Drivers and interrelationships among multiple dimensions of rarity for freshwater fishes

Species can be rare or common in three different dimensions: geographic range size, habitat breadth, and local abundance. Understanding drivers of rarity are not only fundamentally interesting; it is also pertinent for their conservation. We addressed this challenge by analyzing the rarity of 291 native freshwater fishes occurring in ca 3500 independent stream reaches that span a broad environmental gradient across continental USA. Using phylogenetic regression and path analysis, we examined the concordance among the three rarity dimensions, and identified possible mechanisms by which species life‐history, habitat affinities, and biogeography drive variation in rarity. Weak double extinction jeopardies were driven by weakly positive correlations between habitat breadth and local abundance, and between habitat breadth and geographic range size. However, a triple extinction jeopardy was averted as local abundance and range size were not positively linked in our study. This is because large‐river and lacustrine habitat use mediated a trade‐off between local abundance and range size. Large rivers and lacustrine habitats represent important dispersal pathways and refugia that enabled fishes to acquire wide ranges; however, species using these habitats are less abundant overall because they are less adapted to small lotic channels, which comprise the majority of stream habitats in the US. Life‐history traits were key in governing the relationship between abundance and range size as large‐river and lacustrine habitat use were driven by body size, egg size, and parental care. Our analysis contributes novel insights into mechanisms that underlie multiple dimensions of rarity in freshwater fish and informs the prioritization of multiply rare species for conservation.     

How species traits and affinity to urban land use control large-scale species frequency

Aim Although urban areas only occupy c. 2.8% of the earth's land surface, urbanization threatens biodiversity as areas of high human population density often coincide with high biodiversity. Therefore, nature conservation should concentrate on both remote areas and densely populated regions. Protecting rare plant species in rural and urban areas can contribute to the protection of biodiversity. We therefore need to understand why species are rare. Studies on causes of rarity often concentrate on either plant traits or extrinsic threats (such as habitat fragmentation or nitrogen enrichment). However, there are only a few studies that combine traits and extrinsic threats, although such analyses might clarify causes of rarity. We assessed how the affinity of vascular plant species to urban land use (‘urbanity’) interacts with plant traits in determining species frequency. Location Germany, resolution c. 12 km × 11 km. Methods Species with a low frequency may be rare because they occur in rare habitats or because of other reasons, although their habitat is frequent. Therefore, we calculated the frequency of species corrected for habitat frequency, i.e. relative species frequency. We explained relative species frequency by the interactions of species traits and species affinity to urban land use using generalized linear models. Simultaneous autoregressive error models controlled for phylogenetic relationships of species. Results Relative species frequency depends on species affinity to urban land use, independent of the different interactions between traits and urbanity used as predictors. The higher the urbanity the higher is species frequency. Urbanity interacts with species preferences towards temperature and soil acidity. Moreover, dispersal, nitrogen preferences and origin explain relative species frequency, amongst others. Main conclusions Many rare species, especially those preferring cool or acidic habitats might already have disappeared from urban areas. Analyses that combine species traits and environmental effects can explain the causes of rarity and help to derive better conservation strategies.     

Can We Distinguish Plant Species that are Rare and Endangered from Other Plants Using Their Biological Traits?

Understanding the factors responsible for species rarity is crucial for effective species conservation. One possible approach to obtaining information about causes of species rarity is to compare rare and common species. We analyzed the biological and ecological traits of critically endangered (CR) plant species of the Czech Republic. We compared the vegetative, generative and ecological traits of CR species with: i) common closely related species (a form of phylogenetic correction), ii) common closely related species sharing the same habitat (i.e., excluding pairs not sharing the same habitat, because many differences in species traits can be caused by adaptation to a specific habitat type) and iii) all plants of the Czech Republic. Information about species traits was mainly obtained from literature and databases. Comparison with common closely related species showed that CR species are smaller, flower for shorter periods, and have higher proportions of self-compatibility and higher terminal velocities. CR species also differ in their mode of dispersion, and their ecology and distribution. Comparison with species from the same habitat gave similar results. Comparison with the whole flora produced slightly different results, with additional differences in pollination mode and seed mass. The results of all three types of comparison suggest that critically endangered species of the Czech Republic are small, competitively inferior species, with some differences in the generative part of their life cycle, and occur mainly in open, unproductive habitats.     

