Distinguishing trophic and habitat partitioning among sympatric populations of the estuarine fish Osmerus mordax Mitchill

Sympatric populations of rainbow smelt Osmerus mordax of the St Lawrence middle estuary exhibited distinct morphologies suitable for exploiting either pelagic or benthic habitats in the absence of consistent morphological features related to specific diets. The magnitude of morphological differentiation was elevated and constant over 3 years even though statistically significant differences were observed among samples within populations. Both populations were spatially segregated in the estuary but some population mixing occurred at low densities of rainbow smelt. Diet analysis revealed that both populations opportunistically feed on large macrozooplankton. An amended version of Schoener's D was developed and used to compare the magnitude of niche overlap. It revealed that the diet of the middle estuary rainbow smelt differed according to the area where samples were collected but no differences existed between populations. The exploitation of ecologically distinct habitats appeared to be effective in maintaining morphologically distinct sympatric populations within an estuarine setting in the absence of diet differentiation.     

Niche partitioning and morphospace in early stages of two sympatric Diogenichthys species (Myctophidae)

Diet and morphospace of larval stages of two sympatric lanternfish Diogenichthys atlanticus and D. laternatus from the south-east Pacific Ocean were compared and the covariance between both variables was assessed for each species. Diogenichthys atlanticus stomach contents consisted mainly of copepod nauplii and digested remains and this species had a broader niche than D. laternatus, in which stomach contents were highly digested. No dietary overlap was found between both species. The covariance between skull shape and diet for D. atlanticus was given by a wider mouth gape related to the presence of copepod nauplii, whilst for D. laternatus, a shorter snout and posteriorly displaced eye were related to the presence of highly digested stomach contents. Interspecific differences between diets and skull shapes suggest that both species may have undergone morphological or niche divergence to avoid competition, such as feeding at different hours or depth stratification.     

Niche breadth and resource partitioning by four sympatric species of bark beetles (Coleoptera: Scolytidae)

Summary In a North Carolina population of Diamorpha smallii, densities were artificially manipulated by thinning stands to predetermined densities. Individuals in areas of high density produce fewer seeds than individuals at low density in the same habitat. Three components of fecundity, the number of flowers per plant, the number of fruits per plant, and the number of seeds per fruit, were negatively correlated with density. The relationship of components of fecundity to density were used to develop a model of population regulation for this species. These results and predictions from the model are discussed in relation to other density studies.     

Multisite and multispecies measures of overlap,co‐occurrence,and co‐diversity

Several indices measure the association or segregation between two species and the similarity or differentiation between two sets of species. These indices are based on the overlap in the distribution of species (measured with the number of co‐occurrences) or on the overlap in species composition of sites (measured with the number of species that are shared between two sites). This paper shows that when evaluating more than two species the number of overlaps and the number of pairwise co‐occurrences are not equal, as it is the case for two species. Equivalently, when comparing more than two species assemblages, the number of overlaps differ from the number of instances of species sharing by pairs of sites (the ‘co‐diversities’). Considering this distinction, two different types of multispecies and multisite indices can be derived: indices of general overlap and indices of co‐occurrence or co‐diversity. Here I present a complete series of the two types of indices that correspond to the popular Jaccard, Sørensen, and Simpson two‐species or two‐site indices. Indices of general overlap are defined by three parameters (the total number of species, the total number of sites, and the total number of occurrences), whereas indices of co‐occurrence or co‐diversity depend on those parameters plus an additional one that is defined by the values of species richness or range size. Consequently, the two types of indices respond differently to null models, depending on the parameters that are fixed or randomized. All indices correlate well with the mean of the traditional indices calculated pair by pair, and the correspondence is extremely close for the new indices of co‐occurrence and co‐diversity. These properties should be useful in clarifying some of the confusion that exists in the current discussion about the advantages and disadvantages of pairwise vs community‐wide approaches in the analysis of diversity.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Habitat and resource partitioning among Indo‐Pacific bottlenose dolphins in Moreton Bay,Australia

