Does habitat fragmentation reduce genetic diversity and subpopulation connectivity?

The estimation of levels of genetic variation has received considerable attention because it is generally thought to be indicative of overall species vitality and the potential for evolutionary responses to environmental changes. Here, we use allozymes markers and two distinct collections of Cakile maritima, an annual species from sandy coastal habitats (2000 generation and 2005 generation collected from 9 populations in their natural habitats), to assess the magnitude of expected genetic change. We compared genetic diversity between generations (all populations combined), and then between populations at each generation. Based on 13 loci scored from the eight enzymes examined, a high genetic diversity was detected at both the population and generation level as compared to other herbaceous species. However, allelic richness reduction in the 2005 generation suggested restricted gene flow and a high risk of future genetic bottlenecks, if larger tracts of coastal areas disappear. Most loci showed deviation from Hardy‐Weinberg equilibrium due to excess of heterozygotes in all populations suggesting that this species has an allogamic mode of reproduction. It appears most likely that this species has experienced a recent decrease in population size, and that genetic drift in small populations has resulted in a loss of alleles occurring at low frequency. Despite the deterioration process, maintenance of high genetic diversity suggests that there are some ecological factors determining population structure.     

What determines the relationship between plant diversity and habitat productivity?

The relationship between biodiversity and habitat productivity has been a fundamental topic in ecology. Although the relationship between these parameters may exhibit different shapes, the unimodal shape has been frequently encountered. The decrease in diversity at high productivity has usually been attributed to competitive exclusion. We suggest that evolutionary history and dispersal limitation may be even more important in shaping the diversity–productivity relationship. On a global scale, unimodal diversity–productivity relationships dominate in temperate regions, whereas positive relationships are more common in the tropics. This difference can be accounted for by contrasting evolutionary history. Temperate regions have smaller species pools for productive habitats since these habitats have been scarce historically for speciation, while the opposite is true for the tropics. In addition, dispersal within a region may limit diversity either due to the lack of dispersal syndromes at low productivity or the low number of diaspores at high productivity. Thereafter, biotic interactions (competition and facilitation) can shape the relationship. All these processes can act independently or concurrently. We recommend that the common approach to examining empirical diversity–environmental relationships should start with the role of large‐scale processes such as evolutionary history and dispersal limitation, followed by influences associated with ecological interactions.     

How does habitat diversity affect the species–area relationship?

Aim   To examine the way in which 'area' and 'habitat diversity' interact in shaping species richness and to find a simple and valid way to express this interaction.
Location   The Natura 2000 network of terrestrial protected areas in Greece, covering approximately 16% of the national territory.
Methods   We used the Natura 2000 framework, which provides a classification scheme for natural habitat types, to quantify habitat heterogeneity. We analysed data for the plant species composition in 16,143 quadrats in which 5044 species and subspecies of higher plants were recorded. We built a simple mathematical model that incorporates the effect of habitat diversity on the species–area relationship (SAR).
Results   Our analysis showed that habitat diversity was correlated with area. However, keeping habitat diversity constant, species richness was related to area; while keeping area constant, species richness was related to habitat diversity. Comparing the SAR of the 237 sites we found that the slope of the species–area curve was related to habitat diversity.
Main conclusions   Discussion of the causes of the SAR has often focused on the primacy of area per se versus habitat heterogeneity, even though the two mechanisms are not mutually exclusive and should be considered jointly. We find that increasing habitat diversity affects the SAR in different ways, but the dominant effect is to increase the slope of the SAR. While a full model fit typically includes a variety of terms involving both area and habitat richness, we find that the effect of habitat diversity can be reduced to a linear perturbation of the slope of the species accumulation curve.     

Does habitat heterogeneity affect the diversity of epigaeic arthropods in vineyards?

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Effects of habitat complexity on forest beetle diversity: do functional groups respond consistently?