Book Review

Abstract

Most studies on population ecology of bryophytes have involved common species. However, some studies have compared life history strategies in rare and common species. We review the life history strategies (life expectancy, sexual and asexual reproduction, spore production, spore size and dispersal) for species that are rare in relation to pattern and persistency of suitable habitat patches. In particular, we discuss the dynamics on two levels, within and among localities, for different categories of rare species. We predict that most rare species will be found to have restricted dispersal capacities but higher than average life expectancies of local subpopulations. Natural rarity is distinguished from human-induced rarity and species rare for the latter reason are distinguished as 'threatened'.     

Abundance and habitat segregation in Mediterranean grassland species: the importance of seed weight

Abstract. We analysed the relationship between seed traits (weight, shape and dispersal structures) and the abundance and habitat segregation of Mediterranean grassland species. To take into account possible correlations with other plant traits, the study also includes 5 vegetative traits (growth form, plant longevity, clonality, onset of flowering and plant size) of commonly accepted functional importance. Data were recorded for 85 species from dehesa grasslands in central Spain. Species abundance was measured in upper (dry and less productive, high stress) and lower (moist and more productive, low stress) slope zones in the same area. Habitat segregation was estimated using an index based on the relative frequencies of species in upper and lower slope zones. Multiple regression models were fitted using species, as well as phylogenetically independent contrasts, as data points. Annual small‐seeded species without specialised dispersal structures are over‐represented in dehesa grasslands. Abundance was negatively related to seed weight in upper slope zones. None of the recorded plant traits were related to abundance in the lower slope zones. Habitat segregation was mainly related to seed weight, but also to some vegetative traits. Annual, early flowering and small‐seeded species were relatively more abundant in the upper than the lower slope zones. This pattern is independent of phylogeny. Our results suggest that in dry Mediterranean grasslands, abundance of many species is determined by dispersal (production of numerous small seeds) rather than by competitive ability.     

The reproductive biology of the Mediterranean endemic Cyclamen balearicum Willk. (Primulaceae)

Knowledge of the reproductive biology of endemic plants improves our understanding of how mating system may be related to patterns of species abundance and provides a basis for the development of rational conservation programmes. In this paper we present natural population data on the floral biology and reproductive ecology of the endemic Mediterranean species Cyclamen balearicum Willk. This is a long-lived, diploid perennial herb which occurs in southern France in five fragmented and isolated regions and on the Balearic Islands of Mallorca, Menorca, Ibiza, Cabrera and Draponera. Our observations indicate a particularly scarce pollinator activity (rare syrphid visits) and dispersal by ants over small distances. A controlled pollination experiment in a natural population showed that in southern France C. balearicum is fully self-compatible and that selfing is autonomous and probably delayed (i.e. following opportunities for outcrossing). The proximity of stigmas and anthers will favour autonomous selfing. The high pollen/ovule ratio indicates nevertheless that C. balearicum has a mixed mating system. Patterns of variation in stigma-anther separation and pollen production per flower suggest that not only has the current mating system of the species evolved from an outcrossing ancestor but that due to the fragmentation and isolation of populations greater levels of selfing have evolved in southern France (and to an intermediate degree on Ibiza and Menorca). On the island of Mallorca where larger continuous belts of C. balearicum habitat still exist the species has floral traits indicating a more outcrossed mating system. To our knowledge this is the first paper to document such trends in floral traits in the endemic component of the Mediterranean flora.     

