Species‐habitat associations change in a subtropical forest of China

Question: Do tree and shrub species in an evergreen broadleaf forest show similar habitat associations across different life stages? Location: A 24‐ha evergreen broadleaf forest plot in a heterogeneous landscape in Gutianshan National Nature Reserve, Zhejiang Province, Eastern China. Methods: Species having positive associations with four habitat types (low valley, low ridge, upper valley, and upper ridge) at three life stages (sapling, juvenile, and mature stages) were compared for 60 tree and shrub species using torus‐translation tests. Results: A total of 117 significant positive associations with the four habitats were observed at the three life stages (43, 41, and 33 at the sapling, juvenile, and mature stages, respectively). For the 52 species significantly associated with habitats, only 16 were associated with the same habitat across all three stages. The majority of associated species at the juvenile stage (34 out of 40) were associated with the same habitat at their sapling stage, whereas half of species at the mature stage had consistent associations with the same habitat at their sapling stage. More species were associated with the upper ridge at the sapling and juvenile stages compared to the mature stage. Conversely, more species were associated with the low valley at the mature stage compared to the sapling and juvenile stages. Conclusions: Our results indicate that species ecological habitat associations can differ between developmental stages beyond 1 cm DBH, as most species habitat preferences were consistent from the sapling stage to the juvenile stage but changed at the mature stage.     

Species–topography association in a species-rich subtropical forest of China

Habitat specialization has been considered as a primary factor in determining the distribution of species. In this study, we investigated species–habitat associations while controlling for spatial neighbourhood effects in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. Habitat specialization was measured by topographic variation and its effects on species distributions were modelled at three different spatial scales (10×10, 20×20 and 25×25 m²) using log-linear regression models and randomization tests. Our results showed: (1) 83% of the species were related to at least one or more topographic variables. Among them, 66%, 60%, 65% and 70% were closely dependent on slope, aspect, elevation and convexity, respectively. (2) Topographic variables have much stronger non-linear (quadratic and cubic) effects on species distributions than linear effects. (3) The effects of topographic heterogeneity on the distribution of shrubs species are smaller than on the distribution of canopy species, and smaller effects were also found in less abundant species. (4) There was a strong neighbourhood effect on species distribution: In 85% of the species, abundance in a focal quadrat was significantly correlated with abundance in the neighbour quadrats. We conclude that habitat specialization plays an important role in maintaining the diversity of this species-rich subtropical forest.     

Species’ thermal preferences affect forest bird communities along landscape and local scale habitat gradients

The spatial distributions of species, and the resulting composition of local communities, are shaped by a complex interplay between species’ climatic and habitat preferences. We investigated this interaction by analyzing how the climatic niches of bird species within given communities (measured as a community thermal index, CTI) are related to vegetation structure. Using 3129 bird communities from the French Breeding Bird Survey and an information theoretic multimodel inference framework, we assessed patterns of CTI variation along landscape scale gradients of forest cover and configuration. We then tested whether the CTI varies along local scale gradients of forest structure and composition using a detailed data set of 659 communities from six forests located in northwestern France. At landscape scale, CTI values decreased with increasing forest cover, indicating that bird communities were increasingly dominated by cold‐dwelling species. This tendency was strongest at low latitudes and in landscapes dominated by unfragmented forest. At local scale, CTI values were higher in mature deciduous stands than in conifer or early stage deciduous stands, and they decreased consistently with distance from the edge of forest. These trends underpin the assertion that species’ habitat use along forest gradients is linked with their climatic niche, although it remains unclear to what extent it is a direct consequence of microclimatic variation among habitats, or a reflection of macroscale correlations between species’ thermal preferences and their habitat choice. Moreover, our results highlight the need to address issues of scale in determining how habitat and climate interact to drive the spatial distribution of species. This will be a crucial step towards accurate predictions of changes in the composition and dynamics of bird communities under global warming.     

Species–area curve and distance–decay relationships indicate habitat thresholds of ectomycorrhizal fungi in an old‐growth Pseudotsuga menziesii landscape

Aim

Ectomycorrhizal fungi (EMF) are a diverse and essential biota of forests that are vulnerable to species loss through reductions in late‐seral habitat. We examined how the spatial ecology of this biota, particularly distance–decay and species–area relationships, could better inform habitat thresholds for EMF conservation planning.

Location

Southeast Vancouver Island near Victoria, British Columbia, Canada.

Methods

Using a stratified sampling design, 11 plots (0.15 ha in size) were established at 0.05–17.5 km apart across 2,800 ha of mesic old‐growth Pseudotsuga menziesii var. menziesii and Tsuga heterophylla forests. EMF communities were compiled through molecular analysis of root tips and sporocarps.

