SPECIES FLOCK IN THE NORTH AMERICAN GREAT LAKES: MOLECULAR ECOLOGY OF LAKE NIPIGON CISCOES (TELEOSTEI: COREGONIDAE: COREGONUS)

Studies on north temperate fish species indicate that new habitat availability following the last ice sheet retreat has promoted ecological speciation in postglacial lakes. Extensive ecophenotypic polymorphisms observed among the North American Great Lakes ciscoes suggest that this fish group has radiated through trophic adaptation and reproductive isolation. This study aims at relating the ecomorphological and genetic polymorphisms expressed by the Lake Nipigon ciscoes to evaluate the likelihood of an intralacustrine divergence driven by the exploitation of alternative resources. Morphological variation and trophic and spatial niches are characterized and contrasted among 203 individuals. Genetic variation at six microsatellite loci is also analyzed to appraise the extent of genetic differentiation among these morphotypes. Ecomorphological data confirm the existence of four distinct morphotypes displaying various levels of trophic and depth niche overlap and specialization. However, ecological and morphological variations were not coupled as expected, suggesting that trophic morphology is not always predictive of ecology. Although extensive genetic variability was observed, little genetic differentiation was found among morphotypes, with only one morph being slightly but significantly differentiated. Contrasting patterns of morphological, ecological, and genetic polymorphisms did not support the hypothesis of ecological speciation: the most ecologically different forms were morphologically most similar, while the only genetically differentiated morph was the least ecologically specialized. The low levels of genetic differentiation and the congruence between θ and φ estimates altogether suggest a recent (most likely postglacial) process of divergence and/or high gene flow among morphs A, C, and D, whereas higher φ estimates for comparison involving morph B suggest that this morph may be derived from another colonizing lineage exchanging little genes with the other morphs. Patterns of ecophenotypic and genetic diversity are also compatible with a more complex evolutionary history involving hybridization and introgression.     

Dietary and Temporal Niche Differentiation in Tropical Ants—Can They Explain Local Ant Coexistence?

How species with similar ecological requirements avoid competitive exclusion remains contentious, especially in the species‐rich tropics. Niche differentiation has been proposed as a major mechanism for species coexistence. However, different niche dimensions must be studied simultaneously to assess their combined effects on diversity and composition of a community. In most terrestrial ecosystems, ants are among the most abundant and ubiquitous animals. Since they display direct, aggressive competition and often competitively displace subordinate species from resources, niche differentiation may be especially relevant among ants. We studied temporal and trophic niche differentiation in a ground ant community in a forest fragment in French Guiana. Different baits were presented during day and night to assess the temporal and dietary niches of the local species. They represented natural food resources such as sugars, carrion, excrements, seeds, and live prey. In addition, pitfalls provided a background measure of ant diversity. The communities attracted to the different baits significantly differed from each other, and even less attractive baits yielded additional species. We detected species specialized on living grasshoppers, sucrose, seeds, or dead insects. Community‐level differences between day and night were larger than those between baits, and many species were temporally specialized. In contrast to commonness, foraging efficiency of species was correlated to food specialization. We conclude that many ant species occupy different temporal or dietary niches. However, for many generalized species, the dietary, and temporal niche differentiation brought forward through our sampling effort, cannot alone explain their coexistence.     

Functional traits reveal the expansion and packing of ecological niche space underlying an elevational diversity gradient in passerine birds

Variation in species richness across environmental gradients may be associated with an expanded volume or increased packing of ecological niche space. However, the relative importance of these alternative scenarios remains unknown, largely because standardized information on functional traits and their ecological relevance is lacking for major diversity gradients. Here, we combine data on morphological and ecological traits for 523 species of passerine birds distributed across an Andes-to-Amazon elevation gradient. We show that morphological traits capture substantial variation in species dietary (75%) and foraging niches (60%) when multiple independent trait dimensions are considered. Having established these relationships, we show that the 14-fold increase in species richness towards the lowlands is associated with both an increased volume and density of functional trait space. However, we find that increases in volume contribute little to changes in richness, with most (78%) lowland species occurring within the range of trait space occupied at high elevations. Taken together, our results suggest that high species richness is mainly associated with a denser occupation of functional trait space, implying an increased specialization or overlap of ecological niches, and supporting the view that niche packing is the dominant trend underlying gradients of increasing biodiversity towards the lowland tropics.     

