A multi-gene phylogeny of Cephalopoda supports convergent morphological evolution in association with multiple habitat shifts in the marine environment

ABSTRACT: BACKGROUND: The marine environment is comprised of numerous divergent organisms living under similar selective pressures, often resulting in the evolution of convergent structures such as the fusiform body shape of pelagic squids, fishes, and some marine mammals. However, little is known about the frequency of, and circumstances leading to, convergent evolution in the open ocean. Here, we present a comparative study of the molluscan class Cephalopoda, a marine group known to occupy habitats from the intertidal to the deep sea. Several lineages bear features that may coincide with a benthic or pelagic existence, making this a valuable group for testing hypotheses of correlated evolution. To test for convergence and correlation, we generate the most taxonomically comprehensive multi-gene phylogeny of cephalopods to date. We then create a character matrix of habitat type and morphological characters, which we use to infer ancestral character states and test for correlation between habitat and morphology. RESULTS: Our study utilizes a taxonomically well-sampled phylogeny to show convergent evolution in all six morphological characters we analyzed. Three of these characters also correlate with habitat. The presence of an autogenic photophore is correlated with a pelagic habitat, while the cornea and accessory nidamental gland correlate with a benthic lifestyle. Here, we present the first statistical tests for correlation between convergent traits and habitat in cephalopods to better understand the evolutionary history of characters that are adaptive in benthic or pelagic environments, respectively. DISCUSSION: Our study supports the hypothesis that habitat has influenced convergent evolution in the marine environment: benthic organisms tend to exhibit similar characteristics that confer protection from invasion by other benthic taxa, while pelagic organisms possess features that facilitate crypsis and communication in an environment lacking physical refuges. Features that have originated multiple times in distantly related lineages are likely adaptive for the organisms inhabiting a particular environment: studying the frequency and evolutionary history of such convergent characters can increase understanding of the underlying forces driving ecological and evolutionary transitions in the marine environment.     

Spatial variation of reef fishes and the relative influence of biotic and abiotic habitat traits

Patterns of distribution of reef fishes were examined across three spatial scales and related to habitat traits along 25 km of the northern Portuguese coast. Response variables included the multivariate assemblage structure, the total number of taxa and individuals, and the abundance of single groups categorized according to their preference for the benthic, proximo-benthic or pelagic environment, feeding and reproductive behaviour. Habitat traits included topographic elements (small and large ‘drops’ like cracks and crevices) and the extent of dominant morpho-functional types of macroalgae (kelp, large foliose, small erect, turf-forming filamentous, and encrusting). All fish responses were characterized by the largest variance at the smallest scale (among transects tens m apart), followed by that among reefs (hundreds m to 1 km apart) and almost null variance among sites (some km apart). Small and large ‘drops’ of the substratum explained, respectively, considerable variation of assemblage structure and the total abundance of individuals, while the extent of bare rock influenced the richness of taxa and that of benthic fishes, fishes feeding on sessile invertebrates and fishes laying benthic eggs or having nesting behaviour. Combinations of abiotic and biotic structural attributes of reefs influenced proximo-benthic fishes, the predators of mobile animals and fishes releasing pelagic eggs. The here reported associations between patterns of distribution of reef fishes and habitat traits have implications for the design of future protection schemes suitable to guarantee the conservation of reef fish communities and of the processes responsible for their variation. Within the SLOSS (single-large vs. several-small) debate in the design of marine reserves, for example, effective protection to the studied reef fishes would be provided by a set of small reserves, rather than a single large which might be appropriate for fishes having wider home ranges.     