Investigating resource partitioning among mobile marine predators such as cetaceans is challenging. Here we integrate multiple methodologies (analyses of habitat use, stable isotopes and trace elements) to assess ecological niche partitioning amongst two genetically divergent sympatric subpopulations (North and South) of Indo‐Pacific bottlenose dolphins (Tursiops aduncus) in Moreton Bay, Australia. Comparisons of the mean locations (latitude, longitude) and environmental variables (distance from sandbanks, distance from shore and water depth) observed at sightings of biopsy‐sampled individuals indicated that the North subpopulation occurred in the northwestern bay in significantly deeper water than the South subpopulation, which was found in southeastern nearshore waters and closer to sandbanks. Ratios of stable carbon and nitrogen isotopes in skin samples suggested that North dolphins foraged on higher trophic level prey in relatively more pelagic, offshore habitats, while South dolphins foraged on lower trophic prey in more nearshore, demersal and/or benthic habitats. Habitat partitioning was also reflected in higher blubber concentrations of most of the 13 measured trace elements, in particular lead, in the coastal South compared to the more pelagic North dolphins. These findings indicate that genetic subpopulations of bottlenose dolphins in Moreton Bay are adapted to different niches.     

Strategies of resource partitioning between two sympatric puffer fishes in a tropical hypersaline estuary,Brazil

The strategies of food partitioning among two abundant estuarine puffer fishes, Sphoeroides greeleyi (Gilbert, 1900) and Sphoeroides testudineus (Linnaeus, 1758), were investigated in a tropical hypersaline estuary. We examined the stomachs of 946 fishes collected with a beach seine across three zones (upper, middle and lower estuary) along with a salinity gradient. The highest abundances of species were recorded in the upper and middle estuary. The diet was comprised mostly by benthic organisms, mainly Bivalvia, Gastropoda, and Brachyura. However, in a hypersaline estuary, these species develop strategies of resource partitioning in three ways: 1) asymmetry effects; 2) habitat selection; and 3) trade-offs among age classes. Diet variations according to fish age classes indicated that the largest individuals showed a decrease in their consumption of small preys (Amphipods, Copepods, and Ceratopogonidae) and an increase in their consumption of larger preys (Bivalvia and Decapoda). This behavior of switching diet towards larger prey was related to the functional trade-offs in swimming capacity, and feeding mode used to capture prey. The middle and upper estuary are important feeding grounds for puffer fishes, as demonstrated by their influence on prey distributions and habitat type. Diet breadth indicated that S. greeleyi tended to be a generalist, whereas S. testudineus tended to be a specialist. Thus, the strategies of partitioning seem to have importance for puffer fish populations in this hypersaline environment.     

Resource partitioning of sympatric small mammals in an African forest‐grassland vegetation mosaic

It is often hypothesized that two species competing for the same resource cannot stably coexist unless they partition their resources in space and time. More recently stable isotope analyses have complemented traditional, observation‐based niche research by conceptualizing many of the characteristics of communities, for example, trophic niche width and the partitioning of resources. Here we quantify resource partitioning of sympatric small mammal species in an African ecosystem by analysing stable isotope ratios of hair collected from a South African forest‐grassland vegetation mosaic, and combine this with known spatial and temporal behavioural data to interpret community competition and resource partitioning. We observe niche separation to different degrees across the entire community, with different species displaying either unique isotopic dietary preferences, or partitioning resources in space and/or time. δ¹³C values were more enriched in species that inhabited afromontane grassland compared with those that inhabited afromontane forest, a reflection of the dominant vegetation in each habitat. Contrary to expectations, arboreal rodents occupied higher trophic positions than terrestrial rodents and approaching δ¹⁵N values similar to insectivorous shrews, suggesting that arboreal rodents feed on items such as arthropods enriched in ¹⁵N. While grassland species display phenotypic plasticity in terms of dietary preferences, small mammals that occurred in forests display narrow niche preferences, suggesting these species may be particularly sensitive to habitat modifications. Our results illustrate that the use of stable isotopes can be used in conjunction with spatial and temporal behavioural knowledge to elucidate resource partitioning in small African mammal communities.     

