Lizard scales in an adaptive radiation: variation in scale number follows climatic and structural habitat diversity in Anolis lizards

Lizard scales vary in size, shape and texture among and within species. The overall function of scales in squamates is attributed to protection against abrasion, solar radiation and water loss. We quantified scale number of Anolis lizards across a large sample of species (142 species) and examined whether this variation was related either to structural or to climatic habitat diversity. We found that species in dry environments have fewer, larger scales than species in humid environments. This is consistent with the hypothesis that scales reduce evaporative water loss through the skin. In addition, scale number varied among groups of ecomorphs and was correlated with aspects of the structural microhabitat (i.e. perch height and perch diameter). This was unexpected because ecomorph groups are based on morphological features related to locomotion in different structural microhabitats. Body scales are not likely to play an important role in locomotion in Anolis lizards. The observed variation may relate to other features of the ecomorph niche and more work is needed to understand the putative adaptive basis of these patterns. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 570–579.     

Nowhere to run: the role of habitat openness and refuge use in defining patterns of morphological and performance evolution in tropical lizards

For species from open habitats with little cover and few refugia, selection should favour morphologies that enhance performance at tasks that enable rapid movement across open areas. Similarly, selection should also favour traits that enable rapid access and movement within suitable refugia. This study examined the relationship between habitat openness, refuge use, morphology and performance of 19 species representing 23 populations of tropical Lygosomine skink. Species from this group occupy a wide array of habitats from open forest and open rocky intertidal zones to high‐altitude heaths and dense, closed forests. Species that occupied open habitats were faster at sprinting, climbing and had better cling ability than species from more cluttered, closed habitats. In addition, species from habitats that used rock crevices as refuges had enhanced sprinting ability. This study shows the importance of both habitat openness and refuge type in the evolution of both the morphology and performance in lizards.     

Non-random interactions within and across guilds shape the potential to coexist in multi-trophic ecological communities

Theory posits that the persistence of species in ecological communities is shaped by their interactions within and across trophic guilds. However, we lack empirical evaluations of how the structure, strength and sign of biotic interactions drive the potential to coexist in diverse multi-trophic communities. Here, we model community feasibility domains, a theoretically informed measure of multi-species coexistence probability, from grassland communities comprising more than 45 species on average from three trophic guilds (plants, pollinators and herbivores). Contrary to our hypothesis, increasing community complexity, measured either as the number of guilds or community richness, did not decrease community feasibility. Rather, we observed that high degrees of species self-regulation and niche partitioning allow for maintaining larger levels of community feasibility and higher species persistence in more diverse communities. Our results show that biotic interactions within and across guilds are not random in nature and both structures significantly contribute to maintaining multi-trophic diversity.     

Density dependence and habitat preference shape seedling survival in a subtropical forest in central China

Aims Seedlings are vulnerable to many kinds of fatal abiotic and biotic agents, and examining the causes of seedling dynamics can help understand mechanisms of species coexistence. To disentangle the relative importance of neighborhood densities, habitat factors and phylogenetic relatedness on focal seedling survival, we monitored the survival of 5306 seedlings of 104 species>15 months. We address the following questions: (i) How do neighborhood densities, habitat variables and phylogenetic relatedness affect seedling survival? What is the relative importance of conspecific densities, habitat variables and phylogenetic relatedness to seedling survival? (ii) Does the importance of the neighborhood densities, habitat variables and phylogenetic relatedness vary among growth forms, leaf habits or dispersal modes? Specially, does the conspecific negative density dependence inhibit tree and deciduous seedlings more compared with shrub and evergreen species? Does density dependence affect the wind and animal-dispersed species equally?Methods We established 135 census stations to monitor seedling dynamics in a 25-ha subtropical forest plot in central China. Conspecific and heterospecific seedling density in the 1-m 2 seedling plot and adult basal area within a 20-m radius provided neighborhood density variables. Mean elevation, convexity and aspect of every 5- × 5-m grid with seedling plots were used to quantify habitat characteristics. We calculated the relative average phylodiversity between focal seedling and heterospecific neighbors to quantify the species relatedness in the neighborhood. Eight candidate generalized linear mixed models with binominal error distribution were used to compare the relative importance of these variables to seedling survival. Akaike's information criteria were used to identify the most parsimonious models.Important findings At the community level, both the neighborhood densities and phylogenetic relatedness were important to seedling survival. We found negative effects of increasing conspecific seedlings, which suggested the existence of species-specific density-dependent mortality. Phylodiversity of heterospecific neighbors was negatively related to survival of focal seedlings, indicating similar habitat preference shared among phylogenetically closely related species may drive seedling survival. The relative importance of neighborhood densities, habitat variables and phylogenetic relatedness varied among ecological guilds. Conspecific densities had significant negative effect for deciduous and wind-dispersed species, and marginally significant for tree seedlings>10cm tall and animal-dispersed species. Habitat variables had limited effects on seedling survival, and only elevation was related to the survival of evergreen species in the best-fit model. We conclude that both negative density-dependent mortality and habitat preference reflected by the phylogenetic relatedness shape the species coexistence at seedling stage in this forest.     

