A survey method for evaluating drought-sensitive bryophytes in fragmented forests: A bryophyte life-form based approach

Drought-sensitive bryophytes are especially vulnerable to edge effects caused by forest fragmentation. Because of increased forest fragmentation, these bryophyte species are declining and are in need of conservation. Considering that a field survey including all bryophyte species is very time consuming, methods that make identification less difficult should be tested for their usefulness in conservation. This paper describes an alternative survey method that utilizes the correlation between bryophyte life-forms and microclimates for evaluating drought-sensitive bryophytes. This survey method was examined using epiphytic bryophyte flora on tree trunks in 27 fragmented forests of Kyoto city, in the western part of Japan. The usefulness of life-forms for this evaluation was discussed based on the correlation of life-forms with species richness and microclimates. The results indicated that, while life-from richness was considered to reflect the heterogeneity of moisture availability and light intensity in bryophyte habitats, a certain life-form category (for example, fans and dendroids) seemed to correlate with drought sensitivity of bryophytes. Considering this correlation, an alternative survey method was proposed that utilizes the richness of hygrophilous life-forms. This survey method seems to be more cost-effective than a traditional all-species survey, and provides an important step toward the conservation of drought-sensitive species.     

Are habitat-tracking metacommunities dispersal limited? Inferences from abundance-occupancy patterns of epiphylls in Amazonian forest fragments

In theory, habitat fragmentation alters plant community dynamics by influencing both local (within patch) and regional (among patch) processes. However, the lengthy generation times of plant taxa relative to the short duration of most experiments has precluded studies from assessing the impact of fragmentation at both local and regional scales. Due to their accelerated life cycles, high rates of local extinction, and naturally patchy substrates, epiphyllous bryophyte assemblages are an appropriate plant guild for empirically testing metacommunity-based predictions associated with habitat fragmentation. By examining the local abundance and regional distribution patterns of 67 epiphyllous (leaf-inhabiting) bryophyte species in an experimentally fragmented landscape in Amazonia, we demonstrate that changes in local abundance wrought by habitat fragmentation are best explained by fragment size rather than proximity to forest edge. Furthermore, evidence of a simultaneous inter-specific decline in epiphyll local abundance and regional distribution in small (1- and 10-ha) forest fragments corroborate with metapopulation-based predictions highlighting the importance of immigration in buffering from patch extinction risk (i.e., the rescue effect). Collectively, these results provide indirect evidence that dispersal limitation, rather than compromised habitat quality attributable to edge effects, likely account for species loss from small tropical forest fragments. Whether dispersal limitation is due to increased insularity from regional sources for epiphyll recolonization or rather to lowered within-fragment dispersal potential is unknown; nonetheless, the long-term persistence of these microscopic plant metacommunities depends on the preservation of rain forest reserves of at least 100-ha in size.     

Bird species abundance–occupancy patterns and sensitivity to forest fragmentation: Implications for conservation in the Brazilian Atlantic forest

Developing a predictive theory for species responses to habitat fragmentation is a large, complex challenge in conservation biology, and meeting this challenge likely requires tailoring predictions to specific habitats and taxa. We evaluate the effects of fragmentation on forest birds living in three distinct forest ecosystems found in Brazilian Atlantic forest: seasonal semi-deciduous forest (SF), mixed rain forest (MF), and dense rain forest (DF). We test the hypotheses that (1) bird species most prevalent in SF (relative to other habitat types) will be least vulnerable to population declines in fragmented SF, and (2) species with stronger affiliations with DF or MF will be relatively more sensitive to fragmentation in SF. Using an exploratory statistical technique called “Rank Occupancy–Abundance Profiles (ROAPs),” we compared distribution and abundance of birds among large “continuous” areas of each forest type, then compared abundances in continuous SF forests with patterns of abundance in small fragments of SF, where edge effects could play a marked role in population dynamics. Overall, 39 species showed substantially lower local abundance, occupancy, or both in SF fragments versus continuous SF. As predicted, a higher proportion of bird species associated with DF appeared sensitive to fragmentation in SF; by contrast, species most abundant in SF and MF were similarly abundant in fragmented SF. Our study demonstrates how quantifying distribution and abundance in diverse habitats may enhance managers’ ability to incorporate species-specific responses to human disturbances in their conservation plans, and points out ways that even small reserves may have significant conservation value.     

