Alan J. Kohn

Effect of habitat complexity on population density and species richness in tropical intertidal predatory gastropod assemblages

SummaryStructural complexity of the habitat significantly increases population density and number of species in assemblages of predatory gastropod molluscs (families Conidae, Muricidae, Mitridae and Vasidae) on intertidal, generally smooth, horizontal limestone platforms fringing tropical Pacific islands. The important topographic features are physical (depressions partly filled with coral rubble) and biotic (thick algal turf binding sand). Higher population density and species richness in areas with than without such natural refuges, and following experimental addition of artificial refuges on portions of habitat lacking them support this hypothesis. Two species of Drupa differ from the other species present in not utilizing refuges during times of physical stress; this is attributed to their depressed shell and broad, tenacious foot. Highest gastropod densities occur in steep-sided depressions and those containing much coral rubble and sand, suggesting that these are important qualities of refuges. We believe this is the first demonstration of how specific environmental factors affect population density and species diversity of benthic invertebrates in a coral reef-associated habitat.

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.  2001 The Linnean Society of London

Ecology of Conus on eastern Indian Ocean fringing reefs: Diversity of species and resource utilization

The most diverse assemblages of the genus Conus known occur on fringing coral reefs in Thailand and Indonesia. As many as 27 congeneric species of these gastropods inhabit a single reef; in all, we examined 1,350 individuals of 48 species. Several attributes of the populations we observed conform to expectations of a model of ecological characteristics of bench and reef Conus proposed by Kohn (1971a). Number of species (S) averaged 15, and species diversity (H″) averaged 2.3 in the most heterogeneous habitat type — topographically complex, subtidal reef platforms (Type III habitat). Both species richness and evenness of distribution of individuals among species contribute strongly to H″. Fewer congeners and greater numerical dominance by single species characterize more homogeneous habitats. On subtidal reef platforms with large areas of sand substrate and less coral limestone (Type I–III habitat), mean values were S=10, and H″=1.6. In the one intensively studied, truncated reef-limestone platform (Type II–III intermediate habitat), S=13 and H″=1.4. Summed population density of all Conus species in Type III and I–III habitats is similar (0.02 to 0.05 individuals /m2) and comparable to estimates from similar habitats elsewhere in the Indo-West Pacific region. Mean density (0.7/m2) and other population attributes in Type II–III habitat more closely resemble those of Type II than Type III habitats in general. We combined analysis of species diversity and other attributes of assemblages in habitats of different environmental complexity with analysis of microhabitat and food-resource utilization, in order to demonstrate the extent to which specialization on different resources occurs in assemblages differing in diversity and habitat type. In the habitats studied, co-occurring species of Conus specialized to a greater extent on different prey species than on different microhabitat patches, but degree of microhabitat specialization was greater than in similarly complex habitats with assemblages of lower diversity elsewhere in the Indo-West Pacific region. While most Conus species preyed primarily on a different species or higher taxon of polychaetes, diets are not more specialized or dissimilar than in similar habitats elsewhere. Degree of specialization on different prey is not correlated with Conus species diversity in the different types of habitats studied. The data lead to the conclusion that differential predation is as important — and differential microhabitat utilization is more important — in permitting coexistence of potentially competing congeners, compared with conditions in habitats of comparable heterogeneity that support fewer congeners farther from the center of the Indo-West Pacific region. Pairwise comparisons of congeners indicate that many species pairs have low or no overlap in both microhabitat and food utilization. Members of species pairs with high overlap in microhabitat utilization typically eat different prey organisms, and those with similar diets typically occupy different habitats or microhabitats. This applies to molluscivorous as well as vermivorous species. Information on the diets of 11 species is reported here for the first time. Of 48 Indo-West Pacific Conus species whose food is now known, 35 prey on polychaetes, 2 on enteropneusts, 6 on gastropods, and 5 on fishes. Vermivorous Conus prey on relatively few of the polychaete species present in the environments. Species eaten represent only 12% of a total estimated polychaete population density of 27,000 individuals /m2. Certain very abundant polychaetes may be protected from predation by Conus by their small size, others by their long tubes. Two new aspects of size-selective predation by Conus are reported: (1) Although comparisons of predation rate with prey standing-crop suggest that food is plentiful, selective predation on the largest prey individuals present suggests that only small proportions of prey-species populations may have large enough body size to repay foraging effort by the Conus present; (2) composition of the diet changes qualitatively with increase in body size in several vermivorous Conus species; shifting by larger individuals to larger prey species could be documented in C. ebraeus.