Evolutionary bottlenecks in brackish water habitats drive the colonization of fresh water by stingrays

Species richness in freshwater bony fishes depends on two main processes: the transition into and the diversification within freshwater habitats. In contrast to bony fishes, only few cartilaginous fishes, mostly stingrays (Myliobatoidei), were able to colonize fresh water. Respective transition processes have been mainly assessed from a physiological and morphological perspective, indicating that the freshwater lifestyle is strongly limited by the ability to perform osmoregulatory adaptations. However, the transition history and the effect of physiological constraints on the diversification in stingrays remain poorly understood. Herein, we estimated the geographic pathways of freshwater colonization and inferred the mode of habitat transitions. Further, we assessed habitat‐related speciation rates in a time‐calibrated phylogenetic framework to understand factors driving the transition of stingrays into and the diversification within fresh water. Using South American and Southeast Asian freshwater taxa as model organisms, we found one independent freshwater colonization event by stingrays in South America and at least three in Southeast Asia. We revealed that vicariant processes most likely caused freshwater transition during the time of major marine incursions. The habitat transition rates indicate that brackish water species switch preferably back into marine than forth into freshwater habitats. Moreover, our results showed significantly lower diversification rates in brackish water lineages, whereas freshwater and marine lineages exhibit similar rates. Thus, brackish water habitats may have functioned as evolutionary bottlenecks for the colonization of fresh water by stingrays, probably because of the higher variability of environmental conditions in brackish water.     

The evolution of underwater flight: The redistribution of pectoral fin rays,in manta rays and their relatives (Myliobatidae)

Batoids are a diverse clade of flat cartilaginous fishes that occur primarily in benthic marine habitats. The skates and rays typically use their flexible pectoral fins for feeding and propulsion via undulatory swimming. However, two groups of rays have adopted a pelagic or bentho‐pelagic lifestyle and utilize oscillatory swimming—the Myliobatidae and Gymnuridae. The myliobatids have evolved cephalic lobes, anteriorly extended appendages that are optimized for feeding, while their pectoral fins exhibit several modifications that likely arose in association with functional optimization of pelagic cruising via oscillatory flight. Here, we examine variation in fin ray distribution and ontogenetic timing of fin ray development in batoid pectoral fins in an evolutionary context using the following methods: radiography, computed tomography, dissections, and cleared and stained specimens. We propose an index for characterizing variation in the distribution of pectoral fin rays. While undulatory swimmers exhibit symmetry or slight anterior bias, we found a posterior shift in the distribution of fin rays that arose in two distinct lineages in association with oscillatory swimming. Undulatory and oscillatory swimmers occupy nonoverlapping morphospace with respect to fin ray distribution illustrating significant remodeling of pectoral fins in oscillatory swimmers. Further, we describe a derived skeletal feature in anterior pectoral fins of the Myliobatidae that is likely associated with optimization of oscillatory swimming. By examining the distribution of fin rays with clearly defined articulation points, we were able to infer evolutionary trends and body plan remodeling associated with invasion of the pelagic environment. Finally, we found that the number and distribution of fin rays is set early in development in the little skate, round stingray, and cownose ray, suggesting that fin ray counts from specimens after birth or hatching are representative of adults and therefore comparable among species.     

Patterns of lineage diversification in the genus Naso (Acanthuridae)

The evolutionary history of the reef fish genus Naso (F. Acanthuridae) was examined using a complete species-level molecular phylogeny of all recognized (19) species based on three loci (one nuclear ETS2 and two mitochondrial 16S, cyt b). This study demonstrates that distinct foraging modes and specialized body shapes arose independently at different times in the evolutionary history of the genus. Members of the subgenus Axinurus, characterized by a scombriform morphology, caudal fin structure and pelagic foraging mode, were consistently placed basal to the remaining Naso species, suggesting that pelagic foraging is plesiomorphic and benthic foraging derived in this genus. We used a genus-level phylogeny (nuclear marker, ETS2), which included several taxa from all other acanthurid genera, to obtain a range of age estimates for the most recent common ancestor of the genus Naso. These age estimates (range of 52-43.3 MY) were then used to estimate divergence times (by nonparametric rate smoothing method) of the node giving rise to extant Naso species using the combined sequence data (from all loci). The reconstruction of the pattern of divergence of extant species indicates two sequences of events. The basal species characterized by pelagic foraging modes arose during the Eocene and Oligocene. Most of the remaining Naso species, including those characterized by benthic foraging, arose over a period of 20 MY during the Miocene. Diversification during this period was associated with major plate tectonic and glaciation events, resulting in changes in sea level, ocean temperature and productivity regimes. Regardless of the foraging mode exhibited, all species of Naso have a caudal propulsive unit similar to that observed in pelagic scombriform fishes, a legacy of the basal position of the subgenus Axinurus in the phylogeny of the genus.     