Impacts of Industrial Polluters on Bryophytes: a Meta-analysis of Observational Studies

Identifying the factors responsible for the diversity of responses of biota to industrial pollution is crucial for predicting the fates of polluted ecosystems. A meta-analysis based on 49 field studies conducted around 47 point polluters demonstrated that the individual (growth and reproduction) and community (abundance and species richness) characteristics of bryophytes in polluted habitats are reduced to about a half of the values observed in unpolluted sites. Non-ferrous smelters cause a stronger reduction in species richness and larger changes in species composition than other types of polluters. The magnitudes of the effects of pollution on the abundances of individual bryophyte species are not linked with their taxonomic position, life form or Ellenberg indicator values for light, moisture and nitrogen. The variation in species?? responses to pollution is mostly explained by differences in their reproductive characteristics; bryophyte species that possess special forms of vegetative reproduction and those that produce abundant sporophytes are more successful in polluted habitats. Ranking of bryophyte species according to their sensitivity to pollution is independent of the type of the polluter. Changes in bryophyte cover follow changes in tree cover, but not changes in the cover of the vascular field layer in the same pollution gradients. Pollution impacts cause stronger adverse effects on bryophytes in warmer climates.     

Integrating demographic and genetic approaches in plant conservation

We summarize the problems that populations of formerly common plants may encounter when habitat fragmentation isolates them and reduces population size. Genetic erosion, inbreeding depression, Allee-effects on reproductive success, catastrophes and environmental stochasticity are illustrated with studies on species that have recently become rare in The Netherlands due to habitat fragmentation. These clearly indicate that population viability is negatively affected. We also show that in the recent literature (since 1980), most studies on the conservation of rare plants have addressed population genetic structure and relationships between genetic variation and population size. Though important, these studies are not suitable for assessing the importance of genetics for population viability. In turn, demographic studies can detect changes in vital rates in small populations, but cannot reveal underlying genetic causes. Fitness and demographic studies are also well-represented in the literature, but remarkably few studies have attempted to integrate empirical demographic and genetic studies. We discuss two approaches to fill this very important lacuna in our knowledge. One of these constructs matrix-projection models on the basis of demographic censuses of—if possible—large and viable populations, and combines these with the results of experiments to determine inbreeding effects on demographic transitions and, subsequently, population growth and extinction. The other approach is to demographically monitor experimentally created small and large populations with low and high genetic variation and measure their actual growth rates and probabilities of extinction. We conclude that demography and demographic-genetic experiments should play a central role in plant conservation genetics.     

Plant species extinction debt in a temperate biodiversity hotspot: Community, species and functional traits approaches

Destruction and fragmentation of (semi-) natural habitats are considered the main causes of biodiversity loss worldwide. Plant species may exhibit a slow response to fragmentation, resulting in the development of an extinction debt in fragmented plant communities. The detection of extinction debt is of primary importance in habitat conservation strategies. We applied two different approaches proposed in the literature to identify extinction debt in South-East Belgium calcareous grasslands. The first method compared species richness between stable and fragmented habitat patches. The second explored correlations between current species richness and current and past landscape configurations using multiple regression analyses. We subsequently examined results generated by both methods. In addition, we proposed techniques to identify species that are more likely to support extinction debt and associated functional traits. We estimated a respective extinction debt of approximately 28% and 35% of the total and specialist species richness. Similar results were obtained from both methods. We identified 15 threatened specialist species under the current landscape configuration. It is likely the landscape configuration no longer supports the species habitat requirements. We demonstrated that non-clonal species are most threatened, as well as taxa that cannot persist in degraded habitats and form only sparsely distributed populations. We discussed our results in light of other studies in similar habitats, and the overall implications for habitat conservation.     

