Above- and belowground interactions drive habitat segregation between two cryptic species of tropical trees

In the lowlands of central Panama, the Neotropical pioneer tree Trema micrantha (sensu lato) exists as two cryptic species: "landslide" Trema is restricted to landslides and road embankments, while "gap" Trema occurs mostly in treefall gaps. In this study, we explored the relative contributions of biotic interactions and physical factors to habitat segregation in T. micrantha. Field surveys showed that soils from landslides were significantly richer in available phosphorus and harbored distinct arbuscular mycorrhizal fungal (AMF) communities compared to gap soils. Greenhouse experiments designed to determine the effect of these abiotic and biotic differences showed that: (1) both landslide and gap species performed better in sterilized soil from their own habitat, (2) the availability of phosphorus and nitrogen was limiting in gap and landslide soils, respectively, (3) a standardized AMF inoculum increased performance of both species, but primarily on gap soils, and (4) landslide and gap species performed better when sterilized soils were inoculated with the microbial inoculum from their own habitat. A field experiment confirmed that survival and growth of each species was highest in its corresponding habitat. This experiment also showed that browsing damage significantly decreased survival of gap Trema on landslides. We conclude that belowground interactions with soil microbes and aboveground interactions with herbivores contribute in fundamental ways to processes that may promote and reinforce adaptive speciation.     

Spatial clustering of habitat structure effects patterns of community composition and diversity

Natural ecosystems often show highly productive habitats that are clustered in space. Environmental disturbances are also often nonrandomly distributed in space and are either intrinsically linked to habitat quality or independent in occurrence. Theoretical studies predict that configuration and aggregation of habitat patch quality and disturbances can affect metacommunity composition and diversity, but experimental evidence is largely lacking. In a metacommunity experiment, we tested the effects of spatially autocorrelated disturbance and spatial aggregation of patch quality on regional and local richness, among-community dissimilarity, and community composition. We found that spatial aggregation of patch quality generally increased among-community dissimilarity (based on two measures of beta diversity) of communities containing protozoa and rotifers in microcosms. There were significant interacting effects of landscape structure and location of disturbances on beta diversity, which depended in part on the specific beta diversity measures used. Effects of disturbance on composition and richness in aggregated landscapes were generally dependent on distance and connectivity among habitat patches of different types. Our results also show that effects of disturbances in single patches cannot directly be extrapolated to the landscape scale: the predictions may be correct when only species richness is considered, but important changes in beta diversity may be overlooked. There is a need for biodiversity and conservation studies to consider the spatial aggregation of habitat quality and disturbance, as well as connectivity among spatial aggregations.     

Spatial and temporal variability modify density dependence in populations of large herbivores

A central challenge in ecology is to understand the interplay of internal and external controls on the growth of populations. We examined the effects of temporal variation in weather and spatial variation in vegetation on the strength of density dependence in populations of large herbivores. We fit three subsets of the model ln(Nt) = a + (1 + b) x ln(N(t-1)) + c x ln(N(t-2)) to five time series of estimates (Nt) of abundance of ungulates in the Rocky Mountains, USA. The strength of density dependence was estimated by the magnitude of the coefficient b. We regressed the estimates of b on indices of temporal heterogeneity in weather and spatial heterogeneity in resources. The 95% posterior intervals of the slopes of these regressions showed that temporal heterogeneity strengthened density-dependent feedbacks to population growth, whereas spatial heterogeneity weakened them. This finding offers the first empirical evidence that density dependence responds in different ways to spatial heterogeneity and temporal heterogeneity.     

Spatial memory, habitat auto-facilitation and the emergence of fractal home range patterns