Diet overlap and resource partitioning among three forage fish species in Coorong,the largest inverse estuary in Australia

Protracted drought in southern Australia has degraded the system function of the Coorong estuary and intensified resource competition among forage fish species. The present study investigates the gut content, prey composition and dietary overlap of three forage fishes: smallmouth hardyhead (Atherinosoma microstoma), Tamar River goby (Afurcagobius tamarensis) and sandy sprat (Hyperlophus vittatus) influenced by environmental variation in the Murray Estuary and Coorong. The prey species identified in fish stomachs were dominated by crustaceans (amphipods, ostracods and harpacticoids), but nematodes and acanthocephalans were also common in all forage fishes. The diet of the sandy sprat and Tamar River goby highly overlapped (α = 0.8) in the Murray Estuary and all three forage fishes showed potential diet overlap (α ≥ 0.6) in the North Lagoon. Spatiotemporal variation of prey diversity was observed in smallmouth hardyhead while temporal variation of prey diversity was observed in sandy sprat and Tamar River goby. Overall, the prey abundance was temporally variable and predominantly regulated by salinity, pH, dissolved oxygen, water transparency and chlorophyll a in the Murray Estuary and Coorong. This study adds to our knowledge on dietary overlap and resource partitioning among small-bodied forage fishes mediated by environmental factors in the Murray Estuary and Coorong.     

Niche overlap and diet composition of three sympatric coastal dolphin species in the southwest Atlantic Ocean

Sympatric species are expected to differ in ecological requirements to minimize niche overlap and avoid competition. Here we assess the trophic interactions among three coexisting dolphin species from southern Brazil: the franciscana dolphin (Pontoporia blainvillei), the Guiana dolphin (Sotalia guianensis), and the Lahille's bottlenose dolphin (Tursiops truncatus gephyreus). We evaluated temporal variation in carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values of bone collagen to examine potential dietary shifts resulting from increased fishing activity over the past three decades. We estimated the degree of niche overlap among these species and the contribution of potential prey sources to their diet. δ¹⁵N values were consistent among species and across years, while δ¹³C values increased for Guiana dolphins and decreased for bottlenose dolphins, suggesting changes in diet and/or foraging habitats through time. The similar δ¹³C and δ¹⁵N values and the high niche overlap between Guiana and bottlenose dolphins indicate that these species are primarily feeding on demersal prey. The franciscana diet is primarily composed of pelagic prey, resulting in a lower niche overlap in comparison with the other dolphin species. Our study provides further information about the foraging ecology of this unique dolphin community in southern Brazil with implications for its management and conservation.     

Comparative phylogeography of two co‐distributed but ecologically distinct rainbowfishes of far‐northern Australia

Aim

To test the influence of historical and contemporary environment in shaping the genetic diversity of freshwater fauna we contrast genetic structure in two co‐distributed, but ecologically distinct, rainbowfish; a habitat generalist (Melanotaenia splendida) and a habitat specialist (M. trifasciata).

Location

Fishes were sampled from far northern Australia (Queensland and Northern Territory).

Methods

We used sequence data from one mitochondrial gene and one nuclear gene to investigate patterns of genetic diversity in M. splendida and M. trifasciata to determine how differences in habitat preference and historical changes in drainage boundaries affected patterns of connectivity.

Results

Melanotaenia splendida showed high levels of genetic diversity and little population structure across its range. In contrast, M. trifasciata showed high levels of population structure. Whereas phylogeographic patterns differed, both species showed a strong relationship between geographical distance and genetic differentiation between populations. Melanotaenia splendida showed a shallower relationship with geographical distance, and genetic differentiation was best explained by stream length and a lower scaled ocean distance (11.98 times coast length). For M. trifasciata, genetic differentiation was best explained by overwater distance between catchments and ocean distance scaled at 1.16 × 10⁶ times coast length.