Theory meets reality: How habitat fragmentation research has transcended island biogeographic theory

Island biogeography theory (IBT) provides a basic conceptual model for understanding habitat fragmentation. Empirical studies of fragmented landscapes often reveal strong effects of fragment area and isolation on species richness, although other predictions of the theory, such as accelerated species turnover in fragments, have been tested less frequently. As predicted by IBT, biota in fragments typically ‘relax’ over time towards lower species richness. Beyond these broad generalizations, however, the relevance of IBT for understanding fragmented ecosystems is limited. First, IBT provides few predictions about how community composition in fragments should change over time, and which species should be most vulnerable. Second, edge effects can be an important driver of local species extinctions and ecosystem change, but are not considered by IBT. Third, the matrix of modified vegetation surrounding fragments—also ignored by IBT—can strongly influence fragment connectivity, which in turn affects the demography, genetics, and survival of local populations. Fourth, most fragmented landscapes are also altered by other anthropogenic changes, such as hunting, logging, fires, and pollution, which can interact synergistically with habitat fragmentation. Finally, fragmentation often has diverse impacts on ecosystem properties such as canopy-gap dynamics, carbon storage, and the trophic structure of communities that are not considered by IBT. I highlight these phenomena with findings from fragmented ecosystems around the world.     

Population demography and fecundity do not decline with habitat fragmentation in the rainforest tree Macadamia integrifolia (Proteaceae)

Habitat fragmentation is often associated with reduced levels of fitness and local extinction of plant species, and consequently poses a major threat to the persistence of species worldwide. The majority of demography-based fragmentation studies to date have focussed primarily on fragmentation impacts on individual plant fecundity. Here we investigate the impact of habitat fragmentation on the demography (plant height classes and density) and key population dynamic processes for the rainforest tree species Macadamia integrifolia (Proteaceae). Raceme and fruit production and seedling emergence across fragmented sites exceeded that in more intact sites with no apparent difference in short-term mortality rates. Fecundity of flowering trees did not appear to be affected by fragmentation. Instead, overall reproductive output in fragmented sites was enhanced relative to undisturbed sites due to a higher proportion of reproductively active individuals. The probability of flowering and fruiting was negatively correlated with the projected foliage cover (PFC) surrounding individual trees, and average PFC was significantly lower in small and medium fragments, suggesting light availability as a potential contributor to the trends observed here.This study demonstrates that the short-term effects of habitat fragmentation on population viability may not necessarily be detrimental for some species, and highlights the importance of assessing not only the fecundity of flowering individuals but also the proportion of individuals reproducing within fragments.     

Bryophyte (moss and liverwort) conservation under remnant canopy in managed forests

Management of forest for timber values presents potential threats for forest floor bryophytes, as localized disturbances are applied across landscapes. Dispersal limitation may exacerbate local extirpation, by preventing recolonization within a cut-block rotation period. Populations of forest floor bryophytes that persist under those patches of tree canopy remaining after clear-cutting could reduce dispersal distances and thereby contribute to conservation of species across the landscape. We examined bryophyte guilds (liverworts, forest-habitat mosses and colonist-pioneer mosses) and community composition in relation to habitat quality (microclimate and substrate) in five treatment classes in New Brunswick Acadian forest, 4 years after harvest. Four potential refugium classes with intact substrate were examined: three were characterized by remnant canopy height, one was treeless. These were compared to clear-cut areas with substrate disturbance. Microclimate (temperature, total daily photosynthetically active radiation and vapour pressure deficit) differed significantly between areas with and without remnant canopy, but differed little among refugium classes. This suggests that any remnant canopy moderates microhabitat relative to treeless areas. Liverworts and forest-habitat mosses were more frequent under remnant canopy than in open and clear-cut areas, with 25 species present only under remnant canopy. Environmental variation explained approximately 24% of bryophyte pattern, highlighting the potential influence of the pre-harvest community, which we could not document. In the absence of substrate disturbance, patches of remnant canopy provide potential refugia for some forest-habitat bryophytes. Characteristics of effective refugia (size and shape) should be determined by assessments of their impacts on: (i) change in bryophyte communities in refugia relative to natural dynamics and (ii) recolonization of adjacent areas.     

