Patterns and correlates of plant diversity differ between common and rare species in a neotropical dry forest

Determining which factors affect species richness is important for conservation theory and practice. However, richness of common and rare species may be affected by different factors. We use an extensive inventory of woody plants from a tropical dry forest landscape in Yucatan, Mexico to assess the unique effects of environmental variables, spatial dependence of sampling sites, forest stand age and the combined effect of all groups of variables on species richness of woody plants with different levels of rarity (common, intermediate, rare, very rare)—according to their abundance, habitat specificity and spatial distribution range in the landscape. Analyzing separately common species and those with different levels of rarity uncovered contrasting patterns and correlates of species richness that were not apparent when focusing on all woody plants. In particular, richness of common and intermediate species was influenced mainly by environmental factors, whereas richness of very rare species was affected mostly by the unique effect of spatial dependence of sampling sites, suggesting a main role of environmental filtering and dispersal limitation, respectively. However, common and very rare species also responded inversely to some landscape metrics, revealing contrasting environmental preferences of these groups of species. These contrasting results suggest different underlying mechanisms and the need for very different conservation strategies. Therefore, basic and applied research on tropical forest biodiversity should consider separately species with different levels of rarity, focusing on which factors control variation in each level, and paying special attention to very rare species, generally the most specious and vulnerable to local extinction.     

Amphibian Species Contribute Similarly to Taxonomic,but not Functional and Phylogenetic Diversity: Inferences from Amphibian Biodiversity on Emei Mountain

Understanding the relationships between species,communities,and biodiversity are important challenges in conservation ecology.Current biodiversity conservation activities usually focus on species that are rare,endemic,distinctive,or at risk of extinction.However,empirical studies of whether such species contribute more to aspects of biodiversity than common species are still relatively rare.The aim of the present study was to assess the contribution of individual amphibian species to different facets of biodiversity,and to test whether species of conservation interest contribute more to taxonomic,functional,and phylogenetic diversity than do species without special conservation status.To answer these questions,19 000 simulated random communities with a gradient of species richness were created by shuffling the regional pool of species inhabiting Emei Mountain.Differences of diversity values were then computed before and after removing individual species in these random communities.Our results indicated that although individual species contributed similarly to taxonomic diversity,their contribution to functional and phylogenetic diversity was more idiosyncratic.This was primarily driven by the diverse functional attributes of species and the differences in phylogenetic relationships among species.Additionally,species of conservation interest did not show a significantly higher contribution to any facet of biodiversity.Our results support the claims that the usefulness of metrics based only on species richness is limited.Instead,assemblages that include species with functional and phylogenetic diversity should be protected to maintain biodiversity.     

Miniaturizing landscapes to understand species distributions

Species’ geographic distributions shape global patterns of biodiversity and therefore have long been of interest to ecology and conservation. Theory has generated valuable hypotheses about how landscape structure, dispersal, biotic interactions and evolution shape range dynamics, but most predictions have not been tested on real organisms because key variables are difficult to isolate, replicate or manipulate in natural ecosystems. An exciting and rapidly emerging approach is to extend classical microcosm and mesocosm systems to create experimental ‘micro-landscapes’. By enabling researchers to manipulate geographic features of interest, replicate landscapes, control colonization and follow dynamics across evolutionary timescales, micro-landscapes allow explicit tests of the ecological and evolutionary underpinnings of species distributions. Here we review the micro-landscape systems being used to advance biogeography, the major insights they have generated thus far, and the features that limit their application to some scenarios. We end by highlighting important questions about species’ biogeography that are ripe for testing with experimental micro-landscapes, particularly those of immediate concern given rapid global change, such as range contractions and constraints to range expansion.     