Ecomorphology: Experimental Functional Anatomy for Ecological Problems

It is generally believed that the functional design of an organismrelates to its ecology, yet this ecomorphological paradigm hashistorically suffered from the lack of a rigorous frameworkfor its implementation. I present a methodology for experimentallyexploring the ecological consequences of variation in morphology.The central idea is that morphology influences ecology by limitingthe ability of the individual to perform key tasks in its dailylife. Inthis scheme the effect of morphological variation onbehavioral performance is first tested in laboratory experiments.As the behavioral capability of an individual defines the rangeof ecological resources that it can potentially make use of(the potential niche), the second step in the scheme involvescomparing the potential niche of an individual to actual patternsof resource use (the realized niche). This permits a quantitativeassessment of the significance of an organism's maximal capabilitiesin determining actual patterns of resource use. An example is presented from work on the feeding biology offishes in the family Labridae (wrasses and parrotfishes). Mostlabrids feed by crushing shelled prey in their powerful pharyngealjaws. This example explores the dietary consequences of variationin crushing strength amongand within species. Crushing strengthwas estimated from biomechanical analyses of the crushingapparatusin several species, and these predictions of relative strengthwere tested in laboratory feeding experiments with hard-shelledprey. Morphology accurately predicted relative crushing ability,and the final section of the study explored the effect of variationin crushing ability on diet. Within each of three species crushingstrength appears to underlie a major ontogentic dietary switchfrom soft-bodied prey to a diet dominated by hard-shelled prey.In each species this switch occurred at about the same crushingstrength, around 5 Newtons (N), in spite of the fact that thiscrushing strength is achieved by the three species at differentbody sizes. Diet breadth increases during ontogeny in each species,until a crushing strength of 5 N is achieved, when diet breadthbegins to decline. The strongest fishes specialized almost entirelyon molluscs and sea urchins. Thus, these labrids take advantageof ontogenetic and interspecific differences in crushing strengthby including harder and harder prey in their diet, and ultimatelyspecializing on hard prey types. The specialized organizationof the labrid pharyngeal jaws can be viewed as a key innovationthat has permitted this lineage of fishes to invade the mollusceating niche, a relatively empty trophic niche within the highlyspeciose and diverse communities of coral reef fishes.     

Morphological-dietary relationships within two assemblages of marine demersal fishes

In this study we have used morphological characters related to feeding and prey capture and dietary data to investigate the trophic organization within two assemblages of marine demersal fishes. Morphological and dietary disparities within fish assemblages were estimated from species similarities based on Euclidean distances plus species projections on the principal axes from multivariate analyses. The analyses of the morphological variables indicated that species in each assemblage comprised morphologically distinct groups strongly influenced by trophically linked characters. Stomach content analyses revealed that fish species in each assemblage were classified into three basic feeding groups: polychaete-shrimp feeders, crab feeders and fish feeders. These results indicated that food resource partitioning was operating within each assemblage. However, when morphological and trophic data were compared no significant correlations were found. The results did not particularly support the ecomorphological hypothesis that dietary differences are due to morphological differences, since similar diets do not correspond to similar morphologies. The patterns of trophic organization within the fish assemblages examined, possibly reflect differences among species due not only to the effect of ecological demands on morphology but also by their evolutionary history and constructional constraints imposed by phylogeny. This revised version was published online in August 2006 with corrections to the Cover Date.     