Molecular pedomorphism underlies craniofacial skeletal evolution in Antarctic notothenioid fishes

Background  

Pedomorphism is the retention of ancestrally juvenile traits by adults in a descendant taxon. Despite its importance for evolutionary change, there are few examples of a molecular basis for this phenomenon. Notothenioids represent one of the best described species flocks among marine fishes, but their diversity is currently threatened by the rapidly changing Antarctic climate. Notothenioid evolutionary history is characterized by parallel radiations from a benthic ancestor to pelagic predators, which was accompanied by the appearance of several pedomorphic traits, including the reduction of skeletal mineralization that resulted in increased buoyancy.     

Morphological divergence between three Arctic charr morphs – the significance of the deep‐water environment

Morphological divergence was evident among three sympatric morphs of Arctic charr (Salvelinus alpinus (L.)) that are ecologically diverged along the shallow‐, deep‐water resource axis in a subarctic postglacial lake (Norway). The two deep‐water (profundal) spawning morphs, a benthivore (PB‐morph) and a piscivore (PP‐morph), have evolved under identical abiotic conditions with constant low light and temperature levels in their deep‐water habitat, and were morphologically most similar. However, they differed in important head traits (e.g., eye and mouth size) related to their different diet specializations. The small‐sized PB‐morph had a paedomorphic appearance with a blunt head shape, large eyes, and a deep body shape adapted to their profundal lifestyle feeding on submerged benthos from soft, deep‐water sediments. The PP‐morph had a robust head, large mouth with numerous teeth, and an elongated body shape strongly related to their piscivorous behavior. The littoral spawning omnivore morph (LO‐morph) predominantly utilizes the shallow benthic–pelagic habitat and food resources. Compared to the deep‐water morphs, the LO‐morph had smaller head relative to body size. The LO‐morph exhibited traits typical for both shallow‐water benthic feeding (e.g., large body depths and small eyes) and planktivorous feeding in the pelagic habitat (e.g., streamlined body shape and small mouth). The development of morphological differences within the same deep‐water habitat for the PB‐ and PP‐morphs highlights the potential of biotic factors and ecological interactions to promote further divergence in the evolution of polymorphism in a tentative incipient speciation process. The diversity of deep‐water charr in this study represents a novelty in the Arctic charr polymorphism as a truly deep‐water piscivore morph has to our knowledge not been described elsewhere.     

Prepared for the future: A strong signal of evolution toward the adult benthic niche during the pelagic stage in Labrid fishes

The morphology of organisms reflects a balance between their evolutionary history, functional demands, and biomechanical constraints imposed by the immediate environment. In many fish species, a marked shift in the selection regime is evident when pelagic larvae, which swim and feed in the open ocean, settle in their adult benthic habitat. This shift is particularly dramatic in coral‐reef fishes, where the adult habitat is immensely complex. However, whether the adult trophic ecotype affects the morphology of early‐life stages is unclear. We measured a suite of 26 functional‐morphological traits in the head and body of larvae from an ontogenetic series of 16 labrid species. Using phylogenetic comparative methods, we reconstructed the location of adaptive peaks of larvae whose adults are associated with different trophic ecotypes. We found that the morphospace occupation in these larvae is largely driven by divergent adaptations to the adult benthic habitats. The disparity between adaptive peaks is achieved early and does not monotonically increase with size. Our findings thus refute the notion that larvae rapidly acquire the trophic‐specific traits during a metamorphic period immediately prior to settlement. This early specialization might be due to the highly complex musculoskeletal system of the head that cannot be rapidly modified.     

