered fish species Anaecypris hispanica is restricted to eight disjunct populations in the Portuguese Guadiana drainage. The genetic structure of these populations was studied in order to determine levels of genetic variation within and among populations and suggest implications for conservation of the species. Based on five microsatellite loci, the null hypothesis of population homogeneity was tested. Tests for genetic differentiation revealed highly significant differences for pairwise comparisons between all populations, and substantial overall population subdivision (F_ST=0.112). All sampled populations contained unique alleles. Our findings indicate marked genetic structuring and emphasise limited dispersal ability. The high levels of genetic diversity detected within and among A. hispanica populations suggest, however, that the observed fragmentation and reduction in population size of some populations during the last two decades, has impacted little on levels of genetic variability. Data imply that most A. hispanica populations should be managed as distinct units and that each has a high conservation value containing unique genetic variation. It is argued that geographic patterns of genetic structuring indicate the existence of eight management units.     

Reduced dispersal of native plant species as a consequence of the reduced abundance of frugivore species in fragmented rainforest

Frugivorous animals disperse the seeds of the majority of rainforest plant species and hence play a key role in the trajectory of rainforest regeneration. This study investigated whether changes in the species composition of the frugivore community in fragmented rainforest in subtropical Australia is likely to impact the dispersal of native plant species. The potential of frugivorous bird and bat species to disperse the seeds of plant species in fragmented rainforest was assessed using published dietary information together with field surveys of frugivore abundance within intact forest, forest fragments and patches of regrowth. Frugivore species with reduced abundance in fragmented rainforest were the only known dispersers of 27 of the 221 native plant species in the data set (12% of species). These frugivore species were also major dispersers of plant species producing fruits wider than 10 mm and species from the families Rubiaceae, Lauraceae, Myrtaceae, Meliaceae, Lamiaceae and Vitaceae. Except for Rubiaceae, these plant taxa are also potentially dispersed by two of the frugivore species that were widespread in fragmented rainforest, Lopholaimus antarcticus and Ptilonorhynchus violaceus, although dispersal rates are likely to be lower in fragmented than in extensive rainforest. Consistent with other regions, large-seeded plants are susceptible to reduced dispersal in fragmented rainforest in subtropical Australia. However, we predict a smaller deficit in seed dispersal in fragmented forests than has been reported from other regions, due to factors such as functional overlap among frugivore species, the ability of many Australian rainforest vertebrates to persist in fragmented rainforest, and a lack of hunting in these forests. Nevertheless, rainforest fragmentation has reduced the abundance of a suite of frugivorous rainforest fauna, which in turn is likely to reduce the dispersal of a certain plant taxa and may alter patterns of plant regeneration in subtropical Australian rainforest fragments.     

Persistence and conservation of a consumer-resource metapopulation with local overexploitation of resources

The monophagous flightless weevil Hadramphus spinipennis causes the frequent local extinction of its host plant Aciphylla dieffenbachii through overexploitation. Both species are endemic and restricted to the Chatham Islands group (New Zealand). The weevil is classified as endangered and the plant as a threatened species. As this herbivore-plant system is locally unstable long-term persistence only appears possible via a metapopulation structure in which subpopulations are connected by dispersal. This paper investigates this hypothesis using a spatially explicit model of a consumer population whose resource is patchily distributed. The parameters are adapted to the H. spinipennis-A. dieffenbachii system. Our model includes local population dynamics and dispersal of the consumer, the destruction of host plant patches due to foraging and their regeneration. The results show that the key factor for long-term persistence is the short-range dispersal of the consumer with high mortality during dispersal. Only this highly inefficient dispersal prevents the synchronisation of local dynamics while ensuring sufficient colonisation of regenerated patches. We also show that although synchronisation through spatially correlated environmental fluctuations may be critical for long-term persistence, it cannot replace the strong effects of dispersal. Thus, in a consumer-resource metapopulation with deterministic local extinction, the strength and spatial scale of consumer dispersal in relation to the spatial and temporal scales of the resource patch dynamics (patch destruction and regeneration, number, size and distance of patches) play a vital role for long-term persistence. The consequence for conservation management is that such metapopulations could in fact profit rather than suffer from decreasing connectivity of resource patches.     

