Effects of species turnover on reserve site selection in a fragmented landscape

Changes in species composition is an important process in many ecosystems but rarely considered in systematic reserve site selection. To test the influence of temporal variability in species composition on the establishment of a reserve network, we compared network configurations based on species data of small mammals and frogs sampled during two consecutive years in a fragmented Atlantic Forest landscape (SE Brazil). Site selection with simulated annealing was carried out with the datasets of each single year and after merging the datasets of both years. Site selection resulted in remarkably divergent network configurations. Differences are reflected in both the identity of the selected fragments and in the amount of flexibility and irreplaceability in network configuration. Networks selected when data for both years were merged did not include all sites that were irreplaceable in one of the 2 years. Results of species number estimation revealed that significant changes in the composition of the species community occurred. Hence, temporal variability of community composition should be routinely tested and considered in systematic reserve site selection in dynamic systems.

Are there latitudinal gradients in species turnover?

Aim To examine the effect on the observed relationship betw een spatial turnover and latitude of both the measure of beta diversity used and the method of analysis.

Location The empirical analyses presented herein are for the New World.

Methods We take the spatial distributions of the owls of the New World as an exemplar data set to investigate the patterns of beta diversity across latitudes revealed by different analytical methods. To illustrate the strengths and weaknesses of alternative measures of beta diversity and different analytical approaches, we also use a simple random distribution model, focusing in particular on the influence of richness gradients and landmass geometry.

Results Our simple spatial model of turnover demonstrates that different combinations of analytical approach and measure of beta diversity can give rise to strikingly different relationships between turnover and latitude. The analyses of the bird data for the owls of the New World demonstrate that this observation extends to real data.

Conclusions For the particular assemblage considered, we present strong evidence that species richness declines at higher latitudes, and there is also some evidence that species turnover is greater nearer the equator, despite conceptual and practical difficulties involved in analysing spatial patterns of species turnover. We suggest some ways of overcoming these difficulties.

Temporal Diversity Patterns and Phenology in Fruit-feeding Butterflies in the Atlantic Forest

The Atlantic Forest deserves special attention due to its high level of species endemism and degree of threat. As in other tropical biomes, there is little information about the ecology of the organisms that occur there. The objectives of this study were to verify how fruit-feeding butterflies are distributed through time, and the relation with meteorological conditions. Species richness and Shannon index were partitioned additively at the monthly level, and beta diversity, used as a hierarchical measure of temporal species turnover, was calculated among months, trimesters, and semesters. Circular analysis was used to verify how butterflies are distributed along seasons and its relation with meteorological conditions. We sampled 6488 individuals of 73 species. Temporal diversity of butterflies was more grouped than expected by chance among the months of each trimester. Circular analyses revealed that diversity is concentrated in hot months (September-March), with the subfamily Brassolinae strongly concentrated in February-March. Average temperature was correlated with total abundance of butterflies, abundance of Biblidinae, Brassolinae and Morphinae, and richness of Satyrinae. The present results show that 3mo of sampling between September and March is enough to produce a nonbiased sample of the local assemblage of butterflies, containing at least 70 percent of the richness and 25 percent of abundance. The influence of temperature on sampling is probably related to butterfly physiology. Moreover, temperature affects resource availability for larvae and adults, which is higher in hot months. The difference in seasonality patterns among subfamilies is probably a consequence of different evolutionary pressures through time.

Climate and soil attributes determine plant species turnover in global drylands

Aim Geographical, climatic and soil factors are major drivers of plant beta diversity, but their importance for dryland plant communities is poorly known. The aim of this study was to: (1) characterize patterns of beta diversity in global drylands; (2) detect common environmental drivers of beta diversity; and (3) test for thresholds in environmental conditions driving potential shifts in plant species composition. Location Global. Methods Beta diversity was quantified in 224 dryland plant communities from 22 geographical regions on all continents except Antarctica using four complementary measures: the percentage of singletons (species occurring at only one site); Whittaker's beta diversity, β(W); a directional beta diversity metric based on the correlation in species occurrences among spatially contiguous sites, β(R2); and a multivariate abundance-based metric, β(MV). We used linear modelling to quantify the relationships between these metrics of beta diversity and geographical, climatic and soil variables. Results Soil fertility and variability in temperature and rainfall, and to a lesser extent latitude, were the most important environmental predictors of beta diversity. Metrics related to species identity percentage of singletons and β(W) were most sensitive to soil fertility, whereas those metrics related to environmental gradients and abundance (β(R2) and β(MV) were more associated with climate variability. Interactions among soil variables, climatic factors and plant cover were not important determinants of beta diversity. Sites receiving less than 178 mm of annual rainfall differed sharply in species composition from more mesic sites (> 200 mm). Main conclusions Soil fertility and variability in temperature and rainfall are the most important environmental predictors of variation in plant beta diversity in global drylands. Our results suggest that those sites annually receiving c. 178 mm of rainfall will be especially sensitive to future climate changes. These findings may help to define appropriate conservation strategies for mitigating effects of climate change on dryland vegetation.