Main conclusions

Connectivity of freshwater populations inhabiting regions periodically interconnected during glacial periods appears to have been affected by ecological differences between species. Species‐specific differences are epitomized here by the contrast between co‐distributed congeners with different habitat requirements: for the habitat generalist, M. splendida, there was evidence for greater historical genetic connectivity with oceans as a weaker barrier to gene exchange in contrast with the habitat specialist, M. trifasciata.     

The effects of morphology,phylogeny and prey availability on trophic resource partitioning in an anuran community

Several factors influence the partitioning of trophic resources in ecological communities, such as morphology, evolutionary history, and resource availability. Although the effects of morphology, phylogeny, and resource availability on trophic ecology have long been explored by theoretical studies, little has been done to empirically test these relationships. Here, we tested whether phylogenetic and morphological distances correlate with trophic niche overlap using a path analysis of multiple partial regression of distance matrices. Also, we tested whether niche breadth is influenced by body size using Phylogenetic Generalized Least Squares analysis. Trophic niche overlap was better explained by morphology per se than by the phylogenetic distance. We also found that predator's body size influences niche breadth calculated considering prey traits and availability, but not when we do not include these availability data. Additionally, trophic niche breadth was usually smaller when we considered prey traits and availability, differently from niche overlap, whose values increased when we did not consider these data. Our findings show that the interpretation of trophic niche in communities changes if we consider availability data, affecting inferences about coexistence and trophic specialization. Our study contributes to understanding trophic specialization and emphasizes the importance of incorporating prey availability and their traits into diet analysis.     

Niche partitioning at emergence of two sympatric top-predator dragonflies,Anax imperator and A. parthenope (Odonata: Aeshnidae)

In natural communities, closely related species are phenotypically similar but usually spatially and/or temporally isolated. In odonates, interspecific competition occurs not only at the larval or adult stage but also during emergence. We investigated the emergence of two sympatric Anax species, focusing on the temporal pattern, vertical stratification, and body size trend over time. Anax imperator started to emerge two weeks earlier than A. parthenope but most of the emergence season overlapped. Both species showed an asynchronous emergence and the median emergence date was 10.3 days earlier in A. imperator. Sex ratio at emergence was not significantly different from 1:1. Body size of both species increased significantly over time, which contrasts many previous studies. The height of exuvia fixation was not significantly different between species but the larger species A. parthenope selected longer supports.     

Trophic niche width and overlap of two sympatric gulls in the southwestern mediterranean

The diets of two potential competitor species, Audouin's Larus audouinii and yellow-legged gulls Larus cachinnans, were examined while they bred at the Chafarinas Islands during 1993, 1994, and 1995. Data were collected during two commercial fishing regimes: (1) trawling and purse seine fisheries, and (2) diurnal trawlers only. Since the food supply for the gulls in this area was heavily reliant on the activity of purse seine fisheries, these contrasting situations allowed us to analyze short-term effects, induced by daily changes in food supply, on niche width, dietary shift, and niche overlap between the two species. Overall, both species relied mainly on fish for food, especially Clupeiforms, in the case of Audouin's gull irrespective of the fishing situation, and in the case of the yellow-legged gull, only when purse seine fishing was in operation. When purse seine boats did not operate (food shortage), yellow-legged gulls broadened their niche, consuming equal amounts of all the feeding resources, and they showed a dietary shift toward a greater consumption of prey from refuse tips. In contrast, Audouin's gulls did not change their niche width, but showed a slight dietary shift away from the consumption of epipelagic fish, compensated by an increase in reliance on benthic-mesopelagic resources. Niche overlap was clearly higher on days when both fishing fleets operated, probably because a superabundant food resource facilitates high overlap without affecting coexistence between the two species. Since our study was developed on the basis of daily variations in food supply, and competition effects are to be expected on a longer-term basis, these changes can be seen as the outcome of the coexistence of two species in stable competitive equilibrium. Received: 4 October 1996 / Accepted: 20 May 1997     

Seed production and germination in two rare and three common co‐occurring Acacia species from south‐east Australia