The coexistence of fish species in streams: relationships between assemblage attributes and trophic and environmental variables

Diet overlap and niche breadth are well-known species traits from trophic ecology that can assist in explaining how species interact and coexist as well as the ecological mechanisms that influence biodiversity. In the present study, we analyzed the relationships between these trophic variables and indicators of resource availability with some attributes of fish assemblages (species richness, Shannon diversity index, evenness, density and individual body size). The physical and chemical characteristics of the biotopes (topography, water quality and conservation of slopes) were examined to identify possible patterns. Monthly sampling using electrofishing was conducted in 2003 along five streams located in the Cuiabá River watershed. The relationships between environmental variables and attributes of fish assemblages were evaluated using Spearman correlation. Species richness and abundance varied among streams, with higher values (54 and 82 species) found in low-gradient streams that drained small swampy areas discharging in Cuiabá River. Diet overlap showed significant and negative correlations with species richness, the diversity index, abundance, variation in body size and food availability and positive correlations with evenness and niche breadth. Environments that had greater food availability had a reduced variety of food items (smaller amplitude) and distinct values in terms of dietary overlap. Nevertheless, to explain resource partitioning in a fish assemblage (overlap), it is necessary to consider food availability and niche breadth (degree of trophic specialization) beyond the interaction of these variables with species richness and density. In conclusion, high diversity and abundance values were strongly associated with a high degree of trophic specialization (low amplitude of trophic niche) and a small degree of overlap in the diet.     

Feeding ecology and trophic segregation of two sympatric mesopredatory sharks in the heavily exploited coastal ecosystem of the Adriatic Sea

The ecological roles and trophic interactions of two commercially important mesopredatory shark species, Squalus acanthias and Mustelus punctulatus that co‐occur on the continental shelf of the north‐central Adriatic Sea were investigated. Both shark species are dietary specialists, with a significant dietary overlap recorded only during the spring season. They showed different patterns of feeding as they grew: S. acanthias extended its trophic niche with an increase in size, while M. punctulatus developed a more specialized diet. These two sharks partition food resources and reduce niche overlap by foraging at different trophic levels. Mustelus punctulatus is a crustacean feeder, specialized in foraging on scavenging malacostracans frequently found along trawl tracks or on discards in the Adriatic fishing zone. Conversely, S. acanthias prefers small pelagic fishes, which are commercially exploited and in decline. The different foraging strategies adopted by these two species suggest that they should be managed separately. Dietary specialization, direct competition with humans for prey and their higher intrinsic vulnerability make S. acanthias particularly susceptible to the effects of anthropogenic perturbations.     

Functional versatility supports coral reef biodiversity

We explore the role of specialization in supporting species coexistence in high-diversity ecosystems. Using a novel ordination-based method to quantify specialist and generalist feeding structures and diets we examined the relationship between morphology and diet in 120 wrasses and parrotfishes from the Great Barrier Reef. We find that wrasses, despite their morphological diversity, exhibit weak links between morphology and diet and that specialist morphologies do not necessarily equate to specialized diets. The dominant pattern shows extensive overlap in morphology (functional morphospace occupation) among trophic groups; fish with a given morphology may have a number of feeding modes. Such trophic versatility may lay the foundation for both the origins and maintenance of high biodiversity on coral reefs.     

Linking habitat specialization with species' traits in European birds

Ecological specialization provides information about adaptations of species to their environment. However, identification of traits representing the relevant dimensions of ecological space remains challenging. Here we endeavoured to explain how complex habitat specializations relate to various ecological traits of European birds. We employed phylogenetic generalized least squares and information theoretic approach statistically controlling for differences in geographic range size among species. Habitat specialists had narrower diet niche, wider climatic niche, higher wing length/tail length ratio and migrated on shorter distances than habitat generalists. Our results support an expected positive link between habitat and diet niche breadth estimates, however a negative relationship between habitat and climate niche breadths is surprising. It implies that habitat specialists occur mostly in spatially restricted environments with high climatic variability such as mountain areas. This, however, complicates our understanding of predicted impacts of climatic changes on avian geographical distributions. Our results further corroborate that habitat specialization reflects occupation of morphological space, when specialists depend more on manoeuvrability of the flight and are thus more closely associated to open habitats than habitat generalists. Finally, our results indicate that long distance movements might hamper narrow habitat preferences. In conclusion, we have shown that species’ distributions across habitats are informative about their positions along other axes of ecological space and can explain states of particular functional traits, however, our results also reveal that the links between different niche estimates cannot be always straightforwardly predicted.     