Spatial Assortment of Mixed Propagules Explains the Acceleration of Range Expansion

Range expansion of spreading organisms has been found to follow three types: (i) linear expansion with a constant rate of spread; (ii) bi-phase expansion with a faster linear expansion following a slower linear expansion; and (iii) accelerating expansion with a continuously increasing rate of spread. To date, no overarching formula exists that can be applied to all three types of range expansion. We investigated how propagule pressure, i.e., the initial number of individuals and their composition in terms of dispersal ability, affects the spread of a population. A system of integrodifference equations was then used to model the spatiotemporal dynamics of the population. We studied the dynamics of dispersal ability as well as the instantaneous and asymptotic rate of spread. We found that individuals with different dispersal abilities were spatially sorted with the stronger dispersers situated at the expanding range front, causing the velocity of expansion to accelerate. The instantaneous rate of spread was found to be fully determined by the growth and dispersal abilities of the population at the advancing edge of the invasion. We derived a formula for the asymptotic rate of spread under different scenarios of propagule pressure. The results suggest that data collected from the core of the invasion may underestimate the spreading rate of the population. Aside from better managing of invasive species, the derived formula could conceivably also be applied to conservation management of relocated, endangered or extra-limital species.     

The spread,establishment and impacts of the spiny water flea, <Emphasis Type="Italic">Bythotrephes longimanus,</Emphasis> in temperate North America: a synopsis of the special issue

More than most sub-disciplines of ecology, the study of biological invasions is characterized by breadth rather than by depth. Studies of expanding ranges of invaders are common, as are post-invasion case studies, but we rarely have a deep understanding of the dynamics and regulators of the processes of invasion and resultant ecological transformations. This is unfortunate because such depth may well be needed to develop targeted, knowledge-based, management plans. In this collection we provide this needed depth of study of the key aspects of the invasion process for the spiny water flea, Bythotrephes longimanus. We do so by presenting the results of the work conducted by researchers in the Canadian Aquatic Invasive Species Network (CAISN), and several of their American and European collaborators over the past half decade. Given its rapid spread in the Great Lakes basin in North America, and the decreases in pelagic biodiversity that have ensued, the last decade has witnessed a surge of research on Bythotrephes. In this collection we learn much about mechanisms and dynamics of its spread, about the key role of humans in that spread, about the importance of Allee effects to establishment and persistence, about choices and parameterization of risk assessment models, about the value of comparing “effects” in native and invaded regions, about complex probable interactions of the invasion with impending changes in the climate, and about the regulators of the invader’s abundance and impacts. There should be much of interest in the collection for aquatic ecologists and invading species biologists alike.     

Abundance–occupancy relationships

1.  The abundance and distribution of species tend to be linked, such that species declining in abundance often tend also to show declines in the number of sites they occupy, while species increasing in abundance tend also to be increasing in occupancy. Therefore, intraspecific abundance–occupancy relationships are commonly positive.
2.  The intraspecific pattern is mirrored by more general positive interspecific abundance–occupancy relationships: widespread species tend to be abundant, and narrowly distributed species rare.
3.  Here, we review recent research on these patterns based on the flora and fauna of the British Isles. We assess their generality, describe what is currently known about their structure, and summarize the results of tests of the several hypotheses proposed to explain their existence.
4.  The positive form generally exhibited by abundance–occupancy relationships, intraspecific or interspecific, has consequences for several areas of applied ecology, including conservation, harvesting, biological invasions and biodiversity inventorying. These implications are discussed briefly.     

Distribution patterns of macrobenthic biodiversity in the intertidal seagrass beds of an estuarine system, and their conservation significance

Intertidal stands of seagrass are important elements in the ecology of many estuaries yet the manner in which their associated macrobenthic biodiversity is distributed throughout any single estuary has never been determined. This has now been attempted for the Knysna estuarine bay in the Garden Route National Park, South Africa, an important site for one vulnerable and declining seagrass, Cape dwarf-eelgrass (Nanozostera capensis), and for several associated animals. Although spanning a salinity range of <5–35, the seagrass beds of this estuary contained a unitary fauna with local variation in frequency and abundance of the various species. Faunal biodiversity was highest along the main estuarine channel in the marine-influenced region and declined both upstream and into the sheltered peripheral zones, overall faunal abundance being inversely correlated with species diversity (though not with species richness). This pattern results in the beds around the 4 km shoreline of a single island near the mouth supporting 91 % of the total macrobenthic invertebrate species present in the system. This situation is discussed in relation to such data as are available from other comparable systems and to the conservation of estuarine seagrass faunal biodiversity. It is concluded that in general and with caveats for some individual species of concern where conservation resources are limited attention would most profitably be focused on the seagrass meadows of downstream estuarine areas.     