Why do juvenile fourhorn sculpin, Triglopsis quadricornis, appear in the pelagic habitat at night?

Synopsis Challenging the generally accepted view of the fourhorn sculpin Triglopsis quadricornis as a cold-stenothermal and true benthic species, novel findings of juveniles in the pelagic habitat of Lake Vättern, Sweden, have been examined and critically evaluated. Young-of-the-year and small-sized, sexually mature individuals, ranging 27–45 and 82–110 mm in total length, were recorded in nocturnal trawl catches from pelagic midwater in August. The sculpin had fed extensively on both benthic and semi-pelagic crustaceans, including large cyclopoid copepods, Mysis relicta, Pallasea quadrispinosa, Monoporeia affinis, Gammaracanthus lacustris, and Saduria entomon. Based on an extensive literature review a series of ecological and evolutionary reasons for a pelagic behavior of this profundal coldwater fish is suggested. Nocturnal, semi-pelagic feeding, and consecutive, thermotactic, vertical migrations up in midwater as shown by Wurtsbaugh & Neverman (1988) in order to maximize energy intake after feeding, are possible adaptations of these juvenile sculpin for minimizing size-dependent mortality due to intra- and interspecific predation and competition from other deepwater fishes.     

Key stages in the evolution of the Antarctic marine fauna

We are beginning to appreciate that the origin of the modern Antarctic marine fauna is related to a series of key events throughout the Cenozoic era. In the first of these, the mass extinction at the Cretaceous–Palaeogene boundary (66 Ma) reset the evolutionary stage and led to a major radiation of modern taxa in the benthic realm. Although this took place in a greenhouse world, there is evidence to suggest that the radiation was tempered by the seasonality of primary productivity, and this may be a time‐invariant feature of the polar regions. Although there could well have been a single, abrupt extinction event at c. 34 Ma, there is also evidence to suggest a phased extinction of various taxa over a period of millions of years. Important new molecular phylogenetic data are indicating that a wide variety of both benthic and pelagic taxa radiated shortly after a second major phase of cooling at c. 14 Ma. Such a phenomenon is linked to a series of major palaeoceanographic changes, which in turn led to a proliferation of diatom‐based ecosystems. Although the modern benthic marine fauna can be traced back some 45–50 Myr, a substantial component of the modern pelagic one may be less than 14 Myr old. The latter is also characterized by assemblages of high abundance but comparatively low species richness and evenness. A distinctive signature of low diversity but high dominance within Antarctic marine assemblages was maintained by the interplay between temperature and primary productivity throughout the Cenozoic.     

Habitat coupling in lake ecosystems

Lakes are complex ecosystems composed of distinct habitats coupled by biological, physical and chemical processes. While the ecological and evolutionary characteristics of aquatic organisms reflect habitat coupling in lakes, aquatic ecology has largely studied pelagic, benthic and riparian habitats in isolation from each other. Here, we summarize several ecological and evolutionary patterns that highlight the importance of habitat coupling and discuss their implications for understanding ecosystem processes in lakes. We pay special attention to fishes because they play particularly important roles as habitat couplers as a result of their high mobility and flexible foraging tactics that lead to inter-habitat omnivory. Habitat coupling has important consequences for nutrient cycling, predator-prey interactions, and food web structure and stability. For example, nutrient excretion by benthivorous consumers can account for a substantial fraction of inputs to pelagic nutrient cycles. Benthic resources also subsidize carnivore populations that have important predatory effects on plankton communities. These benthic subsidies stabilize population dynamics of pelagic carnivores and intensify the strength of their interactions with planktonic food webs. Furthermore, anthropogenic disturbances such as eutrophication, habitat modification, and exotic species introductions may severely alter habitat connections and, therefore, the fundamental flows of nutrients and energy in lake ecosystems.     