Community structure affects trophic ontogeny in a predatory fish

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout‐only systems, (ii) two‐species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three‐species systems (brown trout, Arctic charr, and three‐spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor–prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout‐only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three‐species communities. Our findings revealed that the presence of a small‐sized prey fish species (stickleback) rather than a mixed competitor–prey fish species (charr) was an important factor affecting the ontogenetic niche‐shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.     

Weak phylogenetic effects on ecological niches of Sylvia warblers

To understand the evolution of ecological niches it is important to know whether niche evolution is constrained by phylogeny. We approached this question for Sylvia warblers by testing if closely related species are more similar in 20 ecologically relevant morphological traits than distantly related species. Phylogenetic relatedness was quantified using a molecular phylogeny based on the mitochondrial cytochrome b gene. By Principal Component Analysis (PCA) two major niche axes were extracted. We tested the individual ecomorphological traits and the positions of the species on the PCA axes for phylogenetic effects using Mantel tests. The results demonstrated small but significant phylogenetic effects only for the length of the middle toe, a trait probably correlated with locomotion. In general, however, phylogenetic effects were very weak. This suggests that ecological niches in passerine birds have the potential to evolve rapidly and are not subject to major phylogenetic constraints.     

Effects of hummingbird morphology on specialization in pollination networks vary with resource availability

Specialization of species in interaction networks influences network stability and ecosystem functioning. Spatial and temporal variation in resource availability may provide insight into how ecological factors, such as resource abundance, and evolutionary factors, such as phylogenetically conserved morphological traits, influence specialization within mutualistic networks. We used independent measures of hummingbird abundance and resources (nectar), information on hummingbird traits and plant–hummingbird interactions to examine how resource availability and species' morphology influence the specialization of hummingbirds in three habitat types (forest, shrubs, cattle ranch) sampled over 10 sessions across two years in the southern Andes of Ecuador. Specialization of hummingbird species in the networks was measured by three indices: d' (related to niche partitioning), generality (related to niche width) and PSI (related to pollination services). Specialization indices d', generality and PSI of hummingbird species were influenced by resource availability. All indices indicated that specialization of hummingbirds increased when the availability of resources decreased. Variation in d' was also explained by an interaction between resource availability and bill length; hummingbirds with a long bill switched from being more specialized than other species when resource availability was low to being similarly specialized when availability was high. Overall, our results highlight the importance of ecological and evolutionary factors determining the specialization of species in interaction networks. We demonstrate in particular that ecological gradients in resource availability cause substantial changes in consumers' foraging behavior contingent on their morphology. Changes in pollinator specialization along resource gradients can have impacts on ecosystem functions, such as pollination by animals.     

Evolution of microhabitat association and morphology in a diverse group of cryptobenthic coral reef fishes (Teleostei: Gobiidae: Eviota)

Gobies (Teleostei: Gobiidae) are an extremely diverse and widely distributed group and are the second most species rich family of vertebrates. Ecological drivers are key to the evolutionary success of the Gobiidae. However, ecological and phylogenetic data are lacking for many diverse genera of gobies. Our study investigated the evolution of microhabitat association across the phylogeny of 18 species of dwarfgobies (genus Eviota), an abundant and diverse group of coral reef fishes. In addition, we also explore the evolution of pectoral fin-ray branching and sensory head pores to determine the relationship between morphological evolution and microhabitat shifts. Our results demonstrate that Eviota species switched multiple times from a facultative hard-coral association to inhabiting rubble or mixed sand/rubble habitat. We found no obvious relationship between microhabitat shifts and changes in pectoral fin-ray branching or reduction in sensory pores, with the latter character being highly homoplasious throughout the genus. The relative flexibility in coral-association in Eviota combined with the ability to move into non-coral habitats suggests a genetic capacity for ecological release in contrast to the strict obligate coral-dwelling relationship commonly observed in closely related coral gobies, thus promoting co-existence through fine scale niche partitioning. The variation in microhabitat association may facilitate opportunistic ecological speciation, and species persistence in the face of environmental change. This increased speciation opportunity, in concert with a high resilience to extinction, may explain the exceptionally high diversity seen in Eviota compared to related genera in the family.     