Eco‐evolution on the edge during climate change

We urgently need to predict species responses to climate change to minimize future biodiversity loss and ensure we do not waste limited resources on ineffective conservation strategies. Currently, most predictions of species responses to climate change ignore the potential for evolution. However, evolution can alter species ecological responses, and different aspects of evolution and ecology can interact to produce complex eco‐evolutionary dynamics under climate change. Here we review how evolution could alter ecological responses to climate change on species warm and cool range margins, where evolution could be especially important. We discuss different aspects of evolution in isolation, and then synthesize results to consider how multiple evolutionary processes might interact and affect conservation strategies. On species cool range margins, the evolution of dispersal could increase range expansion rates and allow species to adapt to novel conditions in their new range. However, low genetic variation and genetic drift in small range‐front populations could also slow or halt range expansions. Together, these eco‐evolutionary effects could cause a three‐step, stop‐and‐go expansion pattern for many species. On warm range margins, isolation among populations could maintain high genetic variation that facilitates evolution to novel climates and allows species to persist longer than expected without evolution. This ‘evolutionary extinction debt’ could then prevent other species from shifting their ranges. However, as climate change increases isolation among populations, increasing dispersal mortality could select for decreased dispersal and cause rapid range contractions. Some of these eco‐evolutionary dynamics could explain why many species are not responding to climate change as predicted. We conclude by suggesting that resurveying historical studies that measured trait frequencies, the strength of selection, or heritabilities could be an efficient way to increase our eco‐evolutionary knowledge in climate change biology.     

The effect of habitat on the range expansion of a native and an introduced bird species

Aim Range expansion across a heterogeneous landscape may depend on the habitat selected and used by the expanding species. If habitat selection influences range expansion then localities colonized by a species should contain a greater proportion of favoured habitat (and less non‐habitat) than other nearby localities not colonized. White‐winged doves (Zenaida asiatica) and Eurasian collared doves (Streptopelia decaocto) are two bird species that provide an excellent opportunity to test this hypothesis, because the geographic ranges of both species have been expanding in North America for more than two decades. Location Continental USA. Methods We used distribution data from the North American Breeding Bird Survey to test whether the landscapes occupied by each species contained a greater proportion of favoured habitat (urban land, grassland/pasture, shrub land and cropland) and a lower proportion of non‐habitat (forest land) than landscapes where doves were not found. We tested each species separately in each of three broad expansion areas, namely East, Central and West. We also compared rates of spatial spread between expansion areas and between the two species. Results As predicted, both species tended to occupy landscapes with greater proportions of urban land, shrub land and cropland but with less forest land compared with landscapes without doves, in all three expansion areas. Contrary to prediction, occupied landscapes tended to have slightly less grassland/pasture than unoccupied landscapes. Rates of spread differed between the two species and among expansion areas. Main conclusions Range expansion and the extent to which a species fills or saturates its range are influenced by the habitat ecology of the expanding species. Species colonize localities based on the availability of suitable habitat. However, the role of habitat in a species’ range expansion does depend somewhat on the greater geographical setting. Over large regional and geographical scales, range expansion (rate of spread and saturation) may proceed unevenly, suggesting that range expansion is a very dynamic and context‐specific process.     

The Fate of Cooperation during Range Expansions

Species expand their geographical ranges following an environmental change, long range dispersal, or a new adaptation. Range expansions not only bring an ecological change, but also affect the evolution of the expanding species. Although the dynamics of deleterious, neutral, and beneficial mutations have been extensively studied in expanding populations, the fate of alleles under frequency-dependent selection remains largely unexplored. The dynamics of cooperative alleles are particularly interesting because selection can be both frequency and density dependent, resulting in a coupling between population and evolutionary dynamics. This coupling leads to an increase in the frequency of cooperators at the expansion front, and, under certain conditions, the entire front can be taken over by cooperators. Thus, a mixed population wave can split into an expansion wave of only cooperators followed by an invasion wave of defectors. After the splitting, cooperators increase in abundance by expanding into new territories faster than they are invaded by defectors. Our results not only provide an explanation for the maintenance of cooperation but also elucidate the effect of eco-evolutionary feedback on the maintenance of genetic diversity during range expansions. When cooperators do not split away, we find that defectors can spread much faster with cooperators than they would be able to on their own or by invading cooperators. This enhanced rate of expansion in mixed waves could counterbalance the loss of genetic diversity due to the founder effect for mutations under frequency-dependent selection. Although we focus on cooperator-defector interactions, our analysis could also be relevant for other systems described by reaction-diffusion equations.     