Natural ecosystems: Pattern and process in relation to local and landscape diversity in southwestern Australian woodlands

Natural woodland ecosystems in southwestern Australia are examined in terms of their spatial structure at three different scales. Fine-scale structure of the vegetation and soil surface has a strong influence on water capture and retention, resulting in mostly localised infiltration and use. Stand-scale structure results from the patchy distribution of canopy trees, regeneration patches and a diversity of understorey shrubs which provides a diverse above- and below-ground structure. At the landscape scale, broad vegetation patterns are related to soil and landform changes along catenas, but there are also finer-scale mosaics resulting from disturbance history, individualistic species distributions and more subtle physical/chemical changes. Complex patterns at all these scales confer resilience to the woodland system. Most of this complexity has been lost in the current agricultural system, and it can be hypothesised that many current degradation problems can be related to this. The challenge for the future is to develop complexity and resilience in agricultural systems while maintaining an acceptable level of production.     

Extreme specialization to rocky habitats in Tropidurus lizards from Brazil: Trade‐offs between a fitted ecomorph and autoecology in a harsh environment

Ecomorphological theory indicates that different ecological requirements lead to different organismal designs. Given that species with equal requirements could not coexist, traits leading to more efficient use of resources may be selected to avoid competition among closely related syntopic species, generating specialized ecomorphs. We compared habitat use, diet, thermal biology and morphology among the syntopic Tropidurus semitaeniatus, T. helenae and T. hispidus in the Caatinga of Northeastern Brazil. Tropidurus semitaeniatus and T. helenae are flattened lizards specialized to rocks and rock crevices, whereas T. hispidus has a robust body and generalist habits. We aimed to test the hypothesis that morphological modifications observed in the flattened ecomorphs are related to modifications in diet and habitat use. Also, we hypothesized that specialization to habitat induces morphological modifications, which in turn may constrain lizard performance. Flattened species differed in habitat use, morphology and prey size when compared with the generalist ecomorph. Morphological modifications were related to specializations to rocky habitats and constrained the variety of prey items consumed. This phenotype also reduced their reproductive output when compared with a robust, generalist ecomorph.     

Spatial and temporal abundance,substrate partitioning and species co-occurrence in a guild of Neotropical blackflies (Diptera: Simuliidae)

Blackfly larvae were sampled monthly from two smallNeotropical mountain streams 500 m apart from each other in northernVenezuela during a 15-month period, and ten habitat variables weremeasured at two altitudes (800 and 900 m) to determine local spatialand temporal variations in abundance, stream variables, speciesco-occurrence, and substrate preference within the blackflyguild. The blackfly species composition of each stream was the same,with five species (in order of abundance): Simulium ochraceumWalker s.l., S. paynei Vargas, S. metallicum Bellardicytospecies E, Simulium sp., and S. horacioi Okazawa &Onishi. There were clear differences in community structurebetween streams (not among altitudes), although S. ochraceums.l. was the most abundant species in both habitats. Principal component andcorrelation analyses revealed that structural (great depth, width,flow, and discharge) and chemical (low conductivity) factors werethe stream variables best associated to the spatial distribution ofmost blackfly species. Blackfly abundance was greater in the largerstream. Larvae were most abundant on submerged rocks and fallenleaves, although the patterns of substrate use within the guildrevealed substrate partitioning among most blackfly species.There was one general annual peak of abundance at the end of therainy season. We suggest that blackfly community structure in theseNeotropical headwaters streams, over the small spatial scaleexamined here, is largely influenced by interactions among streamsize and velocity-related factors (width, streamflow, discharge andsubstrate availability-stability) and the rainfall regime.     