Animals interact with their habitat in a manner which involves both negative and positive feedback mechanisms. We apply a specific modeling approach, “multi-scaled random walk”, for the scenario where a spatially explicit positive feedback process emerges from a combination of a spatial memory-dependent tendency to return to familiar patches and a consequently objective or subjective improvement of the quality of these patches (habitat auto-facilitation). In addition to the potential for local resource improvement from physically altering a patch, primarily known from the ecology of grazing ungulates, auto-facilitation from site fidelity may also embed more subtle subjective, individual-specific advantages from patch familiarity. Under the condition of resource superabundance, fitness gain from intra-home range patch fidelity creates a self-reinforcing use of the preferred patches on expense of a broader foraging in a priori equally favorable patches. Through this process, our simulations show that a spatially fractal dispersion of accumulated locations of the individual will emerge under the given model assumptions. Based on a conjecture that intra-home range patch fidelity depends on spatial memory we apply the multi-scaled random walk model to construct a spatially explicit habitat suitability parameter H_ij, which quantifies the dispersion of the generally most constraining resource from the individual's perspective. An intra-home range set of observed H-scores, H_obs, can then be estimated from a simple 2-scale calculation that is derived from the local dispersion of fixes. We show how the spatially explicit habitat utilization index H_obs not necessarily correlates positively with the local density fluctuations of fixes. The H-index solves some well-known problems from using the pattern of local densities of telemetry fixes - the classic utilization distribution - as a proxy variable for relative intra-home range habitat quality and resource selection. A pilot study on a set of telemetry fixes collected from a herd of free-ranging domestic sheep with overlapping summer home ranges illustrates how the H-index may be estimated and interpreted as a first-level approach towards a more extensive analysis of intra-home range habitat resource availability and patch preferences. Spatial memory in combination with site fidelity requires a modeling framework that explicitly describes the property of positive feedback mechanism under auto-facilitation in a spatio-temporally explicit manner.     

Spatial structure and habitat associations of demersal assemblages in the Gulf of Lions: a multicompartmental approach

Experimental trawl surveys were performed in the Gulf of Lions. A multicompartmental approach was used to study the spatial distribution of demersal assemblages and to estimate their associations with two other components of the ecosystem: the benthic macrofauna and substratum type. At the Gulf scale, we found that these components explained 38 and 19% of the total variability of the spatial organization of groundfishes, respectively. Our results showed that groundfish and benthic macrofauna species were primarily distributed along a coast–open sea gradient and secondarily along a longitudinal axis. The longitudinal variability of the spatial organisation of species was maximal in the shelf area, whereas the upper-slope and coastal areas were characterized by strong homogeneity. Although no partitioning could be made based on their spatial distribution, the coastal groundfish assemblages were split into two groups. In the first group, groundfishes were strongly associated with both benthic macrofauna and type of substratum; in the second group, they were strongly independent of these two components. Furthermore, in the Gulf of Lions, the presence of productive zones associated with the Rhône river plume or with shelf-break upwelling seemed to attract some groundfishes. This study illustrates how useful such a multicompartmental approach can be for both fisheries management and community ecology.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

Spatial variability in habitat associations of pre- and post-settlement stages of coral reef fishes at Ishigaki Island,Japan

Successful settlement of pelagic fish larvae into benthic juvenile habitats may be enhanced by a shortened settlement period, since it limits larval exposure to predation in the new habitat. Because the spatial distribution of marine fish larvae immediately prior to settlement versus during settlement was unknown, field experiments were conducted at Ishigaki Island (Japan) using light trap sampling and underwater visual belt transect surveys to investigate the spatial distribution patterns of selected pre- and post-settlement fishes (Acanthuridae, Pomacentridae, Chaetodonidae and Lethrinidae) among four habitats (seagrass bed, coral rubble, branching coral and tabular coral). The results highlighted two patterns: patterns 1, pre- and post-settlement individuals showing a ubiquitous distribution among the four habitats (Acanthuridae) and pattern 2, pre-settlement individuals distributed in all habitats, but post-settlement individuals restricted to coral (most species of Pomacentridae and Chaetodontidae) or seagrass habitats (Lethrinidae). The first pattern minimizes the transition time between the larval pelagic stage and acquisition of a benthic reef habitat, the latter leading immediately to a juvenile lifestyle. In contrast, the second pattern is characterized by high settlement habitat selectivity by larvae and/or differential mortality immediately after settlement.     

Functional traits and the growth-mortality trade-off in tropical trees

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.     

Spatial characteristics of habitat patches and population survival

The effect of the spatial arrangement of habitat patches on the survival of resident populations were considered in a stochastic model using population parameters appropriate to Peromyscus leucopus. The 34 possible arrangements of connections among five otherwise identical patches were simulated in order to determine the survival probabilities and population sizes.The main findings are that populations in completely isolated patches have lower survival probabilities than those in patches that are connected to other patches, are connected to the survival probabilities of populations in connected patches increases with the size of the largest geometric figure of which the patch is a part. The results are discussed in the context of resource management.     