Modelling water transfer by rain-splash in a 3D canopy using Monte Carlo integration

Rain-splash is a very effective process in the short distance dispersal of biotic and abiotic particles leading to potential biological or chemical contamination within crop ecosystems. A physical approach to simulate water transfer by rain-splash in a 3D canopy structure is presented. This new approach is based on the explicit parameterisations of the mechanisms such as raindrop propagation, production and transport of splash droplets and their interception by canopy elements. The model theory and its numerical solution were based on the Monte Carlo integration technique. The calculation of droplet trajectories was validated by comparison with the previous published measured data (R²=0.99 for the terminal velocity and R²=0.93 for the distance travelled by droplets). Simulated patterns of splash dispersal through artificial canopies consisting of vertical cylinders compared well with measured patterns (R²=0.90). This new modelling approach is promising for studying particle dispersal by rain-splash in complex canopy structures.     

Population isolation exacerbates conservation genetic concerns in the endangered Amargosa vole, Microtus californicus scirpensis

The ability of a taxon to maintain adaptive flexibility in a stochastic environment is a function of the genetic diversity within the population. In small, fragmented populations, genetic variation can become depleted more quickly than in larger, more contiguous populations. Characterizing the patterns of genetic variation and differentiation associated with these processes is an important step in establishing conservation priorities. The Amargosa vole, Microtus californicus scirpensis, is an endangered rodent persisting in the small, fragmented marsh complex surrounding the Amargosa River near Death Valley, California. This naturally patchy system has existed since the end of the Pleistocene (approximately 10,000 y.b.p.), however, fragmentation has been exacerbated by recent anthropogenic changes. For this study, I used five nuclear microsatellite loci and the cytochrome-b region of the mitochondrial genome to quantify levels of genetic variation, population substructure, and patterns of gene flow in M.c. scirpensis. These data were compared to a broadly distributed subspecies, Microtus californicus sanctidiegi. Overall levels of nuclear genetic variation were significantly lower in M.c. scirpensis, whether measured in terms of diversity or heterozygosity, compared to more broadly distributed conspecifics. Moreover, only two haplotypes were recovered from the mitochondrial data with over 90% of the observed haplotypes being identical. Despite low genetic diversity, significant genetic subdivision among M.c. scirpensis populations was detected using both pairwise F_ST and Bayesian clustering methods. Furthermore, isolation by distance analyses reveal that an important landscape feature, ephemeral tributaries, is critical for dispersal among population clusters. Recommendations for conservation management are presented.     

Current genetic isolation and fragmentation contrasts with historical connectivity in an alpine lizard (Cyclodomorphus praealtus) threatened by climate change

Assessing levels of genetic diversity, connectivity and historical demography for threatened species provides important information for conservation management. We used a combination of the mitochondrial ND4 gene and seven microsatellite markers to examine both historical and recent population genetic structure and demography of the threatened alpine she-oak skink, Cyclodomorphus praealtus. This species is restricted to the “sky islands” of the Australian alpine region. Based on mtDNA, the New South Wales and Victorian regions are reciprocally monophyletic and highly divergent, with among population variation of 0.9 and net sequence divergence of 4.28%, which suggests that they should be considered separate Evolutionary Significant Units for management purposes. The mtDNA data also indicate historical connectivity between the three Victorian populations. However, a model-based clustering analysis of microsatellite genotypes identified strong population structure in Victoria, with three distinct populations that have no current inter-population gene flow. This suggests that the Victorian populations are effectively isolated from each other, and is indicative of very low dispersal capacity and a high degree of habitat specialisation. This is reinforced by the substantially lower genetic diversity within the lowest elevation population compared to the other higher elevation populations. We found no genetic signature of major changes in effective population size. These data provide a baseline for assessing future impacts of climate change on the genetic structure of this alpine endemic species.     

野燕麦繁殖和抗逆特性及其对小麦的他感效应研究

作为一种分布广泛的农田恶性杂草,野燕麦具有很强的环境胁迫适应能力,并且在种间竞争中常处于优势地位。本试验对野燕麦种群的繁殖与扩散特性、种子与幼苗抗逆特性及其根系分泌物对小麦的异株克生作用进行了研究,以期为找寻治理野燕麦的有效措施提供参考。结果表明:(1)野燕麦具有很强的种子繁殖与扩散能力,开花结实比小麦早,种子产量大、地下储量多、扩散距离远;(2)野燕麦种子对温度的适应范围较广,在室温与变温条件下均可以正常萌发,而冷冻后置于室温环境是其萌发最佳条件,总发芽率可达93.33%;(3)野燕麦种子对盐胁迫具有一定的适应性,在浓度低于1.8%的NaCl溶液中均可正常萌发;(4)野燕麦幼苗对盐胁迫和干旱胁迫具有较强的适应性,随着NaCl浓度和PEG浓度的升高,野燕麦幼苗叶片的脯氨酸含量显著增加(P0.01),过氧化氢酶活性呈现先增加后下降的趋势;(5)野燕麦根系水浸提液对小麦幼苗株高、干重、根长及根系活力均有极显著影响(P0.01),证明野燕麦根系分泌物对小麦幼苗生长具有异株克生作用。以上结果表明,野燕麦的竞争优势主要体现在强大的繁殖扩散能力、较强的抗逆性以及异株克生效应,相关研究可为大田野燕麦防除提供一定理论参考。     