Ecological structure of tropical fish assemblages in wet-zone streams of Sri Lanka

The introduction of four fish species into a depauperate stream system in central Sri Lanka provided a 'natural experiment' that enabled us to determine the ecological structure of wet-zone stream fish assemblages. All the species indigenous to this 'introduction' stream system also co-occur naturally with the introduced species in nearby streams. Analyses of habitat use and dietary requirements revealed that most species in the assemblages were segregated on the basis of macrohabitat, microhabitat and food, regardless of origin of the fishes. Macrohabitat and microhabitat utilizations by species were similar in each stream of the 'introduction' system despite differences in stream conditions. Thus species distributions, relative to each other in ecological space, were consistent among streams, and with the combined data. High overlaps along the resource axes of velocity, depth, substratum and food were few. A major proportion of the high overlaps was due to associations with introduced species; especially Barbus nigrofasciatus and Barbus cumingii. The other two introduced species, Barbus titteya and Rasbora vaterifloris , were more specialized, and interacted relatively less with indigenous species. The indigenous species, however, exhibited pronounced complementarity along three resource axes representing depth, velocity and food. Rasbora vaterifloris and B. nigrofasciatus grew to larger sizes in one stream which had fewer indigenous species, suggesting competitive release. In this stream, the most common indigenous species was a dietary specialist that fed on diatoms, and B. nigrofasciatus , which fed heavily on diatoms in other streams, switched to feeding more on macrophytes. Overall, the data suggest that these assemblages are predictable, co-evolved systems with competition serving as an important structuring force that reinforces species segregation.     

Habitat utilization by coral reef fish: implications for specialists vs. generalists in a changing environment

1. The impact of environmental disturbance and habitat loss on associated species is expected to be dependent on a species' level of specialization. We examined habitat use and specialization of coral reef fish from the diverse and ecologically important family Pomacentridae, and determined which species are susceptible to declines in coral cover due to disturbance induced by crown-of-thorns seastar (COTS, Acanthaster planci L.). 2. A high proportion of pomacentrid species live in association with live coral as adults (40%) or juveniles (53%). Adults of many species had strong affiliations with branching corals, while juveniles favoured plating growth forms, reflecting the sizes of refuge provided by coral types. 3. Juveniles of species that associated with coral had narrower niche breadths than adult conspecifics, due to associations with specific coral types. The especially high coral association and narrower niche breadth of juveniles suggest that the presence of live coral is crucial for many species during early life history, and that disturbance-induced coral loss may have serious flow-on effects on adult abundance. 4. Microhabitat availability was a poor predictor of fish species abundance. Significant correlations between coverage of coral types and abundance of five adults and two juvenile species were detected; however, these relationships explained <35% and <10% of the variation in abundance of adult and juvenile species, respectively. 5. Niche breadth explained 74% of the variation in species' mean response to coral decline and it is clear that disturbance has a greater impact on resource specialists, suggesting that increasing frequency and intensity of coral loss will cause reef fish communities to become dominated by habitat generalists at the expense of coral-dwelling specialists.     

Effects of an adaptive zone shift on morphological and ecological diversification in terapontid fishes

A fundamental goal of evolutionary ecology is understanding the processes responsible for contemporary patterns of morphological diversity and species richness. Transitions across the marine–freshwater interface are regarded as key triggers for adaptive radiation of many clades. Using the Australian terapontid fish family as a model system we employed phylogenetic analyses to compare the rates of ecological (dietary) and morphological evolution between marine and freshwater species of the family. Results suggested significantly higher rates of phenotypic evolution in key dietary and morphological characters in freshwater species compared to marine counterparts. Moreover, there was significant correlation between several of these dietary and morphological characters, suggesting an underlying ecomorphological aspect to these greater rates of phenotypic evolution in freshwater clades. Australia’s biogeographic history, which has precluded colonisation by many of the major ostariophysan fish families that make up much global freshwater fish diversity, appears to have provided the requisite ‘ecological opportunity’ to facilitate the radiation of invading marine-derived fish clades.     