Transcending scale dependence in identifying habitat with resource selection functions

Multi-scale resource selection modeling is used to identify factors that limit species distributions across scales of space and time. This multi-scale nature of habitat suitability complicates the translation of inferences to single, spatial depictions of habitat required for conservation of species. We estimated resource selection functions (RSFs) across three scales for a threatened ungulate, woodland caribou (Rangifer tarandus caribou), with two objectives: (1) to infer the relative effects of two forms of anthropogenic disturbance (forestry and linear features) on woodland caribou distributions at multiple scales and (2) to estimate scale-integrated resource selection functions (SRSFs) that synthesize results across scales for management-oriented habitat suitability mapping. We found a previously undocumented scale-specific switch in woodland caribou response to two forms of anthropogenic disturbance. Caribou avoided forestry cut-blocks at broad scales according to first- and second-order RSFs and avoided linear features at fine scales according to third-order RSFs, corroborating predictions developed according to predator-mediated effects of each disturbance type. Additionally, a single SRSF validated as well as each of three single-scale RSFs when estimating habitat suitability across three different spatial scales of prediction. We demonstrate that a single SRSF can be applied to predict relative habitat suitability at both local and landscape scales in support of critical habitat identification and species recovery.     

Null models reveal preferential sampling, spatial autocorrelation and overfitting in habitat suitability modelling

Nowadays, species are driven to extinction at a high rate. To reduce this rate it is important to delineate suitable habitats for these species in such a way that these areas can be suggested as conservation areas. The use of habitat suitability models (HSMs) can be of great importance for the delineation of such areas. In this study MaxEnt, a presence-only modelling technique, is used to develop HSMs for 223 nematode species of the Southern Bight of the North Sea. However, it is essential that these models are beyond discussion and they should be checked for potential errors. In this study we focused on two categories (1) errors which can be attributed to the database such as preferential sampling and spatial autocorrelation and (2) errors induced by the modelling technique such as overfitting, In order to quantify these adverse effects thousands of nulls models were created. The effect of preferential sampling (i.e. some areas where visited more frequenty than others) was investigated by comparing model outcomes based from null models sampling the actual sampling stations and null models sampling the entire mapping area (Raes and ter Steege, 2007). Overfitting is exposed by a fivefold cross-validation and the influence of spatial autocorrelation is assessed by separating test and training sets in space. Our results clearly show that all these effects are present: preferential sampling has a strong effect on the selection of non-random species models. Crossvalidation seems to have less influence on the model selection and spatial autocorrelation is also strongly present. It is clear from this study that predefined thresholds are not readily applicable to all datasets and additional tests are needed in model selection.     

Seasonal water stress tolerance and habitat associations within four neotropical tree genera

We investigated the relationship between habitat association and physiological performance in four congeneric species pairs exhibiting contrasting distributions between seasonally flooded and terra firme habitats in lowland tropical rain forests of French Guiana, including Virola and Iryanthera (Myristicaceae), Symphonia (Clusiaceae), and Eperua (Caesalpiniaceae). We analyzed 10-year data sets of mapped and measured saplings (stems >150 cm in height and <10 cm diameter at breast height [dbh]) and trees (stems > or =10 cm dbh) across 37.5 ha of permanent plots covering a 300-ha zone, within which seasonally flooded areas (where the water table never descends below 1 m) have been mapped. Additionally, we tested the response of growth, survival, and leaf functional traits of these species to drought and flood stress in a controlled experiment. We tested for habitat preference using a modification of the torus translation method. Strong contrasting associations of the species pairs of Iryanthera, Virola, and Symphonia were observed at the sapling stage, and these associations strengthened for the tree stage. Neither species of Eperua was significantly associated with flooded habitats at the sapling stage, but E. falcata was significantly and positively associated with flooded forests at the tree stage, and trees of E. grandiflora were found almost exclusively in nonflooded habitats. Differential performance provided limited explanatory support for the observed habitat associations, with only congeners of Iryanthera exhibiting divergent sapling survival and tree growth. Seedlings of species associated with flooded forest tended to have higher photosynthetic capacity than their congeners at field capacity. In addition, they tended to have the largest reductions in leaf gas exchange and growth rate in response to experimental drought stress and the least reductions in response to experimental inundation. The corroboration of habitat association with differences in functional traits and, to a lesser extent, measures of performance provides an explanation for the regional coexistence of these species pairs. We suggest that specialization to seasonally flooded habitats may explain patterns of adaptive radiation in many tropical tree genera and thereby provide a substantial contribution to regional tree diversity.     