Ecological and genetic measurements of dispersal in a threatened dragonfly

Leucorrhinia caudalis is a rare dragonfly, threatened throughout its European distribution. The species was formerly widespread in the Swiss lowlands, but only a single population remained in the 1980s. However, a spread has recently been observed, with additional ponds being colonised, sometimes at considerable distance. Despite this evidence of recent long-distance dispersal, it is unknown whether L. caudalis regularly moves among ponds or whether this is a rather rare event. A combination of an ecological mark-resight and a population genetic study was applied to investigate contemporary dispersal and the genetic footprint of the recent population history of L. caudalis in Switzerland. DNA for genetic microsatellite analysis was extracted from exuviae. The mark-resight study and the genetic analysis gave congruent results. They showed that L. caudalis is mostly a sedentary species, with only a few contemporary dispersal events over distances up to 5 km being observed. The genetic analysis was in agreement with the recent population history of the Swiss populations. The oldest and largest population showed large genetic diversity and acted as source population for the recent spread of L. caudalis in Switzerland. Recurrent gene flow among this source population and close populations caused substantial local genetic variation in the latter, as well as low population differentiation. The two recently founded distant populations (?30 km distance) were genetically less diverse and highly differentiated. These distant populations and another recently colonised population also expressed signatures of genetic bottlenecks.     

Does seed dispersal by sheep affect the population genetic structure of the calcareous grassland species Bromus erectus?

This study investigates the contribution of the traditional land-use system “sheep herding” (transhumance) to the gene flow among populations. Three sheep trails with altogether 12 calcareous grassland sites were chosen for the analysis of the tussock grass Bromus erectus, which is known to be the species with the highest rate of dispersal via sheep fleece. Each trail links two to four isolated populations. Allozymes were chosen as genetic markers. Six loci were used, five of which were polymorphic. While genetic variability within populations was found to be quite high (averaged h=0.352), only little between-population differentiation was measured (G_ST=5.5%). Despite this, the UPGMA-dendrogram of Nei's genetic distance shows genetic similarity relations largely congruent with the sheep herding trails. However, a (more) detailed comparison reveals several differences between genetic patterns and trail system. Hence, additional factors other than dispersal by sheep are regarded as being responsible for exchange of genes, and sheep herding is considered to have only minor importance for the genetic patterns of the investigated species. The comparison of Bromus erectus to species with the same life history traits (i.e. pollination by wind, a perennial life cycle, and autopolyploid constitution) shows similar values, thus providing further evidence for the relatively small contribution of dispersal via sheep fleece.     

Genetic structure of soil invertebrate populations: Collembolans,earthworms and isopods

Soil-living animals such as collembolans, earthworms and isopods are considered to be sedentary animals with only limited dispersal capacities. Therefore, gene flow among populations is expected to be low leading to significant population genetic differentiation due to random drift and local adaptation. We reviewed the literature to test this expectation. Our survey reveals a clear signature in the current pattern of genetic variation due to post-glacial colonization events. It also reveals that habitus can be a misleading predictor of dispersal capacity. In some species relatively high gene flow across considerable distances, most likely through passive dispersal (mediated by wind, water flow or animals), might counteract local genetic adaptation and will prevent loss of genetic variation.     

Use of prolamine polymorphism in studying genetic resources of forage grasses

Summary Electrophoresis of single seed prolamines was used for the analysis ofLolium perenne L.,Festuca pratensis Huds., andDactylis glomerata L. populations. Identification and registration of populations was carried out according to the frequencies of occurrence of genotypes with corresponding types of prolamine banding patterns. The publication sums up the problems of applied use of molecular markers for identification and registration of world genetic resources of forage grasses, analysis of the dynamics of population composition and other problems of plant growing, genetics, breeding and seed control. The approaches mentioned in the article are promising for their use in genetic banks as well as at the institutions which store collections of genetic resources of given crops and at universities and breeding stations.     