Habitat fragmentation reduces plant fitness by disturbing pollination and modifying response to herbivory

In fragmented landscapes plant species are often confined to remnants of formerly more widespread habitats, with many of their populations being small and isolated. This study experimentally examined the effects of population size and isolation on pollination, herbivory and reproductive success in the forest herb Phyteuma spicatum (Campanulaceae). In an experiment in which population size and isolation were manipulated using plants from the same origin, population size positively affected pollinator visitation, but did not alter the generally high levels of herbivory. As a result, seed production was higher in large populations. Conversely, plants originating from 14 natural populations of varying size and degree of isolation did not differ in reproductive success when grown in the same environment, suggesting similar attractiveness to pollinators and reproductive potential. The intensity of herbivory, however, was higher in progeny of small populations, at least in terms of the proportion of biomass removed. In both experiments, there were no effects of population isolation. The results suggest (1) that small population size decreases reproductive success via direct negative effects on plant-pollinator interactions, (2) that this pattern is not offset by herbivory, but (3) that herbivory enforces fragmentation effects on pollination by further reducing the number of flowering individuals and (4) that habitat fragmentation may influence plant fitness by affecting plant response to herbivory. The effects of habitat fragmentation on plant populations in present-day landscapes are thus complex, illustrating the need for more integrated studies in conservation biology that take into account both mutualistic and antagonistic plant-animal interactions.     

Phenotypic differentiation in fitness related traits between populations of an extremely rare sunflower: Conservation management of isolated populations

Knowledge of the genetic and demographic consequences of rarity is crucial when evaluating the effects of habitat loss and fragmentation on population viability, and for creating management plans in rare plant species. Reduction in population size and in the number of populations can lead to decreased genetic diversity and increased inbreeding. Genetic diversity is often correlated with fitness and is frequently used to identify populations of greatest conservation concern, or those that may be good candidates for ex situ conservation programs. However, an association between these factors is not always clear, and crossing studies evaluating whether there is phenotypic differentiation among populations in fitness related traits can inform managers of suffering populations or good sources for ex situ materials. Crossing studies can also evaluate the potential for genetic rescue to boost fitness in suffering populations. To address these questions, we conducted two generations of controlled crosses between populations of the extremely rare and fragmented sunflower, Helianthus verticillatus. We measured achene viability, germination, survival, and pollen viability (F₁ only) in 176 F₁ and 159 F₂ families. The populations were differentiated with respect to phenotypic fitness measures with one population having significantly lower achene viability and germination. Also, the potential for genetic rescue was observed as gene flow into the less fit population resulted in higher fitness measures in both the F₁ and F₂. Results are discussed with respect to the importance of combining genetic marker data with crosses and the implications for conservation in disjunct populations of rare species.     

Application of GIS in plant conservation programmes in Portugal

A plant conservation programme is a complex process that requires several sets of studies. The relationships between plant location and its environment play an important role. Nowadays, the use of a GIS constitutes an essential complement for these studies that allows the incorporation of space and the analysis of these ecological interactions. GIS is being used at the Lisbon University Botanical Garden as a tool for conservation programmes on several plant groups and situations. Four case studies are presented in this work: (1) comparing ecological patterns between local and regional scale for the endangered bryophyte Bruchia vogesiaca Schwaegr.; (2) selecting protected areas according to habitat suitability—the case of endangered Portuguese bryophytes; (3) analysing the impact of the alien Carpobrotus edulis (L.) N. E. Br. on endemic plant species at the Berlengas Natural Reserve; and (4) ecogeographical survey for selection of sites for seed collection in order to guarantee a representative sample of the existing genetic diversity. Finally, this work discusses how the implementation of a GIS can help to optimise results and fieldwork effort.     

The effect of habitat fragmentation on communities of mutualists: Amazonian ants and their host plants

The consequences of fragmentation for communities of mutualist partners are for the most part unknown; moreover, most studies addressing this issue have been conducted on plant-pollinator communities. We evaluated how the experimental fragmentation of lowland Amazonian rain forest influenced a community of ant-plant mutualists. We inventoried a total of 1057 myrmecophytes in four fragments and four continuous forest sites; the twelve plant species recorded were occupied by 33 ant morphospecies, of which 11 were obligate plant inhabitants. Neither plant species richness, ant species richness, nor total ant-plant density were significantly lower in forest fragments. However, eight of the plant species, including three of the four most common, had higher mean densities in continuous forest than fragments. Of these four species, only one (Cordia nodosa) had significantly different colonization rates between habitat types, with higher colonization rates of plants in fragments. This may be because the Azteca species it is associated with increases in abundance in forest isolates. Although our results suggest that communities of ant-plant mutualists are likely to persist in fragmented tropical landscapes 25 years after fragment isolation, most species are rare and populations sizes in fragments are extremely low. Environmental and demographic stochasticity could therefore limit long-term population viability. We suggest future studies focus on evaluating how fragmentation has altered herbivore pressure and the dispersal of ants and plants to fragments, since the interaction of these factors is likely to have the greatest impact on long-term patterns of population persistence.     