Niche segregation and habitat specialisation of harpacticoid copepods in a tropical seagrass bed

Several harpacticoid copepod species are adapted to an epiphytic lifestyle. Previous studies on tropical seagrass meiofauna mainly focussed on the epiphytic communities and neglected the benthic component. The present study aims to document the benthic harpacticoid copepod communities sampled from different sediment depth horizons adjacent to five seagrass species in the intertidal and subtidal zone of a tropical seagrass bed (Gazi Bay, Kenya). Two benthic copepod communities could be identified mainly based on the tidal position of the samples: a first community was collected near the intertidal seagrasses Halophila ovalis and Halodule wrightii; a second community occurred near the subtidal seagrasses Thalassia hemprichii, Syringodium isoetifolium and Halophila stipulacea. The first community was mainly determined by sediment characteristics (e.g. skewness), while the second community was split off based on organic matter content (% TOM), nutrient and pigment values. A subtle combination of horizontal and vertical niche segregation was reported for the dominant copepod families. Species of the families Thalestridae, Laophontidae and Diosaccidae were structured by tidal position and showed a strong preference for the subtidal zone. The opposite strategy, i.e. a clear preference for the intertidal zone, was found for copepods belonging to the families Paramesochridae and Canuellidae. In addition, Apodopsyllus africanus (Paramesochridae) was well-adapted to stress and was concentrated in the deeper sediment layers near the subtidal seagrasses. On the other hand, Canuellidae, as filter feeders, were concentrated in the upper centimetres of the sediment. The families Ectinosomatidae and Cletodidae did not show any vertical or horizontal segregation. On the species level, however, clear horizontal niche segregation was detected for the family Cletodidae. In addition to the reported ecological results, the study material was used to evaluate different niche definitions. We found tidal position to be the most important factor forcing harpacticoids to specialise. Sediment depth horizon was less powerful in dividing the families into different guilds (from specialists to generalists) based on standardised niche breadth. The present study documents the subtle habitat partitioning of co-existing species in a limited area and its role in sustaining high biodiversity in the community.     

Seasonal changes in habitat structure underpin shifts in macroalgae-associated tropical fish communities

Habitat shifts play an important role in structuring faunal assemblages; however, research has focused on the influence of random disturbance events and information on how regular seasonal changes to habitat affect marine fauna remains largely unexplored, especially in the tropics. We recorded seasonal changes in the structure of tropical macroalgae fields within the Ningaloo lagoon (Western Australia) and related this to the density, biomass and species richness of fishes that represent key processes: juveniles, predators of juveniles and herbivores. The extent and direction of seasonal changes in macroalgae were inconsistent among sites, creating a highly dynamic habitat matrix across time and space. Species richness and density of fishes were largely maintained where density of holdfasts from canopy-forming macroalgae and/or cover was high across seasons, but shifted markedly in areas of macroalgae habitat loss: suggesting stable habitat structure is critical for the persistence of macroalgae-associated fishes. Our results demonstrate that macroalgae fields that maintain high structural complexity across different seasons are more likely to preserve key ecological processes and therefore warrant greater conservation attention within a spatial management framework.     

Autocorrelation offsets zero-inflation in models of tropical saplings density

Modelling the local density of tropical saplings can provide insights into the ecological processes that drive species regeneration and thereby help predict population recovery after disturbance. Yet, few studies have addressed the challenging issues in autocorrelation and zero-inflation of local density. This paper presents Hierarchical Bayesian Modelling (HBM) of sapling density that includes these two features. Special attention is devoted to variable selection, model estimation and comparison.     

A note on density dependence in population models

One of the most studied phenomena in ecology is density dependent regulation. The model most frequently used to study this behaviour is the theta-logistic model. However, disagreement has developed within the ecology community pertaining to the interpretation of this model’s parameters, and thus as to appropriate values for the parameters to assume. In particular, the parameter θ$\theta$ has been allowed to take negative values, resulting in the ‘growth rate parameter’ estimated to be negative for species which are extant and exhibit no signs of becoming extinct in the short-term. Here we explain this phenomenon by formulating the theta-logistic model in the manner in which the original logistic model was formulated by Verhulst (1838), in doing so providing a simple interpretation of model parameters and thus restrictions on values the parameters may assume. We conclude that θ$\theta$ should (almost always) be restricted to values greater than −1$-1$. This has implications for studies assessing the form of density dependence from data. Additionally, another model appearing in the literature is presented which provides a more flexible model of density dependence at the expense of only one additional parameter.