Temporal variation in saproxylic beetle assemblages in a Mediterranean ecosystem

One of the main challenges in biological conservation has been to understand species distribution across space and time. Over the last decades, many diversity and conservation surveys have been conducted that have revealed that habitat heterogeneity acts as a major factor that determines saproxylic assemblages. However, temporal dynamics have been poorly studied, especially in Mediterranean forests. We analyzed saproxylic beetle distribution at inter and intra-annual scales in a “dehesa” ecosystem, which is a traditional Iberian agrosilvopastoral ecosystem that is characterized by the presence of old and scattered trees that dominate the landscape. Significant differences in effective numbers of families/species and species richness were found at the inter-annual scale, but this was not the case for composition. Temperature and relative humidity did not explain these changes which were mainly due to the presence of rare species. At the intra-annual scale, significant differences in the effective numbers of families/species, species richness and composition between seasons were found, and diversity partitioning revealed that season contributed significantly to gamma-diversity. Saproxylic beetle assemblages exhibited a marked seasonality in richness but not in abundance, with two peaks of activity, the highest between May and June, and the second between September and October. This pattern is mainly driven by the seasonality of the climate in the Mediterranean region, which influences ecosystem dynamics and imposes a marked seasonality on insect assemblages. An extended sampling period over different seasons allowed an overview of saproxylic dynamics, and revealed which families/species were restricted to particular seasons. Recognizing that seasons act as a driver in modelling saproxylic beetle assemblages might be a valuable tool in monitoring and for conservation strategies in Mediterranean forests.

Variability in north temperate dung beetle assemblages at different spatial and temporal scales

This study examined the spatial and temporal variability of dung beetle assemblages across a variety of scales e.g. from the between-pad scale (examining the effects of dung size and type) to larger spatial scales encompassing southern Ireland. Dung beetle assemblage structure as sampled by dung pad cohort samples and dung baited pitfall trapping were compared. Generally, the rank order of abundance of dung beetle species was significantly correlated between pitfall catches and cohort pad samples. Across different dung sizes, in both pitfall catches and cohort pad samples, the relative abundance of species was frequently significantly different, but the rank order of abundance of dung beetle was usually significantly correlated. Considerable variations in pitfall catches at temporal scales of a few days appeared to be closely related to weather conditions and rotational grazing. However, despite considerable variation in absolute abundances between consecutive days of sampling, assemblage structure typically remained very similar. The relationship between dung pad size and dung beetle colonisation was investigated. In field experiments in which pads of different sizes (0.25 L, 0.5 L, 1.0 L and 1.5 L) were artificially deposited, there was a positive relationship between pad size and both biomass and number of beetles colonising dung pads and pitfall traps. In addition, with one exception, the field experiments indicated a general positive relationship between dung pad size and biomass density (dung beetle biomass per unit dung volume). A laboratory experiment indicated that pat residence times of A. rufipes were significantly correlated with dung pad size. Investigation of naturally-deposited cow dung pads in the field also indicated that both larval numbers and densities were significantly correlated with dung pad size. These results were discussed in the context of theory related to aggregation and coexistence of species, and resource utilisation by organisms in ephemeral, patchy resources. The colonisation by dung beetles of dung types from native herbivores (sheep, horse and cow) was investigated in field experiments. There were significant differences between the dung types in the chemical parameters measured, and there were significant differences in abundances of dung beetles colonising the dung types. Sheep dung was typically the preferred dung type. Data from these field experiments, and from published literature, indicated that dung beetle species can display dung type preferences, in terms of comparisons of both absolute and relative abundances. In addition, data from laboratory experiments indicate that both Aphodius larval production and pat residence times tended to be higher in those dung types which were preferred by adult Aphodius in the colonisation experiments. Data from dung-baited pitfall trapping (from this and another study) at several sites (up to 180 km distant) and over a number of years (between 1991 and 1996) were used to investigate spatial and temporal variation in dung beetle assemblage structure and composition (Aphodius, Sphaeridium and Geotrupes) across a range of scales in southern Ireland. Species richness levels, species composition and rank order of abundances were very similar between the assemblages. The temporal variability between seasons within any year exceeded temporal variability between years. DCA ordinations indicated that there was a similar level of variability between assemblage structure from the between-field (~1km) to regional (~180 km) spatial scales, and between year (6 years) temporal scales. At the biogeographical spatial scale, analysis of data from the literature indicated that there was considerable variability at this scale, largely due to species turnover.