Feeding mechanics in Triassic stem-group sauropterygians: the anatomy of a successful invasion of Mesozoic seas

The jaw adductor musculature in Triassic stem-group sauropterygians is reconstructed on the basis of a paradigmatic model of muscle architecture (functional equivalence of sarcomeres) and using invariant traits of the anatomy of the trigeminal jaw adductor muscles in extant reptiles. The reconstructed jaw adductor musculature predicts trophic specializations in stem-group sauropterygians. Suction feeding is a component in prey capture for some benthic feeding, as well as for some pelagic feeding taxa. The differentiation of 'pincer' jaws is correlated with the potential for rapid, snapping bites. There is some evidence for habitat partitioning among Triassic stem-group sauropterygians with respect to trophic specialization. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 135 , 33–63.     

Adaptations of Cave Fishes with some Comparisons to Deep-sea Fishes

I provide my retrospective and prospective views on adaptations of cave fishes. I emphasize the history of my insights into cave adaptation from 45 years of research using surface, cave-spring, and cave species of amblyopsid fishes. My approach has been to use natural experiments and to always consider multiple hypotheses. To clarify evolutionary adaptations, I show the importance of a broad comparative approach which includes studies of morphology, metabolic physiology, foraging behavior, life history, and ecology. And I show that the most important agents of selection, of darkness and attendant low food supply, are best understood in the context of rigor, variability, and predictability. I also present my insights from what I consider the most insightful contributions on deep-sea fishes. The contributions are those of Marshall in studies of morphology in relation to energy economy of pelagic and benthic species, Childress in studies of physiological and biochemical adaptations with depth for pelagic species, and Koslow in studies on population biology and life history of bathybenthic and benthic sea-mount species. Compared to caves, I suggest that the extremes of metabolic and life history adaptations of deep-sea fish are explained by a longer evolutionary history and a much greater habitat range, food supply, and predation risk. Finally, I take a retrospective view of what we have learned about cave fishes. I discuss possible evolutionary mechanisms that can explain the trends with increasing cave adaptation in amblyopsid fishes, especially progenesis and the pleiotropic effects of the stress resistance syndrome. Finally, based on insights from deep-sea fishes, and emerging new techniques, I suggest what cave fish biologists should do in the future.     

Predation-imposed selection on threespine stickleback (Gasterosteus aculeatus) morphology: a test of the refuge use hypothesis

The transition from marine to freshwater life in the threespine stickleback (Gasterosteus aculeatus) is accompanied by complex morphological changes-including reduction in bony armor and change in body shape-but experimental evidence for the selective agents behind these evolutionary transitions is sparse. We investigated whether selection by predatory fish affects threespine stickleback morphology differentially when refuge is absent (pelagic lifestyle-ancestral condition) or present (benthic lifestyle-derived condition). Our results show that selection favors low numbers of lateral plates in habitats with refuge, whereas fully plated individuals have a selective advantage in habitats without refuge. We also found that a decrease in the length of the caudal peduncle increased survival probability, irrespective of habitat. The effect of spine lengths on survival was evident only in a multivariate analysis of selection, implying that it is essential to account for phenotypic and genetic correlations between traits before drawing conclusions about the effects of selection on single traits. Apart from uncovering targets and patterns of predator-induced selection on threespine stickleback morphology, our results provide direct evidence to support the hypothesis that differences in antipredator strategies in pelagic versus benthic sticklebacks could play a role in the repeated, independent cases of plate number reduction following freshwater colonization in this species.     

Edgeworth's legacy of cranial muscle development with an analysis of muscles in the ventral gill arch region of batoid fishes (Chondrichthyes: Batoidea).