Population and geographic range dynamics: implications for conservation planning

Continuing downward trends in the population sizes of many species, in the conservation status of threatened species, and in the quality, extent and connectedness of habitats are of increasing concern. Identifying the attributes of declining populations will help predict how biodiversity will be impacted and guide conservation actions. However, the drivers of biodiversity declines have changed over time and average trends in abundance or distributional change hide significant variation among species. While some populations are declining rapidly, the majority remain relatively stable and others are increasing. Here we dissect out some of the changing drivers of population and geographic range change, and identify biological and geographical correlates of winners and losers in two large datasets covering local population sizes of vertebrates since 1970 and the distributions of Galliform birds over the last two centuries. We find weak evidence for ecological and biological traits being predictors of local decline in range or abundance, but stronger evidence for the role of local anthropogenic threats and environmental change. An improved understanding of the dynamics of threat processes and how they may affect different species will help to guide better conservation planning in a continuously changing world.     

The Biodiversity Crisis: Lessons from Phylogenetic Sagas

The future of the world’s biodiversity involves preservation of individual species and, more importantly, preservation of the natural process by which the biosphere is populated. Inherited history allows species to carry within them the ability to adapt to changing circumstances. But inherited history also sets the limits for adaptation. Evolutionary potential is thus locked within shared history. Extinction removes, speciation replenishes. We must implement conservation policies that mimic the biotic expansion that sets the stage for speciation. If we do not provide space for species to spread out and find their own futures, building biodiversity reserves is tantamount to attempting to maintain standing diversity by blocking evolution. We must preserve as many species, associations, and places as possible in a geographic context large enough so that individual species may expand and contract and evolutionary dynamics can have free rein to shape the future.     

植物群落稀有种维持机制与土壤反馈的研究进展

自Janzen-Connell (J-C)假说提出后半个世纪以来, 生态学家在热带及亚热带森林对该假说开展的大量实证研究表明, 由专性天敌导致的J-C效应所引起的负密度制约是维持森林多样性和决定群落组成的重要驱动力, 该假说成功地解释了热带及亚热带森林的丰富多样性。土壤病原真菌所引起的植物-土壤负反馈是J-C效应最主要的表现形式。然而, 对于植物-土壤负反馈是否能够维持森林群落中的大量稀有种仍然存在许多争议。基于当代物种共存理论的“稀有种优势”假说认为, 只有在满足“可入侵准则” (即物种在稀有时具有种群增加的趋势)的前提下, 稀有种才能在群落中与其他物种长期共存。然而, 当前基于土壤反馈的实验结果与该理论预测相悖, 因此在稀有种的维持机制方面仍存在较大的分歧。本文通过介绍植物-土壤反馈理论, 整合了可能对稀有种维持有较大影响的因素, 包括共生菌根真菌、土壤养分以及植物细根性状等在影响土壤负反馈方面的相关研究, 并对这些因素如何影响群落中物种多度和稀有种在群落中的维持进行了探讨。最后, 我们也从其他角度探讨了一些对稀有种维持的研究。我们认为在未来对稀有种的研究中, 探讨使其长期存续的“优势”和制约其种群扩大的“限制”同等重要, 将当代物种共存理论与新技术、新方法相结合对于探究稀有种的维持机制具有重要的意义, 可为稀有种保护提供理论依据。     

Biodiversity maintenance in food webs with regulatory environmental feedbacks

Although the food web is one of the most fundamental and oldest concepts in ecology, elucidating the strategies and structures by which natural communities of species persist remains a challenge to empirical and theoretical ecologists. We show that simple regulatory feedbacks between autotrophs and their environment when embedded within complex and realistic food-web models enhance biodiversity. The food webs are generated through the niche-model algorithm and coupled with predator-prey dynamics, with and without environmental feedbacks at the autotroph level. With high probability and especially at lower, more realistic connectance levels, regulatory environmental feedbacks result in fewer species extinctions, that is, in increased species persistence. These same feedback couplings, however, also sensitize food webs to environmental stresses leading to abrupt collapses in biodiversity with increased forcing. Feedback interactions between species and their material environments anchor food-web persistence, adding another dimension to biodiversity conservation. We suggest that the regulatory features of two natural systems, deep-sea tubeworms with their microbial consortia and a soil ecosystem manifesting adaptive homeostatic changes, can be embedded within niche-model food-web dynamics.     