Fragmented but not isolated: Contribution of single trees, small patches and long-distance pollen flow to genetic connectivity for Gomortega keule, an endangered Chilean tree

Habitat fragmentation is a major threat to species survival worldwide due to genetic isolation, inbreeding depression, genetic drift and loss of adaptive potential. However the data on how gene-flow changes following habitat fragmentation is contradictory. If there is significant gene-flow between spatially isolated populations then limited conservation resources could be directed away from projects to ‘establish genetic connectivity’ and used to address other consequences of habitat fragmentation.This research focused on an endangered tree species Gomortega keule (Gomortegaceae) in a fragmented landscape in the Central Chile Biodiversity Hotspot and addressed three questions: (1) How far does pollen move between pollen donors and seed trees and what is the shape of the dispersal curve? (2) Do insect pollinators travel outside of forest patches? (3) Do small populations and single trees contribute to genetic connectivity across the landscape?Paternity analysis results show that G. keule’s insect pollinators travel outside of forest patches, over distances of 6 km, beyond the scale of population fragmentation or genetic structure. Pollen moved from small sites and single trees into large sites, as well as in the other direction, indicating these sites play a key role as functioning elements of the wider population and as stepping stones between sites. Fragmentation at the scale investigated has not led to genetic isolation, thus genetic connectivity per se is not a conservation priority. Other consequences of land-use change, specifically continuing habitat loss and population reduction, still threaten the survival of the species.     

Patch size and isolation effects on epiphytic and epiphyllous bryophytes in the fragmented Brazilian Atlantic forest

An inventory of the understory forest epiphyte and epiphyllous bryophyte floras was carried out in eight Atlantic rain forest fragments. The fragments were between 7 and 500 ha in size and belonged to two areas (lowland and submontane) of the state of Pernambuco, northeastern Brazil. The aim of this study was to investigate the effect of habitat fragmentation and changes in the natural landscape on community structure (composition, richness, diversity, and abundance). Although the influence of altitude was noted in this study by increasing richness, diversity, and abundance, it was clear that in some fragments the influence of fragment size and isolation can be more important than this positive environmental factor. Fragment size and isolation affect both communities but the last one seems to be a stronger threatening factor for the epiphylls. The least isolated and the largest fragments housed the richest floras - especially in relation to the epiphylls - and had the greatest proportion of shade species. Habitat fragmentation negatively affected epiphytic and epiphyllous bryophytes and increased the representation of species with larger niches (generalists) while decreasing the representation of species with smaller niches (typically found in shady or in sunny areas). The results suggests that the critical fragment size necessary for bryophyte preservation must be correlated with insularity levels; for epiphytes, however, it is likely that low levels of isolation cannot compensate borderline effects as with the epiphyllous bryophytes.     

Urban domestic gardens (XIII): Composition of the bryophyte and lichen floras, and determinants of species richness

Private, residential gardens form a substantial proportion of the undeveloped land in urban areas. Evaluating their role in supporting biodiversity is crucial to (i) predicting which plant and animal species can persist in towns and cities, (ii) understanding the regional impacts of urbanisation, and (iii) guiding sympathetic garden management by owners. To obtain baseline information on a poorly-studied component of garden biodiversity, we measured the size and composition of the cryptogam assemblages in 61 domestic gardens in the city of Sheffield, UK. A total of 67 bryophyte and 77 lichen taxa were recorded. Bryophytes ranged from 3 to 24 species per garden, with a mean richness of 11.3 species; lichens ranged from 2 to 30, with a mean of 14.9 species. Stone substrates supported the highest lichen richness, although minor substrates contributed unique species. Just over one fifth of bryophyte species were recorded in grass lawns, and these were more widespread than those of other habitats. Most cryptogams were scarce, with around one quarter of both bryophytes and lichens occurring in single gardens, and only 10% were found in more than half of the gardens. Garden area - correlated with substrate richness - and garden altitude were the only two factors explaining variation in cryptogam richness (bryophytes 39.1%, lichens 32.4%). Positive correlations existed among bryophyte, lichen and vascular plant richness, and these were only partially mediated by the effect of garden area. Therefore the opportunity remains for garden owners to support cryptogam richness, the most effective action being to enhance substrate diversity.     