Increasing habitat connectivity in agricultural landscapes as a weed management strategy reconciling ecology and agronomy

While studies have explored how habitat amount drives weed assemblages in agroecosystems, knowledge remains limited of the effects of habitat connectivity. The response-effect trait framework provides insights into the mechanisms underpinning the relationship between landscape structure and the taxonomic diversity and abundance of weed assemblages. This study evaluated how habitat connectivity and habitat amount affect weed diversity and abundance in winter cereal fields, and whether these effects are driven by the functional composition of weed assemblages. We sampled weeds in 27 winter cereal fields. We measured habitat connectivity and habitat amount provided by wooded, grassland and cropland elements. We selected five traits related to the dispersal, establishment, and competitive abilities of weed species likely to respond to landscape structure: seed number per plant, type of reproduction, seed dry mass, plant vegetative height and seed germination rate. The functional composition of weed assemblages was assessed using community weighted mean trait values. Weed diversity and abundance were used as proxies of weed management. The taxonomic approach did not reveal any effect of landscape structure on weed diversity and abundance. Only the grassland elements that contributed to habitat connectivity, and to a lesser extent to habitat amount, drove the functional composition of weed assemblages. High habitat amount favoured species with many seeds, while high habitat connectivity favoured species with fewer seeds, a higher ability to reproduce vegetatively and higher seed germination rates. In turn, higher seed germination rates increased weed evenness and reduced weed abundance. Some of these relationships were influenced by the presence of rare species. Overall, high connectivity provided by grassland elements increases weed evenness and reduces weed abundance by shaping weed functional composition. Our study suggests that land-use planning policies that enhance the connectivity provided by grassland elements could be considered as a weed management strategy reconciling ecology and agronomy.     

Interactions between alien plant species traits and habitat characteristics in agricultural landscapes in Finland

The survival and success of alien plant species is determined by species traits (i.e., invasiveness) and the characteristics of the habitats in the region of introduction (i.e., invasibility). However, little is known about species traits as related to habitat characteristics. We assessed the characteristics of successful invaders and the interaction of environmental factors and life-history traits for alien plant species. The vascular plants were recorded from 52 agricultural landscapes in Finland. We compared the traits of native and alien plant species with Fisher’s exact test and used a three table ordination analysis, RLQ analysis, to relate species traits to environmental conditions. Species were clustered according to their position on the RLQ axes, and the clusters were tested for phylogenetic independence. The successful alien plant species were associated with life form and preferences for moisture and nitrogen, but the trait composition varied according to the habitat type. Two RLQ axes explained 80.5% of the variation, and the species traits were significantly associated with environmental variables. The clustering showed that the occurrence of alien plant species in agricultural habitats was driven by invasion history, traits related to dispersal (dispersal type, seed mass) and habitat preferences, as well as environmental features, such as geographical location, temperature and the quality and disturbance regime of the habitats. All clusters were phylogenetically non-independent. Thus, the clusters of alien species comprised species of diverse taxonomic affinities, although, they shared the traits explaining their occurrence in particular habitats. This information is useful for understanding the link between species traits and the environmental conditions of the habitats, and complexity of the invasion process.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.     

Plant Trait-Species Abundance Relationships Vary with Environmental Properties in Subtropical Forests in Eastern China

Understanding how plant trait-species abundance relationships change with a range of single and multivariate environmental properties is crucial for explaining species abundance and rarity. In this study, the abundance of 94 woody plant species was examined and related to 15 plant leaf and wood traits at both local and landscape scales involving 31 plots in subtropical forests in eastern China. Further, plant trait-species abundance relationships were related to a range of single and multivariate (PCA axes) environmental properties such as air humidity, soil moisture content, soil temperature, soil pH, and soil organic matter, nitrogen (N) and phosphorus (P) contents. At the landscape scale, plant maximum height, and twig and stem wood densities were positively correlated, whereas mean leaf area (MLA), leaf N concentration (LN), and total leaf area per twig size (TLA) were negatively correlated with species abundance. At the plot scale, plant maximum height, leaf and twig dry matter contents, twig and stem wood densities were positively correlated, but MLA, specific leaf area, LN, leaf P concentration and TLA were negatively correlated with species abundance. Plant trait-species abundance relationships shifted over the range of seven single environmental properties and along multivariate environmental axes in a similar way. In conclusion, strong relationships between plant traits and species abundance existed among and within communities. Significant shifts in plant trait-species abundance relationships in a range of environmental properties suggest strong environmental filtering processes that influence species abundance and rarity in the studied subtropical forests.     