A series of studies by Edgeworth demonstrated that cranial muscles of gnathostome fishes are embryologically of somitic origin, originating from the mandibular, hyoid, branchial, epibranchial, and hypobranchial muscle plates. Recent experimental studies using quail-chick chimeras support Edgeworth's view on the developmental origin of cranial muscles. One of his findings, the existence of the premyogenic condensation constrictor dorsalis in teleost fishes, has also been confirmed by molecular developmental studies. Therefore, developmental mechanisms for patterning of cranial muscles, as described and implicated by Edgeworth, may serve as structural entities or regulatory phenomena responsible for developmental and evolutionary changes. With Edgeworth's and other studies as background, muscles in the ventral gill arch region of batoid fishes are analyzed and compared with those of other gnathostome fishes. The spiracularis is regarded as homologous at least within batoid fishes, but its status within elasmobranchs remains unclear; developmental modifications of the spiracularis proper are evident in some batoid fishes and in several shark groups. The peculiar ventral extension of the spiracularis in electric rays and some stingrays may represent convergence, probably facilitating ventilation and/or feeding in both groups. The evolutionary origin of the "internus" and "externus" remains uncertain, despite the fact that a variety of forms of the constrictor superficiales ventrales in batoid fishes indicates an actual medio-ventral extension of the "externus." The intermandibularis is probably present only in electric rays. The "X" muscle occurs only in electric rays and is considered to be Edgeworth's intermandibularis profundus. Its association with the adductor mandibular complex in narkinidid and narcinidid electric rays may relate to its functional role in lower jaw movement. Contrary to common belief, in most batoid fishes as well as some sharks, muscles that originate from the branchial muscle plate and extend medially in the ventral gill arches do exist: the medial extension of the interbranchiales in most batoid fishes and some sharks and the "Y" muscle in the pelagic stingrays Myliobatos and Rhinoptera. The latter is another example of the medial extension of the "internus." Whether the interbranchiales and "Y" muscle are homologous within elasmobranchs and whether homologous with the obliques ventrales and/or transversi ventrales of osteichthyan fishes await further research. Four hypobranchial muscles are recognized in batoid fishes: the coracomandibularis, coracohyoideus, coracoarcualis, and coracohyomandibularis. The coracohyoideus is discrete from the coracoarcualis; its complete structural separation from the latter occurs in several groups of batoid fishes.(ABSTRACT TRUNCATED AT 400 WORDS)     

Origins of diverse feeding ecologies within Conus, a genus of venomous marine gastropods

Specialized predators on polychaetes, fishes, hemichordates or other molluscs, members of the predominantly tropical gastropod genus Conus diversified rapidly during the Miocene to constitute the most species-rich modern marine genus. We used DNA sequence data from mitochondrial and nuclear loci of 76 Conus species to generate species-level phylogenetic hypotheses for this genus and then mapped known diets onto the phylogenies to elucidate the origins and evolutionary histories of different feeding specializations. The results indicate that dramatically new feeding modes arose only a few times, that the most derived feeding modes likely arose in the Miocene, and that much of the known diversity of Conus that was generated during Miocene radiations has survived to the present.     

Ecomorphology of Anolis lizards of the Choco' region in Colombia and comparisons with Greater Antillean ecomorphs

We analysed the ecomorphological relationships in four species of Anolis lizard that occur in the Choco' region in Colombia. The region is one of the most diverse of the Neotropical lowlands. The species were assigned to traditionally recognized Greater Antillean ecomorph categories based on habitat use data. Principal component analyses were carried out to examine correlations between the morphological traits, body size, and habitat use. We found that species are separated in morphological space principally by body size and lamella number. Upon removal of the effect of body size, correlations between morphology and habitat use became apparent. However, when compared with Greater Antillean ecomorphs, we found little evidence of morphological convergence in species occupying similar habitats. The species of the Choco' region are, however, clearly separated in the multidimensional morphological space from the Antillean taxa, and appear to form a separate cluster differentiating principally in body size and the number of lamellae. Mainland species clearly constitute an ecomorphological radiation but apparently this is independent of that of the West Indian fauna. More studies are needed to understand the causes for the independence of evolutionary trajectories on the mainland and the Greater Antilles, and to obtain a better understanding of the ecological and evolutionary processes underlying the radiation of these faunas.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 29–39.     