Diversity of dead wood inhabiting fungi and bryophytes in semi-natural beech forests in Europe

Saproxylic organisms are among the most threatened species in Europe and constitute a major conservation problem because they depend on the most important forestry product - dead wood. Diversity of fungal and bryophyte communities occurring on dead beech trees was analyzed in five European countries (Slovenia, Hungary, The Netherlands, Belgium and Denmark) considering tree level species richness (TLSR), country level species richness (CLSR), frequency distributions of species, occurrence of threatened species and relations between TLSR and decay stage, tree size and countries. Altogether 1009 trees were inventoried in 19 beech dominated forest reserves.The number of fungi in the full dataset was approximately three times larger (456 versus 161 species) and the proportion of low frequent species was higher than among bryophytes. The species richness of bryophytes and fungi was significantly different among countries considering both TLSR and CLSR. In addition the diversity patterns deviated considerably between the two groups of organisms. Slovenian sites appeared to be biodiversity hotspots for bryophytes characterized by high TLSR and CLSR and a high fraction of threatened species. Hungarian sites had somewhat lower bryophyte diversity, while the Atlantic region had deteriorated assemblages. Fungal species richness was very high in Denmark, but the Hungarian and Slovenian sites were richer in threatened and low frequency species. Tree size was better able to explain variation in TLSR in both organism groups than decay stage. TLSR was found to vary significantly between countries but the difference was most considerable in the case of bryophytes.The diversity patterns of both organism groups along the investigated geographical gradient appear to be influenced by both climatic and management related factors (forest history, dead wood availability and continuity, habitat fragmentation). There is no doubt that an increase in the abundance of dead wood in European beech forests will benefit diversity of saproxylic fungi and bryophytes, especially if a continuous presence of large diameter logs are secured within individual stands.     

Does time or habitat make old-growth forests species rich? Bryophyte richness in boreal Picea mariana forests

The relative importance of time since disturbance and habitat variables in creating diversity in old-growth forests will influence conservation strategies. However, the independent roles of these factors are not well understood, as they are rarely examined independently. This study examines the respective roles of habitat variables and time (stand age) in determining bryophyte diversity in Picea mariana (Mill. (BSP)) forests. Bryophytes are frequently used as indicators of old-growth forest, but their true dependence on forest continuity is unknown. Bryophytes were classified into taxonomic-habitat guilds: true mosses (forest), forest liverworts, bog liverworts and sphagna (bog). Diversity increased with age and peaked at approximately 275 years since fire, driven by liverworts. Multiplicative habitat modeling indicated that time and habitat played different roles for the different taxonomic-habitat guilds. True mosses and forest liverworts were primarily influenced by habitat variables, while sphagna and bog liverworts were influenced by time and habitat variables. The models for sphagna were particularly strong, indicating that many important factors were included, while forest liverwort models were particularly weak. This unexplained variability may represent site specific random factors, such as secondary disturbances that create habitat in a fully occupied space. Overall, high richness was created by small species dependent on habitat variables and chance factors for establishment. Therefore, time since disturbance was not the primary factor limiting richness in these forests. These results suggest that in boreal North America, where forest fragmentation is limited and recent, conservation strategies that emphasize habitat variables rather than forest continuity may be effective for some bryophytes.     

A GIS-based survey for the conservation of bryophytes at the landscape scale

Geographical information system (GIS) data on landscape features and land use were collected to predict bryophyte diversity and conservation value in order to determine the factors that favour bryophytes at a large geographical scale and propose the relevant conservation measures. Total species diversity and diversity in species of high conservation value were highly correlated, and the landscape features promoting them were the proportion of military lands, steep slopes, and broadleaf woodland. Military lands seemed to be especially important for the conservation of endangered species highly specialized to open habitats maintained by the appropriate level of disturbance. Woodland cover was also as a key factor for bryophyte diversity but landscape heterogeneity, such as steep slopes with a range of contrasting ecological conditions, was required to reach the highest species numbers. The GIS-based approach presented here may help focusing the attention on sites exhibiting the appropriate landscape features in terms of conservation, which is especially relevant in the context of the European network `Natura 2000' for designating, conserving, and managing the sites of high biological value.