The impact of neutrality, niche differentiation and species input on diversity and abundance distributions

We present a spatially-explicit generalization of Hubbell's model of community dynamics in which the assumption of neutrality is relaxed by incorporating dispersal limitation and habitat preference. In simulations, diversity and species abundances were governed by the rate at which new species were introduced (usually called 'speciation') and nearly unaffected by dispersal limitation and habitat preference. Of course, in the absence of species input, diversity is maintained solely by niche differences. We conclude that the success of the neutral model in predicting the abundance distribution has nothing to do with neutrality, but rather with the species-introduction process: when new species enter a community regularly as singletons, the typical J-shaped abundance distribution, with a long tail of rare species, is always observed, whether species differ in habitat preferences or not. We suggest that many communities are indeed driven by the introduction process, accounting for high diversity and rarity, and that species differences may be largely irrelevant for either.     

Long‐term colonization ecology of forest‐dwelling species in a fragmented rural landscape – dispersal versus establishment

Species colonization in a new habitat patch is an efficiency indicator of biodiversity conservation. Colonization is a two‐step process of dispersal and establishment, characterized by the compatibility of plant traits with landscape structure and habitat conditions. Therefore, ecological trait profiling of specialist species is initially required to estimate the relative importance of colonization filters. Old planted parks best satisfy the criteria of a newly created and structurally matured habitat for forest‐dwelling plant species. We sampled species in 230 ancient deciduous forests (source habitat), 74 closed‐canopy manor parks (target habitats), 151 linear wooded habitats (landscape corridors), and 97 open habitats (isolating matrix) in Estonia. We defined two species groups of interest: forest (107 species) and corridor specialists (53 species). An extra group of open habitat specialists was extracted for trait scaling. Differing from expectations, forest specialists have high plasticity in reproduction mechanisms: smaller seeds, larger dispersules, complementary selfing ability, and diversity of dispersal vectors. Forest specialists are shorter, less nutrient‐demanding and mycorrhizal‐dependent, stress‐tolerant disturbance‐sensitive competitors, while corridor specialists are large‐seeded disturbance‐tolerant competitors. About 40% of species from local species pools have immigrated into parks. The historic forest area, establishment‐related traits, and stand quality enhance the colonization of forest specialists. The openness of landscape and mowing in the park facilitate corridor specialists. Species traits in parks vary between a forest and corridor specialist, except for earlier flowering and larger propagules. Forest species are not dispersal limited, but they continue to be limited by habitat properties even in the long term. Therefore, the shady parts of historic parks should be appreciated as important forest biodiversity‐enhancing landscape structures. The habitat quality of secondary stands can be improved by nurturing a heterogeneous shrub and tree layer, and modest herb layer management.     

You don’t belong here: explaining the excess of rare species in terms of habitat,space and time

Ecological communities are composed of a few common and several rare species. Many studies have evaluated the shape of abundance distribution curves, but few studies have assessed the causes of rarity. Using a dataset of stream macroinvertebrates, we investigated whether the excess of rare species in three focal communities of stones in riffles were common 1) in other habitats at the same stream site and period of sampling (environment), 2) in other stream sites in the same habitat and period of sampling (space), and 3) in other years in the same stream site and habitat (time). We observed that around 28% of the rare species were common in other habitats (environment), stream sites (space) or years (time). Among the three factors, rarity was mostly explained by habitat type, whereas a significant portion of the rare species in riffles were common in pools, submerged roots of terrestrial plants or in partially submerged moss patches. This result suggests that the presence in non‐optimum habitat is a strong determinant of the rarity observed in natural communities and most rare species are due to sampling artifacts or accidentally sampled transient species.     