Habitat Isolation Reduces the Temporal Stability of Island Ecosystems in the Face of Flood Disturbance

Natural ecosystems are increasingly exposed to multiple anthropogenic stressors, including land-use change, deforestation, agricultural intensification, and urbanisation, all of which have led to widespread habitat fragmentation, which is also likely to be amplified further by predicted climate change. The potential interactive effects of these different stressors cannot be determined by studying each in isolation, although such synergies have been largely ignored in ecological field studies to date. Here, we use a model system of naturally fragmented islands in a braided river network, which is exposed to periodic inundation, to investigate the interactive effects of habitat isolation and flood disturbance. Food web structure was similar across the islands during periods of hydrological stability, but several key properties were altered in the aftermath of flood disturbance, based on distance of the islands from the regional source pool of species: taxon richness and mean food chain length declined with habitat isolation after flooding, while the proportion of basal species increased. Greater species turnover through time reflected the slower process of re-colonisation on the more distant islands following disturbance. Increased variability of several food web properties over a 1-year period highlighted the reduced temporal stability of isolated habitat fragments. Many of these effects reflected the differential successes of predator and prey species at re-colonising the islands: even though larger, more mobile consumers may reach the more distant islands first, they cannot establish populations until the lower trophic levels have successfully reassembled. These results highlight the susceptibility of fragmented ecosystems to environmental perturbations. © 2013 Elsevier Ltd.

Spatio-temporal distribution of calling male frogs in Morro do Diabo State Park (Southeastern Brazil): implications for conservation and management in a seasonally dry tropical forest

The aims of this study are to determine how anuran calling males are spatially and temporally distributed in the Morro do Diabo State Park (MDSP), and to test for indicator species in different breeding habitats. We found high species turnover among breeding habitats of different hydroperiods. Our results indicate that spatial partitioning is more important than seasonal occurrence in permitting species coexistence at MDSP. Indicator Species Analysis identified one species associated with a local stream, one species associated with permanent ponds, two species associated with temporary ponds, and three species associated with a semipermanent pond. Guilds of species do not appear to stratify their distributions throughout the year, since most species occur during the rainy and warm months and are thus temporally overlapping. The main conservation implication of our work is the need to protect the different areas of the park where these unique breeding habitats, and their associated anuran assemblages, are found.

Species interactions and random dispersal rather than habitat filtering drive community assembly during early plant succession

Theory on plant succession predicts a temporal increase in the complexity of spatial community structure and of competitive interactions: initially random occurrences of early colonising species shift towards spatially and competitively structured plant associations in later successional stages. Here we use long-term data on early plant succession in a German post mining area to disentangle the importance of random colonisation, habitat filtering, and competition on the temporal and spatial development of plant community structure. We used species co-occurrence analysis and a recently developed method for assessing competitive strength and hierarchies (transitive versus intransitive competitive orders) in multispecies communities. We found that species turnover decreased through time within interaction neighbourhoods, but increased through time outside interaction neighbourhoods. Successional change did not lead to modular community structure. After accounting for species richness effects, the strength of competitive interactions and the proportion of transitive competitive hierarchies increased through time. Although effects of habitat filtering were weak, random colonization and subsequent competitive interactions had strong effects on community structure. Because competitive strength and transitivity were poorly correlated with soil characteristics, there was little evidence for context dependent competitive strength associated with intransitive competitive hierarchies.