Conserving rare species can have high opportunity costs for common species

Conservation practitioners face difficult choices in apportioning limited resources between rare species (to ensure their existence) and common species (to ensure their abundance and ecosystem contributions). We quantified the opportunity costs of conserving rare species of migratory fishes in the context of removing dams and retrofitting road culverts across 1,883 tributaries of the North American Great Lakes. Our optimization models show that maximizing total habitat gains across species can be very efficient in terms of benefits achieved per dollar spent, but disproportionately benefits common species. Conservation approaches that target rare species, or that ensure some benefits for every species (i.e., complementarity) enable strategic allocation of resources among species but reduce aggregate habitat gains. Thus, small habitat gains for the rarest species necessarily come at the expense of more than 20 times as much habitat for common ones. These opportunity costs are likely to occur in many ecosystems because range limits and conservation costs often vary widely among species. Given that common species worldwide are declining more rapidly than rare ones within major taxa, our findings provide incentive for triage among multiple worthy conservation targets.     

Inferences from Common Species Communities for Selecting Conservation Areas

We aimed at identifying probabilistic areas of high biodiversity value over a large spatial scale, e.g., an entire country (France) within the temperate region, that could work as valuable conservation areas for both rare and common species. We aimed at identifying areas where four measures on bird community overpass a selected threshold value, by using probability interpolation models. The four variables considered were the rare species number, and three measures related to common bird community: relative abundance, estimated species richness and composition originality. For the latter, we developed an indicator that discriminates original from more ordinary compositions of common bird communities, accounting for the number of representatives in each species. This indicator was positively correlated to rare breeding species number, so that original composition of common bird communities allows us to identify areas also supporting the rarest species. Areas with high probabilities of two indicators reaching their threshold values represented 2.9% of continental France. Most double-indicator areas were those with high species richness and high relative abundance, then with original communities and high rare species number. The originality indicator was revealed valuable to identify the most suitable areas that could ensure the preservation of both rare and common species, at a national scale. By preserving sites supporting original common bird communities, conservationists would ensure the protection of rare and common species.     

Are there adequate data to assess how well theories of rarity apply to marine invertebrates?

Understanding the biology of rare species is a very important part of conservation biology. Most of our current understanding of rarity has, however, come from studies of terrestrial plants, birds, mammals and some insects. Freshwater and marine habitats are underrepresented in published studies of rare species or conservation biology. We therefore have little knowledge about how well our understanding of what makes particular species rare and how rare species persist applies to marine invertebrates which form a major component of coastal biodiversity. In this review, I examine some theories about rarity with respect to intertidal and shallow subtidal invertebrates to identify whether there are adequate data to apply these theories to marine invertebrates and how well such theories apply. The general conclusions are that the lack of quantitative data on abundances, ranges, habitat-requirements, dispersal and connectedness among populations for marine invertebrates means that their status as rare species cannot really be assessed appropriately. It is also unlikely that, without extensive sampling programmes and considerable expense, adequate data could be obtained for these small, cryptic animals, which typically have very patchy, variable and unpredictable patterns of distribution and abundance. Intertidal and subtidal assemblages are diverse, including species with many different life-histories from many phyla, occupying the same suite of habitats. It is therefore suggested that future research on rare organisms in marine habitats should build upon the long and successful history of experimental marine studies to test specific hypotheses about processes influencing rarity in the field. Such studies would not only add a new dimension to our current understanding of rarity, but would also provide badly-needed data on the status of rare marine invertebrates. abundances, invertebrates, marine, range, rarity     

Identifying priority sites for the conservation of freshwater fish biodiversity in a Mediterranean basin with a high degree of threatened endemics