Genetic insights into population recovery following experimental perturbation in a fragmented landscape

We investigated the mechanism of population recovery in the Australian bush rat, Rattus fuscipes, following experimental perturbation in a fragmented landscape. Our study involved genetic monitoring over 24 months following the removal of animals from seven sites by intense trapping. A total of 171 bush rats was removed and statistical analysis confirmed a significant knockdown. At one site, local extinction followed the perturbation, while population sizes at 24 months varied from 37% to 90% of the original population size at the remaining sites. The outcomes of genetic analysis at 11 microsatellite loci and mtDNA indicated that population recovery was achieved predominantly by residual animals: recovery populations were genetically more similar to their respective pre-treatment population than near neighbours; assignment tests detected few immigrants; there was no influx of new immigrant alleles and haplotypes. This finding is consistent with emerging evidence for restricted gene flow in bush rats. The local extinction observed at one site indicate that, under severe perturbation restricted dispersal limits opportunities for, and the rate of, population recovery. Thus, while the bush rat appears resilient to substantial population size perturbation, once a critical minimum population size threshold has been reached, this species may be susceptible to local extinction. We concluded that some widespread species currently predicted to be extinction-resilient may, in fact, be at risk if localized extinctions occur and dispersal limitations prevent recolonisation.     

不同种群草鱼遗传结构的TRAP分析

应用TRAP(targetregionamplifiedpolymorphism)技术,对草鱼(Ctenopharyngodonidella)野生种群和两个人工养殖种群的遗传结构进行了比较。从15个引物组合中,筛选出7个扩增效果比较好的引物组合,在3个种群中共检测到103个扩增位点,其中野生种群、淡水中心种群和前洲种群的多态性位点分别为67、55和46,表明人工养殖草鱼种群的遗传多样性有所降低。与野生种群相比较发现,人工养殖种群仅33.98%位点的基因频率保持不变,表明人工养殖种群的遗传结构发生了改变。淡水中心种群、前洲种群与野生种群间的遗传距离分别为0.0421和0.0809。引物组合Ga5-800-E5扩增的结果发现,人工养殖种群中存在扩增位点明显减少的区域,这为寻找鉴定草鱼野生种群和养殖种群的遗传标记提供了依据。     

Use of a spectrum model and satellite cloud data in the simulation of wheat stripe rust (Puccinia striiformis) dispersal across the Tasman Sea in 1980

Long distance dispersal of plant pathogens has been simulated using different approaches. Models to predict dispersal of airborne spores often use simplified methods to estimate solar radiation in order to estimate mortality of spores. We incorporated a spectrum model into a Lagrangian approach to simulate dispersal of wheat stripe rust from Australia to New Zealand across the Tasman Sea. To estimate cloud cover conditions, satellite data were also used in our study. The combination of a spectrum model and satellite cloud data made it easy to identify sites at which viable spores were deposited. In our study, 0.6% of deposition near New Zealand followed exposure to UV radiation less than our mortality threshold (0.9 MJ m⁻²). Model spores of wheat stripe rust arrived at sites within 40 km from Otama, New Zealand, where wheat stripe rust was first reported in November 1980. It appeared that the uredospores were transported under an overcast cloud condition, which would increase the chance of successful infection on host crops. Further studies on the relationship between solar UV radiation and mortality of pathogen uredospores would facilitate the use of the spectrum model and satellite data to predict dispersal of airborne spores.     

Dispersal of breeding, adult male Phocarctos hookeri: Implications for disease transmission, population management and species recovery

Dispersal impacts on a range of population parameters making it a key piece of information in species conservation. Despite its importance, dispersal is poorly characterized for many species: pinnipeds are no exception. Understanding dispersal patterns of the New Zealand sea lion Phocarctos hookeri is crucial in the conservation management of the species as its recovery to a non-threatened status hinges on range recolonisation. In this study, we examined the movements of breeding adult male New Zealand sea lions within and following the breeding season of the 2002/03 austral summer using a novel multi colony approach. Based on resightings of 202 individually identifiable adult males, we found (1) a previously unappreciated, high level of dispersal by adult territorial males between breeding colonies during the pupping period and (2) that breeding males disperse to the extremes of the species’ range at the end of female oestrous. Our findings are contrary to the current paradigm of otariid breeding behaviour, which is believed to consist of prolonged, uninterrupted male territoriality based on intense male-male competition and sustained fasting. Adult male dispersal between colonies and across the species range has important implications for adult males as vectors of disease in three recent epizootics, species management and species recovery via recolonisation as males are apparently remaining part of a localized, vulnerable breeding population.     