Light and shape: a contribution to demonstrate morphological differences in diurnal and nocturnal teleosts

Light intensity is an important environmental factor affecting the structure of fish assemblages during the day-night cycle. Light influences how organisms perceive their environment, modulating their intraspecific and interspecific relationships. The relationship between light intensity variations and biological cycles should be observed at the level of organismal morphology. In this study the relationship between activity rhythms, thus light intensity experienced by fish in the period of major activity and external morphology, have been investigated. The morphological traits of 97 selected fish species were compared in order to determine the existence of a common morphological plan in agreement with their diurnal or nocturnal activity rhythm. Species sorting was performed by maximizing the diversity of activity rhythm, habitat choice, ecology, and trophic habits within the same family, to assess the importance of the day-night cycle on species morphology in relation to other environmental features. The morphological characters selected for the geometric morphometric analysis were body profile and the position of mouth, eye, pelvic, pectoral, dorsal, and caudal fin. The present analysis allowed different consensus forms for nocturnal and for diurnal species to be identified. Two-block Partial Least Squares analysis was then performed for the purpose of modeling the covariation between the form and two important external variables (ecology and activity).     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Exploring the nature of ecological specialization in a coral reef fish community: morphology,diet and foraging microhabitat use

Patterns of ecological specialization offer invaluable information about ecosystems. Yet, specialization is rarely quantified across several ecological niche axes and variables beyond the link between morphological and dietary specialization have received little attention. Here, we provide a quantitative evaluation of ecological specialization in a coral reef fish assemblage (f. Acanthuridae) along one fundamental and two realized niche axes. Specifically, we examined ecological specialization in 10 surgeonfish species with regards to morphology and two realized niche axes associated with diet and foraging microhabitat utilization using a recently developed multidimensional framework. We then investigated the potential relationships between morphological and behavioural specialization. These relationships differed markedly from the traditional ecomorphological paradigm. While morphological specialization showed no relationship with dietary specialization, it exhibited a strong relationship with foraging microhabitat specialization. However, this relationship was inverted: species with specialized morphologies were microhabitat generalists, whereas generalized morphotypes were microhabitat specialists. Interestingly, this mirrors relationships found in plant–pollinator communities and may also be applicable to other ecosystems, highlighting the potential importance of including niche axes beyond dietary specialization into ecomorphological frameworks. On coral reefs, it appears that morphotypes commonly perceived as most generalized may, in fact, be specialized in exploiting flat and easily accessible microhabitats.     

Intersexual segregation in foraging microhabitat use by Magellanic Woodpeckers (Campephilus magellanicus): Seasonal and habitat effects at the world's southernmost forests

Animals facing seasonal food shortage and habitat degradation may adjust their foraging behaviour to reduce intraspecific competition. In the harsh environment of the world's southernmost forests in the Magellanic sub‐Antarctic ecoregion in Chile, we studied intersexual foraging differences in the largest South American woodpecker species, the Magellanic Woodpecker (Campephilus magellanicus). We assessed whether niche overlap between males and females decrease when food resources are less abundant or accessible, that is, during winter and in secondary forests, compared to summer and in old‐growth forests, respectively. We analysed 421 foraging microhabitat observations from six males and six females during 2011 and 2012. As predicted, the amount of niche overlap between males and females decreased during winter, when provisioning is more difficult. During winter, males and females (i) used trees with different diameter at breast height (DBH); (ii) fed in trunk sections with different diameters; and (iii) fed at different heights on tree trunks or branches. Vertical niche partitioning between sexes was found in both old‐growth and secondary forests. Such a niche partitioning during winter may be a seasonal strategy to avoid competition between sexes when prey resources are less abundant or accessible. Our results suggest that the conservation of this forest specialist, dimorphic and charismatic woodpecker species requires considering differences in habitat use between males and females.     