Results

The EMF community was comprised of many Cortinarius, Piloderma, Russula and Tricholoma species typical of mesotrophic habitat. A total of 238 EMF species were observed, of which 86 species were detected only once. The ratio of average species richness per plot (84 taxa) to total richness was low at 0.35, and inherent stochasticity of the EMF community was estimated to be 31% community dissimilarity for species incidence. Distance decay of EMF communities was nonlinear, with an estimated slope break at 2.6 km, followed by a largely unchanging trend in β‐diversity. Accumulated species–area curves were fitted best by the cumulative Weibull sigmoid model, and the asymptote (367 species) at approx. 50 ha was consistent with nonparametric estimates of γ‐diversity (342–362 spp.).

Main conclusions

Old‐growth forests host an impressive amount of EMF diversity, and many of the Ramaria, Inocybe and Russula species are likely to be endemic to the Pacific Northwest. Both niche‐ and neutral‐based processes influenced EMF community composition, resulting in a minimum threshold of 50 ha (1.8% of the sample area) for capturing γ‐diversity. These spatial patterns will help design and evaluate conservation efforts, such as retention forestry, to sustain fully diverse EMF communities over managed landscapes.

     

Species‐specific stomatal response of trees to drought – a link to vegetation dynamics?

Question: Is stomatal regulation specific for climate and tree species, and does it reveal species‐specific responses to drought? Is there a link to vegetation dynamics? Location: Dry inner alpine valley, Switzerland Methods: Stomatal aperture (θ_E) of Pinus sylvestris, Quercus pubescens, Juniperus communis and Picea abies were continuously estimated by the ratio of measured branch sap flow rates to potential transpiration rates (adapted Penman‐Monteith single leaf approach) at 10‐min intervals over four seasons. Results: θ_E proved to be specific for climate and species and revealed distinctly different drought responses: Pinus stomata close disproportionately more than neighbouring species under dry conditions, but has a higher θ_E than the other species when weather was relatively wet and cool. Quercus keeps stomata more open under drought stress but has a lower θ_E under humid conditions. Juniperus was most drought‐tolerant, whereas Picea stomata close almost completely during summer. Conclusions: The distinct microclimatic preferences of the four tree species in terms of θ_E strongly suggest that climate (change) is altering tree physiological performances and thus species‐specific competitiveness. Picea and Pinus currently live at the physiological limit of their ability to withstand increasing temperature and drought intensities at the sites investigated, whereas Quercus and Juniperus perform distinctly better. This corresponds, at least partially, with regional vegetation dynamics: Pinus has strongly declined, whereas Quercus has significantly increased in abundance in the past 30 years. We conclude that θ_E provides an indication of a species' ability to cope with current and predicted climate.     

Species–area relationship within benthic habitat patches of a tropical floodplain river: An experimental test

The curvilinear relationship between species richness and habitat area (species–area relationship (SAR)) is a fundamental ecological pattern. The relationship is often viewed from a long‐term perspective across relatively large spatial scales, reflecting a balance between immigration and extinction dynamics. We explored whether predictions of SAR also manifest over short time periods (days) in benthic habitat patches of a dynamic floodplain river where littoral faunal assemblages are continuously assembled and disassembled with changing water levels. We examined the relationship of patch size with faunal abundance (i.e. fish and aquatic invertebrates), taxonomic richness, trophic group richness and overall assemblage composition. Strong taxa–area relationships emerged despite the relatively short experimental time period (21 days); larger patches had more taxa and trophic groups. For the smallest patches, taxonomic richness was especially sensitive to abundance of individuals; abundance of individuals was a less important predictor of taxonomic and trophic group richness for the largest patches. Despite the relatively short time frame for study within this temporally dynamic ecosystem, our findings indicate a strong SAR for fishes and macroinvertebrates inhabiting patchy habitats in the littoral zone of this tropical river.     