Spatial ecology of food webs : herbivore-parasitoid communities on the pedunculate oak

Herbivorous insects, their host plants and natural enemies form the largest and most species-rich communities on earth. But what forces structure such communities? Do they represent random collections of species, or are they assembled by given rules? To address these questions, food webs offer excellent tools. As a result of their versatile information content, such webs have become the focus of intensive research over the last few decades. In this thesis, I study herbivore-parasitoid food webs from a new perspective: I construct multiple, quantitative food webs in a spatially explicit setting, at two different scales. Focusing on food webs consisting of specialist herbivores and their natural enemies on the pedunculate oak, Quercus robur, I examine consistency in food web structure across space and time, and how landscape context affects this structure. As an important methodological development, I use DNA barcoding to resolve potential cryptic species in the food webs, and to examine their effect on food web structure. I find that DNA barcoding changes our perception of species identity for as many as a third of the individuals, by reducing misidentifications and by resolving several cryptic species. In terms of the variation detected in food web structure, I find surprising consistency in both space and time. From a spatial perspective, landscape context leaves no detectable imprint on food web structure, while species richness declines significantly with decreasing connectivity. From a temporal perspective, food web structure remains predictable from year to year, despite considerable species turnover in local communities. The rate of such turnover varies between guilds and species within guilds. The factors best explaining these observations are abundant and common species, which have a quantitatively dominant imprint on overall structure, and suffer the lowest turnover. By contrast, rare species with little impact on food web structure exhibit the highest turnover rates. These patterns reveal important limitations of modern metrics of quantitative food web structure. While they accurately describe the overall topology of the web and its most significant interactions, they are disproportionately affected by species with given traits, and insensitive to the specific identity of species. As rare species have been shown to be important for food web stability, metrics depicting quantitative food web structure should then not be used as the sole descriptors of communities in a changing world. To detect and resolve the versatile imprint of global environmental change, one should rather use these metrics as one tool among several.

Distribuição das assembleias de girinos associadas a riachos da Reserva Ecológica de Guapiaçu, Cachoeiras de Macacú, Rio de Janeiro, Brasil