The Guadiana River basin’s freshwater fish species richness, endemicity and threatened status (92% of native species are threatened) highlight the need for a large-scale study to identify priority areas for their conservation. One of the most common problems in conservation planning is the assessment of a site’s relative value for the conservation of regional biodiversity. Here we used a two-tiered approach, which integrates an assessment of biodiversity loss and the evaluation of conservation value through site-specific measures. These measures based on the reference condition approach introduce the ability to make objective comparisons throughout the Guadiana River basin, thus avoiding a priori target areas. We identified a set of biodiversity priority areas of special conservation significance—which contain rare taxa as well as intact fish communities—because of their outstanding contribution to the basin’s biodiversity. The inclusion of complete sub-basins in these priority areas might guarantee an optimal solution in terms of spatial aggregation and connectivity. However, the high spatial fragmentation to which the Guadiana River basin is submitted due to river regulation highlights the necessity of a systematic approach to evaluate the capability of the identified priority areas to maintain the Guadiana’s freshwater fish biodiversity. Handling editor: R. H. Norris     

Butterflies of European islands: the implications of the geography and ecology of rarity and endemicity for conservation

Depending on their faunal content islands can function as important ‘vehicles’ for conservation. In this study, we examine data on 440 butterfly species over 564 European islands in 10 island groups. To determine the status of the butterfly fauna, we have adopted two approaches, island-focused and species-focused, examined using principal components analysis and regression modelling. In the former, we relate species richness, rarity and endemicity to island geography (area, elevation, isolation and location in latitude and longitude); in the latter, species occurrence on islands is examined in relation to distribution, range, range boundaries, and altitudinal limits on the continent as well as species’ ecology (number of host plants) and morphology (wing expanse). Species on islands are also assessed for their status on the continental mainland, their distributional dynamics (extinctions, distribution changes) and conservation status (Red Data Book, European Habitat Directive, Species of European Conservation Concern and Bern Convention listing. Unexpectedly, we find that a large fraction of the European butterfly species is found on the islands (63.4%; 59% on small islands) comprising some 6.2% of the land area of Europe. Although species occurring on the islands tend, on the whole, to have lower conservation status and are not declining over Europe, 45 species are endemics restricted to the islands. Species richness shows only a weak locational pattern and is related as expected to isolation from the continental source and island area; but, both rarity and endemicity have distinctive geographical bias to southern Europe, on islands now under increasing pressure from climate change and increasingly intensive human exploitation. The vulnerability of species on islands is emphasised in the relationship of island occurrence (% occurrence and presence/absence of species on any island) with continental distributions. A large proportion of the variation (84%) is accounted by continental distribution, the southern range limit and lower altitudinal limit. Most species (69%) occur on very few islands (<5%). In view of ongoing species dynamics on islands, migrations and extinctions of species, island repositories of species depend in large part on conservation of butterflies at continental sources. The unique faunas and rare species on islands also depend on appropriate concern being given to the island faunas. Conservation of European islands is thus a two-way process, sustaining sources and conserving island refuges. Residuals from the regressions (islands with more or fewer species, rare and endemic species; species occurring more or less frequently than expected on islands) provide warning signals of regions and islands deserving immediate attention.     

How local extinction changes rarity: an example with Sonoran Desert fishes

That spatially rare species may be predisposed to extinction is a common tenet of ecology. However, the opposite side of the relationship – how extinction alters spatial rarity – remains little explored. We used an extensive biodiversity database to contrast patterns of spatial rarity of a biogeographic assemblage of native Sonoran fishes before and after an extensive, decades-long wave of extirpations. Focusing on 25 fish species native to the Lower Basin of the Colorado River, we analyzed two key aspects of spatial rarity: 1) species range sizes (expressed as kilometers of stream reach occupied) and 2) species' co-occurrence patterns. Native fish species that were spatially rare historically suffered disproportionate losses in occurrences. However, endemic species did not suffer increased losses relative to non-endemic (but still native) species of comparable rarity. Species' geographic range sizes were concordant through time, with spatially rare species remaining rare after extensive extirpations relative to species that were historically more widespread. In contrast, extirpations greatly disrupted patterns of species co-occurrence on both local and regional scales. Over 50% of the species pairs that historically co-occurred (in the same 5  km reach) no longer co-occur anywhere in the Lower Basin, and species pairs that infrequently co-occurred in historic times suffered greater proportional losses than did more widely co-occurring pairs. Such changes in the relationship between spatial rarity and species richness deserve attention because they inhibit conservation planning (decreasing the efficiency of reserve design) and reduce interaction diversity altering opportunities for long-term co-evolutionary change.