Dietary partitioning promotes the coexistence of planktivorous species on coral reefs

Theories involving niche diversification to explain high levels of tropical diversity propose that species are more likely to co‐occur if they partition at least one dimension of their ecological niche space. Yet, numerous species appear to have widely overlapping niches based upon broad categorizations of resource use or functional traits. In particular, the extent to which food partitioning contributes to species coexistence in hyperdiverse tropical ecosystems remains unresolved. Here, we use a molecular approach to investigate inter‐ and intraspecific dietary partitioning between two species of damselfish (Dascyllus flavicaudus, Chromis viridis) that commonly co‐occur in branching corals. Species‐level identification of their diverse zooplankton prey revealed significant differences in diet composition between species despite their seemingly similar feeding strategies. Dascyllus exhibited a more diverse diet than Chromis, whereas Chromis tended to select larger prey items. A large calanoid copepod, Labidocera sp., found in low density and higher in the water column during the day, explained more than 19% of the variation in dietary composition between Dascyllus and Chromis. Dascyllus did not significantly shift its diet in the presence of Chromis, which suggests intrinsic differences in feeding behaviour. Finally, prey composition significantly shifted during the ontogeny of both fish species. Our findings show that levels of dietary specialization among coral reef associated species have likely been underestimated, and they underscore the importance of characterizing trophic webs in tropical ecosystems at higher levels of taxonomic resolution. They also suggest that niche redundancy may not be as common as previously thought.     

Rarity in Australian desert lizards

Most species of Australian desert lizards are uncommon. Possible causes of rarity are examined, including body size as measured by snout–vent length (SVL), fecundity, number of sites occupied, habitat niche breadth, microhabitat niche breadth, dietary niche breadth, and average total niche overlap with other species. Rare species tend to be larger with lower fecundities than abundant species and they occur at fewer sites. Many, but not all, uncommon species are specialists, either in habitat, microhabitat, or diet. The niche breadth hypothesis, which states that abundant species should be generalists whereas specialized species should be rare, is tested, but rejected as a general explanation for rarity. Some uncommon species exhibit high overlap with other species suggesting that they may experience diffuse competition. However, no single cause of rarity can be identified, but rather each species has its own idiosyncratic reasons for being uncommon. Multivariate analyses show distinct ecological differences between abundant and uncommon species.     

Does morphology predict behavior? Correspondence between behavioral and morphometric data in a Tyrant‐flycatcher (Tyrannidae) assemblage in the Santa Marta Mountains,Colombia

ABSTRACT Morphology is commonly used as a predictor of ecological relationships among species when studying local assemblages of Neotropical birds. Nevertheless, most evidence supporting ecomorphological correspondence in birds comes from studies of communities and not from local assemblages and, moreover, from temperate latitudes. To increase our understanding of ecomorphological correspondence in Neotropical assemblages, we used three multivariate approaches to evaluate correspondence between morphological and foraging behavior data in a tyrant‐flycatcher assemblage (N= 12 species) in the Santa Marta Mountains in Colombia. Principal components analyses revealed similar species ordinations when using morphological measurements (beak size and shape, tarsus length, wing length, and tail length) or behavioral data (behavioral types of searching for prey and prey capture) separately. Discriminant function analyses tested the ability of morphological traits to predict foraging behavior, showing that more than 90% of all measured individuals (N= 267) were correctly classified in previously defined categories of search and attack behavior. Finally, Canonical correlation analyses revealed a significant correlation between morphological data and two independent datasets of search and attack behavior. Our results demonstrate that morphology can accurately predict ecology in an assemblage of Neotropical tyrannids, and similar results have been reported in previous studies of temperate Tyrant‐flycatchers. Our results also show that bill size and shape, wing length, and tarsus length are the best predictors of foraging behavior in this assemblage. Testing for ecomorphological correspondence in other Neotropical taxa would help identify subsets of phenotypic traits that could be used for a practical, but reliable, determination of ecological relationships within different assemblages.