The importance of rare species: a trait‐based assessment of rare species contributions to functional diversity and possible ecosystem function in tall‐grass prairies

The majority of species in ecosystems are rare, but the ecosystem consequences of losing rare species are poorly known. To understand how rare species may influence ecosystem functioning, this study quantifies the contribution of species based on their relative level of rarity to community functional diversity using a trait‐based approach. Given that rarity can be defined in several different ways, we use four different definitions of rarity: abundance (mean and maximum), geographic range, and habitat specificity. We find that rarer species contribute to functional diversity when rarity is defined by maximum abundance, geographic range, and habitat specificity. However, rarer species are functionally redundant when rarity is defined by mean abundance. Furthermore, when using abundance‐weighted analyses, we find that rare species typically contribute significantly less to functional diversity than common species due to their low abundances. These results suggest that rare species have the potential to play an important role in ecosystem functioning, either by offering novel contributions to functional diversity or via functional redundancy depending on how rare species are defined. Yet, these contributions are likely to be greatest if the abundance of rare species increases due to environmental change. We argue that given the paucity of data on rare species, understanding the contribution of rare species to community functional diversity is an important first step to understanding the potential role of rare species in ecosystem functioning.     

Local above‐ground persistence of vascular plants: Life‐history trade‐offs and environmental constraints

Questions: 1. Which plant traits and habitat characteristics best explain local above‐ground persistence of vascular plant species and 2. Is there a trade‐off between local above‐ground persistence and the ability for seed dispersal and below‐ground persistence in the soil seed bank? Locations: 845 long‐term permanent plots in terrestrial habitats across the Netherlands. Methods: We analysed the local above‐ground persistence of vascular plants in permanent plots (monitored once a year for ca. 16 year) with respect to functional traits and habitat preferences using survival statistics (Kaplan‐Meier analysis and Cox’ regression). These methods account for censored data and are rarely used in vegetation ecology. Results: Local above‐ground persistence is determined by both functional traits (especially the ability to form long‐lived clonal connections) and habitat preferences (especially nutrient requirements). Above‐ground persistence is negatively related to the ability for dispersal by wind and to the ability to accumulate a long‐term persistent soil seed bank (‘dispersal through time’) and is positively related to the ability for dispersal by water. Conclusions: Most species have a half‐life expectation over 15 years, which may contribute to time lags after changes in habitat quality or ‐configuration (‘extinction debt’). There is evidence for a trade‐off relationship between local above‐ground persistence and below‐ground seed persistence, while the relationship with dispersal in space is vector specific. The rate of species turnover increases with productivity.     

Effect of habitat area and isolation on plant trait distribution in European forests and grasslands

A number of studies show contrasting results in how plant species with specific life‐history strategies respond to fragmentation, but a general analysis on whether traits affect plant species occurrences in relation to habitat area and isolation has not been performed. We used published data from forests and grasslands in north‐central Europe to analyse if there are general patterns of sensitivity to isolation and dependency of area for species using three traits: life‐span, clonality, and seed weight. We show that a larger share of all forest species was affected by habitat isolation and area as compared to grassland species. Persistence‐related traits, life‐span and clonality, were associated to habitat area and the dispersal and recruitment related trait, seed weight, to isolation in both forest and grassland patches. Occurrence of clonal plant species decreased with habitat area, opposite to non‐clonal plant species, and long‐lived plant species decreased with grassland area. The directions of these responses partly challenge some earlier views, suggesting that further decrease in habitat area will lead to a change in plant species community composition, towards relatively fewer clonal and long‐lived plants with large seeds in small forest patches and fewer clonal plants with small seeds in small grassland patches. It is likely that this altered community has been reached in many fragmented European landscapes consisting of small and isolated natural and semi‐natural patches, where many non‐clonal and short‐lived species have already disappeared. Our study based on a large‐scale dataset reveals general and useful insights concerning area and isolation effects on plant species composition that can improve the outcome of conservation and restoration efforts of plant communities in rural landscapes.     