Os girinos são organismos diversos e abundantes nos pequenos riachos de cabeceira de florestas tropicais e constituem importantes componentes da diversidade biológica, da trófica e funcional dos sistemas aquáticos. Diferentes características estruturais e limnológicas dos ambientes aquáticos influenciam a organização das assembleias de girinos. Embora o estágio larvar dos anuros seja o mais vulnerável de seu ciclo de vida, sujeito a elevadas taxas de mortalidade, as pesquisas sobre girinos na região neotropical ainda são pouco representativas diante da elevada diversidade de anfíbios desta região e ferramentas que permitam a sua identificação ainda são escassas. Nesta tese, dividida em três capítulos, apresento uma compilação das informações relacionadas aos principais fatores que afetam as assembleias de girinos na região tropical (Capítulo 1), a caracterização morfológica dos girinos encontrados nos riachos durante o estudo e uma proposta de chave dicotômica de identificação (Capítulo 2) e avalio a importância relativa da posição geográfica e da variação temporal de fatores ambientais locais sobre as assembleias de girinos, assim como a correlação entre as espécies de girinos e as variáveis ambientais de 10 riachos, ao longo de 15 meses, nas florestas da REGUA (Capítulo 3). Há pelo menos oito tendências relacionadas à distribuição das assembleias de girinos: (1) o tamanho dos riachos e a diversidade de microhabitats são importantes características abióticas influenciando a riqueza e a composição de espécies; (2) em poças, o gradiente de permanência (e.g., hidroperíodo) e a heterogeneidade do habitat são os principais fatores moldando as assembleias de girinos; (3) a composição de espécies parece ser um parâmetro das assembleias mais relevante do que a riqueza de espécies e deve ser primeiramente considerado durante o planejamento de ações conservacionistas de anuros associados a poças e riachos; (4) a predação parece ser a interação biótica mais importante na estruturação das assembleias de girinos, com predadores vertebrados (e.g. peixes) sendo mais vorazes em habitats permanentes e predadores invertebrados (e.g. larvas de odonata) sendo mais vorazes em ambientes temporários; (5) os girinos podem exercer um efeito regulatório, predando ovos e girinos recém eclodidos; (6) o uso do microhabitat varia em função da escolha do habitat reprodutivo pelos adultos, presença de predadores, filogenia, estágio de desenvolvimento e heterogeneidade do habitat; (7) os fatores históricos restringem os habitats reprodutivos que uma espécie utiliza, impondo restrições comportamentais e fisiológicas; (8) a variação temporal nos fatores bióticos (e.g., fatores de risco), abióticos (e.g., distribuição de chuvas), e no padrão de reprodução das espécies pode interferir na estrutura das assembleias de girinos tropicais. A variação temporal na heterogeneidade ambiental dos riachos da REGUA resultou na previsibilidade das assembleias locais de girinos, sendo que os parâmetros ambientais explicaram 23% da variação na sua composição. Os parâmetros espaciais explicaram uma porção menor da variação nas assembleias (16%), enquanto uma porção relativamente elevada da variação temporal da heterogeneidade ambiental foi espacialmente estruturada (18%). As variáveis abióticas que apresentaram as maiores correlação com a composição das assembleias de girinos foram a proporção de folhiço e de rochas no fundo do riacho, e secundariamente a profundidade, a condutividade e a temperatura. O gradiente gerado pela proporção de folhiço e de rochas representou a transição entre riachos permanentes e intermitentes. Este gradiente proporcionou o turnover de espécies, o qual também seguiu um gradiente de condutividade, temperatura, profundidade, e em menor extensão, de hidroperíodo e largura, que estiveram fortemente associado ao grau de permanência dos riachos. Estes resultados corroboram tanto a hipótese do controle ambiental, como do controle biótico de comunidades e indicam que a variação temporal da heterogeneidade ambiental e a variação na posição geográfica são importantes para a estruturação local de assembleias de girinos da REGUA. Os resultados também permitiram distinguir entre assembleias de girinos exclusivas de riachos permanentes, exclusivas de riachos intermitentes e aquelas registradas nos dois tipos de riachos. Os resultados deste capítulo são relevantes para compreender em que extensão os efeitos da variação temporal na heterogeneidade ambiental e de processos espaciais afetam localmente a estruturação de assembleias de girinos.

A puzzle with many pieces: the genetic structure and diversity of Phaeocystis antarctica Karsten (Prymnesiophyta)

Strong ocean current systems characterize the Southern Ocean. The genetic structure of marine phytoplankton species is believed to depend mainly on currents. Genetic estimates of the relatedness of populations of phytoplankton species therefore should provide a proxy showing to what extent different geographic regions are interconnected by the ocean current systems. In this study, spatial and temporal patterns of genetic diversity were studied in the circumpolar prymnesiophyte Phaeocystis antarctica Karsten using seven nuclear microsatellite loci. Analyses were conducted for 86 P. antarctica isolates sampled around the Antarctic continent between 1982 and2007. The resultsrevealed highgenetic diversity without singlegenotypes recurringeven amongisolateswithin a bloom or originating from the same bucket of water. Populations of P. antarctica were significantly differentiated among the oceanic regions. However, some geographically distant populations were more closely related to each other than they were to other geographically close populations. Temporal haplotype turnover within regions was also suggested by the multilocus fingerprints. Our data suggest that even within blooms of P. antarctica genetic diversity and population sizes are large but exchange between different regions canbe limited. Positive and significant inbreeding coefficients hint at further regional substructure of populations, suggestingthat patches, once isolated from one another, may not reconnect. These data emphasize that even for planktonic species in a marine ecosystem that is influenced by strong currents, significant breaks in geneflow may occur.