Using presence‐only modelling to predict Asian elephant habitat use in a tropical forest landscape: implications for conservation

Aim Asian elephants, Elephas maximus, are threatened throughout their range by a combination of logging, large scale forest conversion and conflict with humans. We investigate which environmental factors, both biotic and abiotic, constrain the current distribution of elephants. A spatially explicit habitat model is constructed to find core areas for conservation and to assess current threats. Location Ulu Masen Ecosystem in the province of Nanggroe Aceh Darussalam on the island of Sumatra, Indonesia. Methods A stratified survey was conducted at 12 sites (300 transects) to establish the presence of elephants. Presence records formed the basis to model potential habitat use. Ecological niche factor analysis (ENFA) is used to describe their niche and to identify key factors shaping elephant distribution. An initial niche model was constructed to describe elephant niche structure, and a second model focused on identifying core areas only. To assess the threat of habitat encroachment, overlap between the elephants’ optimal niche and the occurrence of forest encroachment is computed. Results Elephants were recorded throughout the study area from sea level to 1600 m a.s.l. The results show that the elephant niche and consequently habitat use markedly deviates from the available environment. Elephant presence was positively related to forest cover and vegetation productivity, and elephants were largely confined to valleys. A spatially explicit model showed that elephants mainly utilize forest edges. Forest encroachment occurs throughout the elephants range and was found within 80% of the elephants’ ecological niche. Main conclusions In contrast to general opinion, elephant distribution proved to be weakly constrained by altitude, possibly because of movement routes running through mountainous areas. Elephants were often found to occupy habitat patches in and near human‐dominated areas. This pattern is believed to reflect the displacement of elephants from their former habitat.     

A shift in the habitat use pattern of a lentic galaxiid fish: an acute behavioural response to an introduced predator

Despite potentially reducing predation mortality, behavioural responses of native species to introduced predators may still have sub-lethal impacts. In video-recorded laboratory trials, we examined the effects of introduced brown trout, Salmo trutta, on the short-term behaviour of a threatened, lake-dwelling galaxiid fish and confirmed a suspected diel pattern in habitat use by this species. We found that Galaxias auratus followed a distinct diel pattern in the use of complex habitats and open water, which was significantly altered by the presence of brown trout. In trials without the introduced predator, G. auratus used complex habitats (rocks or macrophytes) during the day, and open water during the night. In trials with brown trout present, G. auratus spent significantly less time in open water and rarely ventured out of the macrophytes. However, when given the option of using only rocky substrate or open water, which is the more common situation in the lakes to which this galaxiid is endemic, the fish reduced the amount of time they spent in the open water during the night, but still spent more time in open water than when macrophytes were available. Spending the daylight hours amongst the cover of rocks or macrophytes is most likely an adaptation to reduce the risk of predation by visual predators, and the pattern of reduced use of open water habitats in the presence of brown trout is an acute response to the close proximity of the introduced predator. The difference in the nocturnal use of macrophytes and rocks when trout are present may be related to differences in feeding opportunities or success within these habitats.     

Correlations between habitat use and body shape in a phrynosomatid lizard (Urosaurus ornatus): a population-level analysis

Recent ecomorphological studies have shown that the predicted correlations between morphology and ecology on broad taxonomic levels are often obscured when comparing more closely related groups. Among species, comparisons of lizards often indicate very little support for adaptive radiations into novel habitats. As few population level studies have been performed, we compared body, head and limb shape between four populations of Urosaurus ornatus living in structurally distinct habitats (cliffs, rocks, trees and boulders). Surprisingly, clear correlations between habitat use and body shape among populations were found, most of which were in good accordance with a priori biomechanical predictions (e.g. flat body and head for extreme climbers; long distal hindlimb segments for jumpers and runners; narrow body and long tail for tree dwelling lizards). This indicates that populations of Urosaurus ornatus are seemingly 'adapted' to the habitat they live in. However, quantification of performance and behaviour are needed to determine the adaptive nature of these observations.     

Foraging habits in a generalist predator: Sex and age influence habitat selection and resource use among bottlenose dolphins (Tursiops truncatus)

This study examines resource use (diet, habitat use, and trophic level) within and among demographic groups (males, females, and juveniles) of bottlenose dolphins (Tursiops truncatus). We analyzed the δ¹³C and δ¹⁵N values of 15 prey species constituting 84% of the species found in stomach contents. We used these data to establish a trophic enrichment factor (TEF) to inform dietary analysis using a Bayesian isotope mixing model. We document a TEF of 0‰ and 2.0‰ for δ¹³C and δ¹⁵N, respectively. The dietary results showed that all demographic groups relied heavily on low trophic level seagrass‐associated prey. Bayesian standard ellipse areas (SEA_b) were calculated to assess diversity in resource use. The SEA_b of females was nearly four times larger than that of males indicating varied resource use, likely a consequence of small home ranges and habitat specialization. Juveniles possessed an intermediate SEA_b, generally feeding at a lower trophic level compared to females, potentially an effect of natal philopatry and immature foraging skills. The small SEA_b of males reflects a high degree of specialization on seagrass associated prey. Patterns in resource use by the demographic groups are likely linked to differences in the relative importance of social and ecological factors.     