Restricted gene flow in the clonal hepatic Trichocolea tomentella in fragmented landscapes

We studied the genetic diversity, gene flow and population structure among 18 populations of the clonal bryophyte Trichocolea tomentella located in Finland, Lithuania, the UK and Canada using DNA fingerprinting methods. T. tomentella is a habitat-limited, unisexual hepatic, which occupies spring and mesic habitats in woodland. The relatively small populations are increasingly fragmented with a high risk for extinction for extrinsic reasons. The presence of relatively high levels of genetic diversity regardless of population size highlights the role of even small remnant populations as important sources of genetic diversity in T. tomentella. The long-term accumulation of genotypes and somatic mutations may explain the observed levels of diversity. Gene flow among populations seems to be infrequent indicating dispersal limitation also on the relatively small spatial scale. Colonization within populations is not affected by isolation by distance suggesting the occurrence of random short-range dispersal of detached vegetative fragments. The population structure study confirmed the low mortality rates of shoots indicating a long life span of the clones in favourable conditions. Efficient ramet production by branching is likely to operate against interspecific competition. To conclude, T. tomentella appears to persist well in undisturbed habitats due to clonal regeneration, although restricted dispersal capacity is likely to prevent successful (re-)colonization in the potential habitat patches of recovering forest landscapes. The implications of the results for conservation are introduced.     

Population genetic structure in polar bears (Ursus maritimus) from Hudson Bay, Canada: Implications of future climate change

The primary habitat for polar bears is sea ice, yet unlike most of the high Arctic, Hudson Bay undergoes a summer ice-free period that forces all bears ashore until ice forms again in fall. Polar bear populations in the greater Hudson Bay region have been placed in four management units based primarily on data from tag returns from harvested animals, capture-recapture studies, and conventional and satellite telemetry. Our results indicate that there is a high level of gene flow among management units observed using 26 microsatellite loci and analysis of genetic profiles of 377 polar bears. However, individual-based Bayesian analysis identified population genetic structuring into three clusters and significant F_ST differentiation. Specifically, our data suggest differentiation of polar bears sampled from islands in James Bay. These results were in spite of the extensive dispersal capabilities of polar bears that could homogenize the population. Mapping of high-ancestry individuals suggests that two of the three clusters have foci in southern Hudson Bay and may be a result of predictable annual freeze-thaw patterns that are maintaining breeding ‘groups’. Predicted changes in the distribution and duration of sea ice in Hudson Bay suggest that gene flow among these clusters may be reduced in the future.     

Habitat fragmentation reduces genetic diversity and connectivity among toad populations in the Brazilian Atlantic Coastal Forest

Tropical rainforests are becoming increasingly fragmented and understanding the genetic consequences of fragmentation is crucial for conservation of their flora and fauna. We examined populations of the toad Rhinella ornata, a species endemic to Atlantic Coastal Forest in Brazil, and compared genetic diversity among small and medium forest fragments that were either isolated or connected to large forest areas by corridors. Genetic differentiation, as measured by F_ST, was not related to geographic distance among study sites and the size of the fragments did not significantly alter patterns of genetic connectivity. However, population genetic diversity was positively related to fragment size, thus haplotype diversity was lowest in the smallest fragments, likely due to decreases in population sizes. Spatial analyses of genetic discontinuities among groups of populations showed a higher proportion of barriers to gene flow among small and medium fragments than between populations in continuous forest. Our results underscore that even species with relatively high dispersal capacities may, over time, suffer the negative genetic effects of fragmentation, possibly leading to reduced fitness of population and cases of localized extinction.     

Spatial genetic variation of a riparian wolf spider Pardosa agricola (Thorell, 1856) on lowland river banks: The importance of functional connectivity in linear spatial systems

Habitat patches along river systems are often highly isolated and characterised by a high degree of heterogeneity at different spatio-temporal scales. The connectivity between river bank arthropod populations directly adjoining the river channel is often greatly disturbed due to river regulation. While flight-active arthropods easily disperse upstream, less mobile species are expected to show predominant downstream dispersal unless specific upward movements are prevalent. In linear river ecosystems, downstream drift of organisms may therefore prevail with subsequently strong asymmetrical gene flow. We analysed patterns of genetic variation within and among nine spatially structured populations of the highly stenotopic riparian wolf spider (Lycosidae) Pardosa agricola (Thorell, 1856) using Amplified Fragment Length Polymorphism (AFLP) markers. No evidence was found for downstream accumulation of genetic diversity. The high genetic diversity is hypothesised to be the result of historical drift processes. Nearby populations on the same river shore were less genetically differentiated compared with populations further away and/or on the opposite shore. This indicates that short-distance dispersal still occurs on river banks during low water flow levels, but at the same time that the river channel constitutes a physical barrier for species exchange between opposite shores. The rehabilitation of the riparian corridor will increase the amount of suitable habitat, the functional connectivity during periods of low flow-discharges and eventually riparian spider population persistence.