We examined the responses of a beetle assemblage to habitat complexity differences within a single habitat type, Sydney sandstone ridgetop woodland, using pitfall and flight‐intercept trapping. Six habitat characters (tree canopy cover, shrub canopy cover, ground herb cover, soil moisture, amount of leaf litter, and amount of logs, rocks and debris) were scored between 0 and 3 using ordinal scales to reflect habitat complexity at survey sites. Pitfall trapped beetles were more species rich and of different composition in high complexity sites, compared with low complexity sites. Species from the Staphylinidae (Aleocharinae sp. 1 and sp. 2), Carabidae (Pamborus alternans Latreille), Corticariidae (Cartodere Thomson sp. 1) and Anobiidae (Mysticephala Ford sp. 1) were most clearly responsible for the compositional differences, preferring high complexity habitat. Affinities between general functional groupings of pitfall‐trapped beetles and habitat variables were not clear at a low taxonomic resolution (family level). The composition and species richness of flight‐intercept‐trapped beetles were similar in high and low complexity sites. Our study demonstrates that discrete responses of the various functional groups of beetles are strongly associated with their feeding habits, indicated by differing habitat components from within overall composite habitat complexity measures. Although habitat preferences by beetle species may often reflect their foraging habits, clarification of the causal mechanisms underpinning the relationships between habitat complexity and beetles are critical for the development of general principles linking habitat, functional roles and diversity.     

How does landscape context contribute to effects of habitat fragmentation on diversity and population density of butterflies?

Aim Studies on habitat fragmentation of insect communities mostly ignore the impact of the surrounding landscape matrix and treat all species equally. In our study, on habitat fragmentation and the importance of landscape context, we expected that habitat specialists are more affected by area and isolation, and habitat generalists more by landscape context. Location and methods The study was conducted in the vicinity of the city of Göttingen in Germany in the year 2000. We analysed butterfly communities by transect counts on thirty‐two calcareous grasslands differing in size (0.03–5.14 ha), isolation index (2100–86,000/edge‐to‐edge distance 55–1894 m), and landscape diversity (Shannon–Wiener: 0.09–1.56), which is correlated to percentage grassland in the landscape. Results A total of 15,185 butterfly specimens belonging to fifty‐four species are recorded. In multiple regression analysis, the number of habitat specialist (n = 20) and habitat generalist (n = 34) butterfly species increased with habitat area, but z‐values (slopes) of the species–area relationships for specialists (z = 0.399) were significantly steeper compared with generalists (z = 0.096). Generalists, but not specialists, showed a marginally significant increase with landscape diversity. Effects of landscape diversity were scale‐dependent and significant only at the smallest scale (landscape context within a 250 m radius around the habitat). Habitat isolation was not related to specialist and generalist species numbers. In multiple regression analysis the density of specialists increased significantly with habitat area, whereas generalist density increased only marginally. Habitat isolation and landscape diversity did not show any effects. Main conclusions Habitat area was the most important predictor of butterfly community structure and influenced habitat specialists more than habitat generalists. In contrast to our expectations, habitat isolation had no effect as most butterflies could cope with the degree of isolation in our study region. Landscape diversity appeared to be important for generalist butterflies only.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

The heterospecific habitat copying hypothesis: can competitors indicate habitat quality?

According to the "habitat copying" hypothesis, animals use thereproductive performance of conspecifics to assess habitat suitabilityand choose their future breeding site. This is because conspecificsshare ecological needs and thus indicate habitat suitability.Here, we propose the "heterospecific habitat copying" hypothesis,which states that animals should use public information (i.e.,information derived from the performance of others) from con-and heterospecifics sharing ecological needs. In a correlationalapproach we test some assumptions and predictions of this hypothesiswith a data set from two sympatric bird populations, rollers(Coracias garrulus) and kestrels (Falco tinnunculus), usingthe same nest-boxes and exploiting similar food resources. Sincekestrels are residents and breed earlier, we assumed that theyare dominant over rollers for nest-box acquisition. The environmentappears to be patchy for both species and temporally predictablefor kestrels only. Two results suggest that the use of heterospecificpublic information in breeding habitat selection may be at work:(1) an increase in the reoccupancy probability by kestrels ofprevious roller nests with increasing nest success, and (2)an increase in roller breeding population with increasing localkestrel success. Most of the other observed patterns could beexplained by alternative mechanisms such as natal philopatry,breeding fidelity, conspecific attraction, intraspecific habitatcopying, and the effect of interspecific competition.     

The intermediate disturbance hypothesis--species diversity or functional diversity?