Avian diversity in a Kenyan agroecosystem: effects of habitat structure and proximity to forest

Tropical agroecosystems cover an increasingly large proportion of the Earth’s terrestrial surface. Yet, relatively little is known about the factors that influence their avifauna, especially in areas of high human population density. The potential of tropical farmland for sustaining bird biodiversity, including forest birds, can be influenced by habitat structure and the distance from the nearest forest. We investigated the effect of these two factors on the bird community in the farmland near Kakamega Forest, Kenya. Using point counts, we assessed the number of bird species and individuals on 56 study plots in distances up to about 2,100 m from the forest. We observed a total of 96 bird species in the farmland, 22 of which were forest, 58 shrub-land, and 16 open-country species. High vertical vegetation heterogeneity and a large number of woody plant individuals were related to high species richness of forest and shrub-land birds, whereas open-country birds avoided such areas. The species richness and total number of forest birds declined with increasing distance to the forest. A comparison with the bird community within Kakamega Forest indicated that only a fraction of the forest species could be sustained in the farmland. This suggests that agroecosystems with a diverse habitat structure can support a high diversity of birds, but have only a limited capacity to compensate for forest loss. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Modelling the longitudinal distribution,abundance, and habitat use of the giant freshwater shrimp (Macrobrachium spinipes) in a large intermittent,tropical Australian river to inform water resource policy

1. Water development threatens rivers and their biodiversity. Amphidromous shrimp are particularly vulnerable as they require migration between freshwater and estuaries to complete their life cycle. The Fitzroy River is a large tropical intermittent river undergoing water development that is home to the amphidromous shrimp Macrobrachium spinipes (cherabin), yet little is known about its habitat use and flow-ecology making it difficult to inform sustainable water-take.
2. We investigated habitat associations, distributional patterns suggestive of amphidromy, and the influence of water availability by sampling main channel and floodplain pools along a 350-km river length during 2 contrasting flow years. Applying a size-specific abundance model, we estimated abundance per size class, site, and year. We then predicted abundance at the landscape scale with remotely sensed water to reveal the impact of water availability on the meta-population.
3. Our model revealed that juveniles were in greatest abundance in downstream main channel pools, whereas adults were in greatest abundance in upstream floodplain pools. Abundance varied by year with lower numbers predicted in the low-flow year. Longitudinal and habitat patterns remained when our pool-level results were scaled to the landscape, and the positive relationship of abundance to wet-season flow was strengthened. The predominance of smaller cherabin in the lower reaches of the river provides indirect support for an estuarine nursery and amphidromous life history; however, small individuals observed in landlocked pools, during late dry season suggests possible within-river recruitment.
4. The importance of water development policies that protect wet-season flow and passage along the Fitzroy River is supported by this work. These types of policies are likely to be important for this and other amphidromous shrimp species across Australia, Southeast Asia and further afield. Further research detailing the species life history and describing flow–recruitment relationships will be important contributions to understanding this important taxonomic group and refining policies for current and future water resource development.

     

Size and shape variation in a Lesser Antillean anole, Anolis oculatus (Sauria: Iguanidae) in relation to habitat.

The anole fauna of the Lesser Antilles is depauperate in relation to that of the Greater Antilles, where complex communities characterized by adaptive specialization and convergent structure are present. Much of this adaptation is the result of changes in body size and shape, probably as a result of interspecific competition. Here we present data on variation in size and shape within a solitary Lesser Antillean species occupying an ecologically heterogenous island, and test the hypothesis that natural selection for varying environmental conditions is the cause of this variation. Univariate (analysis of variance), bivariate (analysis of covariance) and multivariate (multiple-group principal component analysis, canonical variate analysis) analysis showed that there is considerable geographic variation in size and shape within Anolis oculatus on Dominica. Partial Mantel tests rejected the null hypothesis of no association between size and shape and environmental variation. The possible proximate and evolutionary mechanisms responsible for these patterns are discussed. Despite these overall associations, a considerable amount of variation in shape appears to be unrelated to environmental variation.