Trophic ecology of three stingrays (Myliobatoidei: Dasyatidae) off the Brazilian north-eastern coast: Habitat use and resource partitioning

Understanding the ecological role of species with overlapping distributions is central to inform ecosystem management. Here we describe the diet, trophic level and habitat use of three sympatric stingrays, Hypanus guttatus, H. marianae and H. berthalutzae, through combined stomach content and stable isotope (δ¹³C and δ¹⁵N) analyses. Our integrated approach revealed that H. guttatus is a mesopredator that feeds on a diverse diet of benthic and epibenthic marine and estuarine organisms, principally bivalve molluscs, Alpheus shrimp and teleost fishes. Isotopic data supported movement of this species between marine and estuarine environments. H. berthalutzae is also a marine generalist feeder, but feeds primarily on teleost fishes and cephalopods, and consequently occupies a higher trophic level. In contrast, H. marianae is a mesopredator specialized on shrimps and polychaetas occurring only in the marine environment and occupying a low niche breadth. While niche overlap occurred, the three stingrays utilized the same prey resources at different rates and occupied distinct trophic niches, potentially limiting competition for resources and promoting coexistence. These combined data demonstrate that these three mesopredators perform different ecological roles in the ecosystems they occupy, limiting functional redundancy.     

Individual variations in habitat use and morphology in brook charr

The specific objectives of this study were to determine if there is individual specialization in habitat use by lacustrine brook charr Salvelinus fontinalis and if so, if specialization is related to fish morphology. Localizations of 28 brook charr equipped with thermosensitive radiotransmitters were recorded during three summers (1991, 1992, and 1993) in two lakes of the Mastigouche Reserve (Québec, Canada). Fifty per cent of the fish were found mainly in the benthic zone (hereafter benthic individuals), 18% in the pelagic zone (pelagic individuals), and 32% travelled regularly between the two zones (generalist individuals). The observed interindividual differences in habitat preference were related to differences in body morphology and coloration: (i) the pectoral fins of benthic and generalist individuals were significantly longer than those of pelagic ones; and (ii) the coloration of the lower flank of benthic and generalist individuals was silver-grey while that of pelagic individuals was red. The results of this study suggest that brook charr inhabiting oligotrophic lakes of the Canadian Shield exhibit trophic polymorphisms, where some individuals are specialists better adapted to feeding in the littoral zone whereas others are specialists better adapted to feeding in the pelagic zone. The potential for reproductive isolation between the two morphs is discussed.     

Trophic Structure of One Deep-sea Benthic Fish Community in the Eastern Canadian Arctic: Application of Food, Parasites and Multivariate Analysis

Synopsis As new arctic marine fisheries develop there is need for a comprehensive ecosystem approach to long-term management. This approach recognizes the importance of community interactions such as food web structure and trophic patterns. We determined whether hierarchical clustering (guild formation) is an effective method of trophic evaluation in deep-sea Artic fish communities using stomach content and parasite data with size class, and evaluated the application of endohelminth communities (parasite species transmitted in the food) as indicators of trophic status. Cluster analysis using food group abundance with size class of fish revealed the presence of 11 guilds within the community, however the same analysis using parasite data showed little correlation between food and parasites. Redundancy analysis (RDA) within the 11 guilds also revealed no significant correlations between food group and parasite abundance suggesting that this type of ordination is not suited for environments containing mainly generalist feeders. RDA of individual taxa without a priori guild designation found that taxa in benthic deep-sea communities are defined by their ability to exploit prey species in more than one habitat zone. Benthic fish species were significantly correlated with benthic food groups and parasites that utilize benthic intermediate hosts whereas benthopelagic–pelagic species fed on a higher diversity of prey species and were infected by a larger number of non-host specific parasites. Eigenanalysis and Monte Carlo results showed that parasites and food groups are highly correlated, indicating that parasite community analysis is an effective tool for predicting feeding strategies in Arctic marine environments. It also suggests that in most cases endoparasite infections alone could be used for trophic evaluation in the absence of stomach content data.