Abstract Seed set, size, viability and germination requirements were investigated for two rare (Acacia ausfeldii and A. willianisonii) and three common (A. pycnantha, A. genistifolia and A.paradoxa) co‐occurring congeners in box‐ironbark eucalypt forests near Bendigo, south‐east Australia to investigate correlates of rarity. Seed size was significantly smaller for the two rare species and germinants were less able to emerge from deeper sowing depths than were the larger seeded common congeners. All species had a strong heat‐stimulated germination response. While the rare A. ausfeldii showed strong germination only at the highest temperature treatment (100°C), the common and widespread A.pycnantha showed strong germination across a broad range of temperatures (60‐100°C), likely to be experienced by soil‐stored seeds during a fire. Seed viability, number of seeds per plant, and number of firm, aborted and eaten seeds per pod varied between species, but the pattern of variation was not related to rarity. Small seed size and a very specific temperature requirement for germination may help to explain rarity in A. ausfeldii, and to a lesser extent in A. willianisonii. Fires are often patchy and heating of the soil is likely to be highly spatially variable, so species with germination responses to a broad range of temperatures have an advantage over those that respond only to a narrow range. A narrower range of soil depths from which seeds can emerge will further reduce the proportion of the seed bank that might recruit following fire. Human impacts on species habitats, such as fragmentation, loss of topsoil through mining, timber harvesting, grazing and urbanization, and consequent reduction in fire intensity, are likely to have further contributed to rarity in these species. The role of pollination and other factors in relation to population size is the subject of further investigation.     

Reproductive biology of two co‐occurring mugilids,Liza argentea and Myxus elongatus,in south‐eastern Australia

The reproductive biology of Liza argentea and Myxus elongatus occurring in two estuaries (Lake Macquarie and St Georges Basin) was found to differ. Gonado‐somatic index values and macroscopic staging of gonads identified the peak spawning period of L. argentea occurred between March and November in Lake Macquarie and January and April in St Georges Basin. In contrast, peak spawning of M. elongatus was concentrated between January and March in both estuaries. Spawning of L. argentea probably occurred in the lower reaches of estuaries as well as in nearshore coastal waters, whereas evidence indicated M. elongatus spawned only in ocean waters. The mean fork length at maturity (L_F50) was greater for females than males in both species, and it also occurred at a larger mean L_F in M. elongatus (males = 230 mm and females = 255 mm) than L. argentea (males = 180 mm and females = 207 mm). Estimates of total potential fecundity were also greater for M. elongatus (425 484–1 157 029) compared to L. argentea (159 933–548 954). Both species had determinate fecundity and displayed a group synchronous pattern of oocyte development, with two distinct size classes of oocytes present in mature ovaries. Liza argentea probably release the larger class of oocytes in one spawning event, but this could not be established for M. elongatus.     

Niche overlap in allotopic and syntopic populations of sexually interacting ground‐hopper species

Abstract There is accumulating evidence that sexual interactions among species (reproductive interference) could have dramatic effects for species’ coexistence. It has been shown that the fitness of individuals can be substantially reduced as a consequence of reproductive interference. This might subsequently lead to displacement of a species (sexual exclusion). On the other hand, some evolutionary and ecological mechanisms might enable species to coexist, such as the divergence of mate recognition systems (reproductive character displacement), habitat partitioning, clumped dispersion patterns or different colonization capabilities. We have previously shown that the two ground‐hopper species Tetrix subulata and Tetrix ceperoi interact sexually in the laboratory as well as in the field. At sites where both species co‐occur niche overlap was high, suggesting that coexistence is maintained by different niche breadths rather than by habitat partitioning. To test the hypothesis that habitat partitioning does not contribute to species’ coexistence, we examined whether allotopic and syntopic populations of these two species differ in niche overlap (competitive release). Our results show that niche overlap is higher in syntopic than in allotopic populations, suggesting that the site‐specific habitat structure (heterogeneity) has a stronger influence on microhabitat utilization than the presence of heterospecifics. Hence, our data do not support the hypothesis that habitat partitioning plays a substantial role for the coexistence of these sexually interacting species.