Modelling the future biogeography of North Atlantic zooplankton communities in response to climate change

Advances in habitat and climate modelling allow us to reduce uncertainties of climate change impacts on species distribution. We evaluated the impacts of future climate change on community structure, diversity, distribution and phenology of 14 copepod species in the North Atlantic. We developed and validated habitat models for key zooplankton species using continuous plankton recorder (CPR) survey data collected at mid latitudes of the North Atlantic. Generalized additive models (GAMs) were applied to relate the occurrence of species to environmental variables. Models were projected to future (2080–2099) environmental conditions using coupled hydroclimatix–biogeochemical models under the Intergovernmental Panel on Climate Change (IPCC) A1B climate scenario, and compared to present (2001–2020) conditions. Our projections indicated that the copepod community is expected to respond substantially to climate change: a mean poleward latitudinal shift of 8.7 km per decade for the overall community with an important species range variation (–15 to 18 km per decade); the species seasonal peak is expected to occur 12–13 d earlier for Calanus finmarchicus and C. hyperboreus; and important changes in community structure are also expected (high species turnover of 43–79% south of the Oceanic Polar Front). The impacts of the change expected by the end of the century under IPCC global warming scenarios on copepods highlight poleward shifts, earlier seasonal peak and changes in biodiversity spatial patterns that might lead to alterations of the future North Atlantic pelagic ecosystem. Our model and projections are supported by a temporal validation undertaken using the North Atlantic climate regime shift that occurred in the 1980s: the habitat model built in the cold period (1970–1986) has been validated in the warm period (1987–2004).

The geographical structure of British bird distributions:diversity, spatial turnover and scale

1. Using data on the spatial distribution of the British avifauna, we address three basic questions about the spatial structure of assemblages: (i) Is there a relationship between species richness (alpha diversity) and spatial turnover of species (beta diversity)? (ii) Do high richness locations have fewer species in common with neighbouring areas than low richness locations?, and (iii) Are any such relationships contingent on spatial scale (resolution or quadrat area), and do they reflect the operation of a particular kind of species-area relationship (SAR)?

2. For all measures of spatial turnover, we found a negative relationship with species richness. This held across all scales, with the exception of turnover measured as beta (sim).

3. Higher richness areas were found to have more species in common with neighbouring areas.

4. The logarithmic SAR fitted better than the power SAR overall, and fitted significantly better in areas with low richness and high turnover.

5. Spatial patterns of both turnover and richness vary with scale. The finest scale richness pattern (10 km) and the coarse scale richness pattern (90 km) are statistically unrelated. The same is true of the turnover patterns.

6. With coarsening scale, locations of the most species-rich quadrats move north. This observed sensitivity of richness 'hotspot' location to spatial scale has implications for conservation biology, e.g. the location of a reserve selected on the basis of maximum richness may change considerably with reserve size or scale of analysis.

7. Average turnover measured using indices declined with coarsening scale, but the average number of species gained or lost between neighbouring quadrats was essentially scale invariant at 10-13 species, despite mean richness rising from 80 to 146 species (across an 81-fold area increase). We show that this kind of scale invariance is consistent with the logarithmic SAR.

Invasive carnivores alter ecological function and enhance complementarity in scavenger assemblages on ocean beaches

Species composition is expected to alter ecological function in assemblages if species traits differ strongly. Such effects are often large and persistent for nonnative carnivores invading islands. Alternatively, high similarity in traits within assemblages creates a degree of functional redundancy in ecosystems. Here we tested whether species turnover results in functional ecological equivalence or complementarity, and whether invasive carnivores on islands significantly alter such ecological function. The model system consisted of vertebrate scavengers (dominated by raptors) foraging on animal carcasses on ocean beaches on two Australian islands, one with and one without invasive red foxes (Vulpes vulpes). Partitioning of scavenging events among species, carcass removal rates, and detection speeds were quantified using camera traps baited with fish carcasses at the dune–beach interface. Complete segregation of temporal foraging niches between mammals (nocturnal) and birds (diurnal) reflects complementarity in carrion utilization. Conversely, functional redundancy exists within the bird guild where several species of raptors dominate carrion removal in a broadly similar way. As predicted, effects of red foxes were large. They substantially changed the nature and rate of the scavenging process in the system: (1) foxes consumed over half (55%) of all carrion available at night, compared with negligible mammalian foraging at night on the fox-free island, and (2) significant shifts in the composition of the scavenger assemblages consuming beach-cast carrion are the consequence of fox invasion at one island. Arguably, in the absence of other mammalian apex predators, the addition of red foxes creates a new dimension of functional complementarity in beach food webs. However, this functional complementarity added by foxes is neither benign nor neutral, as marine carrion subsidies to coastal red fox populations are likely to facilitate their persistence as exotic carnivores.