Phytoplankton dynamics in a shallow eutrophic lake were investigatedover a 3-year period with respect to environmental forces whichdrive species composition and diversity. Diversity was calculatedon the basis of species as well as on the basis of their functionalproperties (the C-R-S concept). Stratification and water columnmixing had a strong impact on phytoplankton composition. Applicationof a similarity–diversity model revealed that a high diversitywas a transient non-stable state, whereas drastic changes orlong-lasting stable environmental conditions are characterizedby low diversity. This effect was more pronounced when the diversitywas calculated on the basis of the phytoplankton species' functionalproperties. Thus, this functional approach supports the intermediatedisturbance hypothesis from field data.     

Phylogeographic structure without pre-mating barriers: Do habitat fragmentation and low mobility preserve song and chorus diversity in a European bushcricket?

When animal species have a strong phylogeographic structure questions arise on the origin, maintenance and future evolutionary trajectory of that structure. One prediction is that phenotypic differences among populations serve as pre-mating barriers should secondary contact occur. Post-mating barriers may also function and ensure further separation of the populations. We tested these predictions in an acoustic insect, the European bushcricket Ephippiger diurnus, that is geographically distributed in separated, genetically isolated populations distinguished by pronounced differences in male songs and chorusing and female preferences for song. We staged mixed-population choruses to examine how males sing when in the company of other populations, and we released females in these choruses to assess their preference for or aversion toward males of their own population versus a different one. We also paired males and females of the same and different populations to test mating success, spermatophore transfer, and oviposition in the various pairings. In most cases males sang as effectively when accompanied by males from another population as when in a single-population chorus, and females overall expressed little aversion toward males from a different population. Inter-population pairs did not mate less frequently, transfer smaller spermatophores, or deposit fewer or smaller eggs than pairs of males and females from the same population. We infer that pre-mating barriers play little role in maintaining phylogeographic structure despite the acoustic differences between populations. Rather, the structure probably reflects the fragmented distribution of suitable habitat and the low mobility of both juveniles and adults. Thus, if secondary contact does occur, and we predict that it will in several key locations owing to ongoing environmental change, coalescence of populations and reduction of genetic and acoustic diversity may follow.     

Does higher connectivity lead to higher genetic diversity? Effects of habitat fragmentation on genetic variation and population structure in a gypsophile

Habitat fragmentation is a major threat to the maintenance of genetic diversity in many plant populations. Genetic effects of population size have received far more attention than the effects of isolation—or connectivity—but both are key components of the fragmentation process. To analyze the consequences of fragment size and connectivity on the neutral genetic variation and population genetic structure of the dominant gypsophile Lepidium subulatum, we selected 20 fragments along two continuous gradients of size and degree of isolation in a fragmented gypsum landscape of Central Spain. We used eight polymorphic microsatellite markers, and analyzed a total of 344 individuals. Populations were characterized by high levels of genetic diversity and low inbreeding coefficients, which agrees with the mainly outcrossing system of L. subulatum and its high abundance in gypsum landscapes. Bayesian clustering methods, pairwise F _ST values and analysis of molecular variance revealed low among-population differentiation, with no significant isolation by distance. However, several genetic diversity indices such as allelic richness, number of effective alleles, expected heterozygosity and number of private alleles were negatively related to population isolation. The higher genetic diversity found on more connected fragments suggests higher rates of gene flow among more connected populations. Overall, our results highlight that fragmentation can have important effects on intra-population genetic processes even for locally abundant, dominant species. This, together with previously documented effects of connectivity on fitness of gypsophile species highlights the importance of including habitat connectivity in management and conservation strategies of this type of semiarid systems.     

How do habitat amount and habitat fragmentation drive time-delayed responses of biodiversity to land-use change?

Land-use change is a root cause of the extinction crisis, but links between habitat change and biodiversity loss are not fully understood. While there is evidence that habitat loss is an important extinction driver, the relevance of habitat fragmentation remains debated. Moreover, while time delays of biodiversity responses to habitat transformation are well-documented, time-delayed effects have been ignored in the habitat loss versus fragmentation debate. Here, using a hierarchical Bayesian multi-species occupancy framework, we systematically tested for time-delayed responses of bird and mammal communities to habitat loss and to habitat fragmentation. We focused on the Argentine Chaco, where deforestation has been widespread recently. We used an extensive field dataset on birds and mammals, along with a time series of annual woodland maps from 1985 to 2016 covering recent and historical habitat transformations. Contemporary habitat amount explained bird and mammal occupancy better than past habitat amount. However, occupancy was affected more by the past rather than recent fragmentation, indicating a time-delayed response to fragmentation. Considering past landscape patterns is therefore crucial for understanding current biodiversity patterns. Not accounting for land-use history ignores the possibility of extinction debt and can thus obscure impacts of fragmentation, potentially explaining contrasting findings of habitat loss versus fragmentation studies.     