Inbreeding depresses short and long distance dispersal in three congeneric spiders

Dispersal is one of the most important precopulatory inbreeding avoidance mechanisms and subject to landscape related selection pressures. In small populations, inbreeding within and between populations may strongly affect population dynamics if it reduces fitness and gene‐flow. While inbreeding avoidance is generally considered to be a key evolutionary driver of dispersal, potential effects of inbreeding on the dispersal process, are poorly known. Here, I document how inbreeding within a population, so by mating among relatives, affects the survivorship and the dispersal behaviour of three congeneric spider Erigone species (Araneae: Linyphiidae) that differ in habitat preference and regional rarity. The three species were chosen as a model because they allow the assessment of both long and short distance dispersal motivation (respectively ballooning and rappelling) under laboratory conditions. Inbreeding reduced both long and short distance dispersal modes in the three congeneric species. Because survival was depressed after inbreeding, with a tendency of reduced survival loss in the rare and highly stenotopic species, energetic constraints are likely to be the underlying mechanism. Inbreeding consequently depresses silk‐related dispersal in three related spiders. This may induce an inbreeding depression vortex with important consequences for range expansion and metapopulation dynamics of aerially dispersing species from highly fragmented landscapes.     

Both traits and phylogenetic history influence community structure in snakes over steep environmental gradients

Assemblages of closely related organisms are generated on axes of deep time diversification, biogeographic processes related to dispersal and habitat filtering, and competition. Using models that account for phylogeny, ecology, and traits, we examine how the interaction among biogeography, habitat filtering, and trait convergence influences community assemblage in Nearctic snakes. With 122 community surveys, environmental niche, trait data including size, diet, parity and habitat preference, and a nearly complete phylogeny of snakes from the United States, we ask 1) do phylogenetic species variability (PSV) and traits change in predictable and correlated ways given ecology and geographic distance, 2) are the measured traits variable within and across communities and how is this related to PSV at local scales, and 3) is there evidence of habitat filtering or trait divergence? Following a general trend of western to eastern North American origin and dispersal of major groups, we similarly show a significant decrease in PSV in this direction but unexpectedly with stable trait variance, showing that traits and phylogenetic variability are disconnected at the community level. We also demonstrate that trait variability and not PSV dominates local communities. Finally, regardless of phylogeny, we show that certain traits, such as reproductive mode (parity) frequency, change within communities in response to steep ecological gradients.     

Both rare and common species make unique contributions to functional diversity in an ecosystem unaffected by human activities

Aim

Rare species typically contribute more to functional diversity than common species. However, humans have altered the occupancy and abundance patterns of many species—the basis upon which we define “rarity.” Here, we use a globally unique dataset from hydrothermal vents—an untouched ecosystem—to test whether rare species over‐contribute to functional diversity.

Location

Juan de Fuca Ridge hydrothermal vent fields, Northeast Pacific Ocean.

Methods

We first conduct a comprehensive review to set up expectations for the relative contributions of rare and common species to functional diversity. We then quantify the rarity and commonness of 37 vent species with relevant trait information to assess the relationship between rarity and functional distinctiveness—a measure of the uniqueness of the traits of a species relative to traits of coexisting species. Next, we randomly assemble communities to test whether rare species over‐contribute to functional diversity in artificial assemblages ranging in species richness. Then, we test whether biotic interactions influence functional diversity contributions by comparing the observed contribution of each species to a null expectation. Finally, we identify traits driving functional distinctiveness using a distance‐based redundancy analysis.

Results

Across functional diversity metrics and species richness levels, we find that both rare and common species can contribute functional uniqueness. Some species always offer unique trait combinations, and these species host bacterial symbionts and provide habitat complexity. Moreover, we find that contributions of species to functional diversity may be influenced by biotic interactions.

Main conclusions

Our findings show that many common species make persistent, unique contributions to functional diversity. Thus, it is key to consider whether the abundance and occupancy of species have been reduced, relative to historical baselines, when interpreting the contributions of rare species to functional diversity. Our work highlights the importance of testing ecological theory in ecosystems unaffected by human activities for the conservation of biodiversity.