Habitat associations of Sterculiaceae trees in a Bornean rain forest plot

Questions: 1. Are trees in a Bornean tropical rain forest associated with a particular habitat? 2. Does the strength of habitat association with the species‐specific optimal habitat increase with tree size? Location : A 52‐ha plot in a mixed dipterocarp forest in a heterogeneous landscape at the Lambir Hills National Park, Sarawak, East Malaysia. Methods : Ten species from the Sterculiaceae were chosen as representative of all species in the plot, on the assumption that competition among closely related species is more stringent than that among more distantly related taxa. Their habitat associations were tested using data from a 52‐ha plot by a torus‐translation test. Results : The torus‐translation test showed that eight out of the ten species examined had significant association with at least one habitat. We could not find negative species‐habitat associations for rare species, probably due to their small sample sizes. Among four species small trees were less strongly associated with habitat than large trees, implying competitive exclusion of trees in suboptimal habitats. The other four species showed the opposite pattern, possibly owing to the smaller sample size of large trees. A habitat had a maximum of three species with which it was significantly positively associated. Conclusions : For a species to survive in population equilibrium in a landscape, habitats in which ‘source’ subpopulations can be sustained without subsidy from adjacent habitats are essential. Competition is most severe among related species whose source subpopulations share the same habitat. On the evidence of source subpopulations identified by positive species‐habitat association, species‐habitat association reduces the number of confamilial competitors. Our results therefore indicate that edaphic niche specialization contributes to coexistence of species of Sterculiaceae in the plot, consistent with the expectations of equilibrium hypotheses.     

Liana habitat associations and community structure in a Bornean lowland tropical forest

Lianas (woody vines) contribute substantially to the diversity and structure of most tropical forests, yet little is known about the importance of habitat specialization in maintaining tropical liana diversity and the causes of variation among forests in liana abundance and species composition. We examined habitat associations, species diversity, species composition, and community structure of lianas at Sepilok Forest Reserve, Sabah, Malaysia in northeastern Borneo among three soil types that give rise to three distinct forest types of lowland tropical rain forest: alluvial, sandstone hill, and kerangas (heath) forest. Alluvial soils are more nutrient rich and have higher soil moisture than sandstone soils, whereas kerangas soils are the most nutrient poor and drought prone. Lianas ≥0.5-cm in diameter were measured, tagged, and identified to species in three square 0.25-ha plots in each forest type. The number of lianas ≥0.5 cm did not differ significantly among forest types and averaged 1348 lianas ha⁻¹, but mean liana stem diameter, basal area, estimated biomass, species richness, and Fisher’s diversity index were all greater for plots in alluvial than sandstone or kerangas forests. Liana species composition also differed greatly among the three habitats, with 71% of species showing significant positive or negative habitat associations. Sandstone forests were intermediate to alluvial and kerangas forests in most aspects of liana community structure and composition, and fewer species showed significant habitat associations with this forest type. Ranking of forest types with respect to liana density, biomass, and diversity matches the ranking in soil fertility and water availability (alluvial > sandstone hill > kerangas). These results suggest that edaphic factors play an important role in maintaining liana species diversity and structuring liana communities.     

Species–area and species–sampling effort relationships: disentangling the effects

Species numbers tend to increase with both the area surveyed (species–area relationship, SAR) and the number of samples taken (species–sampling effort relationship, SSER). These two relationships differ in their nature and underlying mechanisms but are not clearly distinguished in field studies. To discriminate the effects of area (spatial extent) and sampling effort (SE) on species richness, several models explicitly involving both variables were proposed and tested against 13 datasets from marine micro‐, meio‐ and macrobenthos. A combination of power SSER and piecewise power SAR terms was found to have the best fit. The effects of area and SE were both significant, but the former one was noticeably weaker. The SSERs were roughly linear in log‐log space, whereas the SARs demonstrated scale‐dependent behavior with a noticeable threshold (slope breakpoint). Species richness was almost area‐independent below this threshold (the “small area effect”, SAE) but followed typical power‐law SAR beyond the threshold. This effect was similar to the “small island effect” but occurred for arbitrarily delineated areas within continuous habitats. Parameters of the SAR curves depended on organism size. The upper limit of the SAE increased from microorganisms to meiofauna to macrofauna. Also, SAR curves for unicellular groups had significantly lower slopes. SAE is supposed to indicate a spatial range of statistical homogeneity in species composition. Its upper limit corresponds to the characteristic size of a local community (a single habitat occupied by a common species pool). Interpretations of SAR and SSER parameters in terms of α‐ and β‐diversity are proposed. Both SAR and SSER slopes obtained from univariate regressions are overestimated. This upward bias depends on sampling design, decreasing for SAR but increasing for SSER with more unequally spaced samples. Both spatial extent and sampling effort should be taken into account to disentangle properly their effects on diversity.     