Does diversity beget diversity? A case study of crops and weeds

Abstract. The ecological literature is ambiguous as to whether the initial diversity of a plant community facilitates or deters the diversity of colonizing species. We experimentally planted annual crop species in monoculture and polyculture, and examined the resulting weed communities. The species composition of weeds was similar among treatments, but the species richness of weeds was significantly higher in the polycultures than in the monocultures. This supports the ‘diversity begets diversity’ hypothesis. Environmental microheterogeneity, diversity promoters, and ecological equivalency do not seem able to explain the observed patterns.     

Ecological filtering or random extinction? Beta‐diversity patterns and the importance of niche‐based and neutral processes following habitat loss

Although both niche‐based and neutral processes are involved in community assembly, most models on the effects of habitat loss are stochastic, assuming neutral communities mainly affected by ecological drift and random extinction. Given that habitat loss is considered the most important driver of the current biodiversity crisis, unraveling the processes underlying the effects of habitat loss is critical from both a theoretical and an applied perspective. Here we unveil the importance of niche‐based and neutral processes to species extinction and community assembly across a gradient of habitat loss, challenging the predictions of neutral models. We draw on a large dataset containing the distribution of 3653 individuals of 42 species, representing 35% of the small mammal species of the Atlantic Forest hotspot, obtained in 68 sites across three continuously‐forested landscapes and three adjacent 10 000‐ha fragmented landscapes differing in the amount of remaining forest (50%, 30% and 10%). By applying a null‐model approach, we investigated β‐diversity patterns by detecting deviations of observed community similarity from the similarity between randomly assembled communities. Species extinction following habitat loss was decidedly non‐random, in contrast to the notion that fragmented communities are mainly driven by ecological drift. Instead, habitat loss led to a strong biotic homogenization. Moreover, species composition changed abruptly at the same level of landscape‐scale habitat loss that has already been associated with a drastic decline in species richness. Habitat loss, as other anthropogenic disturbances, can thus be seen as a strong ecological filter that increases (rather than decreases) the importance of deterministic processes in community assembly. As such, critical advances for the development of conservation science lie on the incorporation of the relevant niche traits associated with extinction proneness into models of habitat loss. The results also underscore the fundamental importance of pro‐active measures to prevent human‐modified landscapes surpassing critical ecological thresholds.     

Does herbivore diversity depend on plant diversity? The case of California butterflies

It is widely believed that the diversity of plants influences the diversity of animals, and this should be particularly true of herbivores. We examine this supposition at a moderate spatial extent by comparing the richness patterns of the 217 butterfly species resident in California to those of plants, including all 5,902 vascular plant species and the 552 species known to be fed on by caterpillars. We also examine the relationships between plant/butterfly richness and 20 environmental variables. We found that although plant and butterfly diversities are positively correlated, multiple regression, path models, and spatial analysis indicate that once primary productivity (estimated by a water-energy variable, actual evapotranspiration) and topographical variability are incorporated into models, neither measure of plant richness has any relationship with butterfly richness. To examine whether butterflies with the most specialized diets follow the pattern found across all butterflies, we repeated the analyses for 37 species of strict monophages and their food plants and found that plant and butterfly richness were similarly weakly associated after incorporating the environmental variables. We condude that plant diversity does not directly influence butterfly diversity but that both are probably responding to similar environmental factors.     

Whither the diversity bargain?

In Natasha Warikoo’s account, the “diversity bargain” is widespread among white elite American college students. This bargain is tentative support for preferences for underrepresented minorities in college admissions, conditioned on the admitted minority students providing white students with multicultural experiences that signal elite cosmopolitanism. This essay reviews three possible explanations for the pervasiveness of the diversity bargain: campus experiences with the benefits of diversity; socialization into expectations that elites give lip service to the benefits of diversity; and Warikoo’s methodological and analytical choices.