Characterizing fish–habitat associations in streams as the first step in ecological restoration

Abstract Habitat availability is often regarded as the primary factor that limits population and community recovery in degraded ecosystems, and physical habitat is thus often targeted in restoration. The identification of which habitat(s) to attempt to restore is a critical step in the restoration process, but one for which there is often a paucity of useful information. Here we examine the distribution of fish in three lowland streams in Victoria, Australia, that have been degraded by severe sedimentation. We aim to identify habitats that are associated with high abundances of native fish, and that thus might be appropriate to target in habitat restoration. Associations between native fish abundances and physical habitat characteristics were examined at three spatial scales (among streams, among sites and within sites) to determine the types of habitat to which fish respond, and the scales over which these responses occur. Of the four species of native fish found in the streams, three (Galaxias olidus Günther, Gadopsis marmoratus (Richardson) and Nanoperca australis Günther) showed significant habitat associations at small spatial scales (i.e. within sites). In particular, these species were generally found in deeper water, and in close proximity to cover (typically either coarse or fine woody debris or vegetation). Differences in habitat availability among sites and streams were less influential, except in the case of G. marmoratus, which was completely absent from both the ephemeral streams. Although our results suggest that these species collectively respond to habitat at several spatial scales, fish distributions were allied to the presence of habitat structure at the scale of metres, the smallest spatial scale examined. We hypothesize that fish abundances are currently limited by the low availability of habitat at these small spatial scales. It may therefore be possible to increase fish abundances in these creeks by augmenting the amount of available habitat via stream restoration.     

Climate change vulnerability of forest biodiversity: climate and competition tracking of demographic rates

Forest responses to climate change will depend on demographic impacts in the context of competition. Current models used to predict species responses, termed climate envelope models (CEMs), are controversial, because (i) calibration and prediction are based on correlations in space (CIS) between species abundance and climate, rather than responses to climate change over time (COT), and (ii) they omit competition. To determine the relative importance of COT, CIS, and competition for light, we applied a longitudinal analysis of 27 000 individual trees over 6–18 years subjected to experimental and natural variation in risk factors. Sensitivities and climate and resource tracking identify which species are vulnerable to these risk factors and in what ways. Results show that responses to COT differ from those predicted based on CIS. The most important impact is the effect of spring temperature on fecundity, rather than any input variable on growth or survival. Of secondary importance is growing season moisture. Species in the genera Pinus, Ulmus, Magnolia, and Fagus are particularly vulnerable to climate variation. However, the effect of competition on growth and mortality risk exceeds the effects of climate variation in space or time for most species. Because sensitivities to COT and competition are larger than CIS, current models miss the most important effects. By directly comparing sensitivity to climate in time and space, together with competition, the approach identifies which species are sensitive to climate change and why, including the heretofore overlooked impact on fecundity.     

Species–area relationships and biodiversity loss in fragmented landscapes

To estimate species loss from habitat destruction, ecologists typically use species–area relationships, but this approach neglects the spatial pattern of habitat fragmentation. Here, we provide new, easily applied, analytical methods that place upper and lower bounds on immediate species loss at any spatial scale and for any spatial pattern of habitat loss. Our formulas are expressed in terms of what we name the ‘Preston function’, which describes triphasic species–area relationships for contiguous regions. We apply our method to case studies of deforestation and tropical tree species loss at three different scales: a 50 ha forest plot in Panama, the tropical city‐state of Singapore and the Brazilian Amazon. Our results show that immediate species loss is somewhat insensitive to fragmentation pattern at small scales but highly sensitive at larger scales: predicted species loss in the Amazon varies by a factor of 16 across different spatial structures of habitat loss.     

Fungicides for forest trees,shade trees and forest products

The Botanical Review -     

Distributions and habitat associations of the bridled monocle bream Scolopsis bilineatus (Nemipteridae): a demographic approach

This study focussed on the demography and ecology of Scolopsis bilineatus at three locations on the Great Barrier Reef: the Lizard Island Group, Orpheus Island and One Tree Island. Scolopsis bilineatus lived for up to 16 years and had four distinct life‐history stages, which varied in their distribution patterns, habitat use and reproductive behaviour. Pre‐maturational sex change occurred whereby all males were derived from immature females, and males grew faster and larger than females. Small females and larger males generally formed pairs, which influenced their spatial distributions at small scales. Distributions of S. bilineatus were influenced by depth and exposure within reefs, particularly for juveniles, and most fish were found in shallow, sheltered habitats. Abundance was influenced by benthic cover, and was higher in areas of high coral cover and low where algae were abundant. Habitat associations were stronger at the microhabitat scale, and shelter sites were important for adults. Ontogenetic changes in microhabitat associations were found: juveniles occupied sand and rubble, and adults occupied shelters such as caves and overhangs. Adults showed site fidelity for shelter sites over a period of 4 days and returned to specific shelter sites repeatedly. These findings illustrate the importance of understanding the spatial ecology and habitat use of coral reef fishes, particularly with reference to size‐based changes within species.