Molecular phylogeny and evolution of the cone snails (Gastropoda, Conoidea)

We present a large-scale molecular phylogeny that includes 320 of the 761 recognized valid species of the cone snails (Conus), one of the most diverse groups of marine molluscs, based on three mitochondrial genes (COI, 16S rDNA and 12S rDNA). This is the first phylogeny of the taxon to employ concatenated sequences of several genes, and it includes more than twice as many species as the last published molecular phylogeny of the entire group nearly a decade ago. Most of the numerous molecular phylogenies published during the last 15years are limited to rather small fractions of its species diversity. Bayesian and maximum likelihood analyses are mostly congruent and confirm the presence of three previously reported highly divergent lineages among cone snails, and one identified here using molecular data. About 85% of the species cluster in the single Large Major Clade; the others are divided between the Small Major Clade (∼12%), the Conus californicus lineage (one species), and a newly defined clade (∼3%). We also define several subclades within the Large and Small major clades, but most of their relationships remain poorly supported. To illustrate the usefulness of molecular phylogenies in addressing specific evolutionary questions, we analyse the evolution of the diet, the biogeography and the toxins of cone snails. All cone snails whose feeding biology is known inject venom into large prey animals and swallow them whole. Predation on polychaete worms is inferred as the ancestral state, and diet shifts to molluscs and fishes occurred rarely. The ancestor of cone snails probably originated from the Indo-Pacific; rather few colonisations of other biogeographic provinces have probably occurred. A new classification of the Conidae, based on the molecular phylogeny, is published in an accompanying paper.

Hidden diversity in a hyperdiverse gastropod genus: Discovery of previously unidentified members of a Conus species complex

Molecular sequence data are a powerful tool for delimiting species, particularly in cases where morphological differences are obscure. Distinguishing species in the Conus sponsalis complex of tropical marine gastropods has long been difficult, because descriptions and identification has relied exclusively on shell characters, primarily color patterns, and these often appear to intergrade among putative species. Here we use molecular sequence data from two mitochondrial gene regions (16S rRNA and cytochrome oxidase subunit I) and one nuclear locus (a four-loop conotoxin gene) to characterize the genetic discontinuity of the nominal species of this group currently accepted as valid: the Indo-West Pacific C. sponsalis, C. nanus, C. ceylanensis, C. musicus and C. parvatus, and the eastern Pacific C. nux. In these analyses C. nanus and C. sponsalis resolve quite well and appear to represent distinct evolutionary units that are mostly congruent with morphology-based distinctions. We also identified several cryptic entities whose genetic uniqueness suggests species-level distinctions. Two of these fit the original description of C. sponsalis; three forms appear to represent C. nanus but differ in adult shell size or possess a unique shell color pattern.

PREDATION ON EARLY PLEISTOCENE NATICID GASTROPODS IN FIJI

Abstract We determined causes of predation on an early Pleistocene assemblage of ten species of Naticidae in Fiji. Naticids preyed more frequently on the four Natica species (27% of shells drilled) than on Polinices, Eunaticina and Sinum (3% of shells drilled). Tertiary and Cretaceous naticids studied previously are generally less frequently drilled. The absence of incomplete boreholes indicates that drilling predation is typically successful. Predator size (based on borehole diameter) increased directly with prey size in all Natica species. Incidence of drilling predation decreased slightly with increasing prey shell size, but Natica species do not appear able to outgrow drilling predation. Unsuccessful crustacean predation, indicated by scar lines and shell repair, occurred frequently. In the eight commonest naticid species, 17–48% of the shells had healed fractures. The results identify two trophic links in the fossil community, naticids as generally successful predators and crustaceans as often unsuccessful predators, both on naticid prey.

Radula Tooth Structure of the Gastropod Conus imperialis Elucidated by Scanning Electron Microscopy

Scanning electron microscopy of the hollow, harpoon-like radula tooth of the toxoglossan gastropod Conus has elucidated the structure and relationships of its component parts: apex, cutting edge, barbs, serration, adapical and basal openings of the lumen, external and internal folds of the shaft, and base. The functional roles of these components in prey capture are proposed.

Escape responses of three herbivorous gastropods to the predatory gastropod Conus textile

Summary The herbivorous gastropods Strombus canarium, Lambis lambis, and Trochus pyramis evince escape responses, probably mediated by distance chemoreception, in the presence of the predatory gastropod Conus textile. In Strombus and Lambis, the response has two components: increase in the absolute rate of locomotion, and direction of locomotion away from the predator. When placed in the presence of C. textile, T. pyramis began moving, generally in the direction it faced. In control observations, a specimen of S. canarium moved at a mean rate of 2·7 mm/sec, by the leaping mode of locomotion characteristic of this and related genera. In the presence of C. textile, the rate increased to 6·8 mm/sec, due primarily to shortening of the lag or recovery period between leaps. In the presence of C. textile, a specimen of L. lambis moved at a rate of 11·9 mm/sec in the same manner as Strombus. In the presence of C. textile, a specimen of T. pyramis progressed at a rate of 1·9 mm/sec, by pedal locomotory waves. Comparison with absolute rates of locomotion by pedal locomotory waves in other marine gastropod molluscs leads to the conclusion that the leaping of Strombus and Lambis does not enable these gastropods to maintain a more rapid long-term rate of movement, but it confers a striking increase in agility, and a single leap provides an almost instantaneous increase in distance from a potential predator.

Cryptic Species Differentiated in Conus ebraeus, a Widespread Tropical Marine Gastropod

Anomalous mitochondrial and nuclear gene sequences in individuals of the widely distributed tropical marine gastropod Conus ebraeus that were not distinguishable by shell shape and color pattern characters suggested the presence of a second, cryptic species. We tested this hypothesis by genetic, morphological, and ecological comparisons of additional individuals from the site in Okinawa where the two forms co-occurred. Radular tooth size and shape, prey type in nature, and microhabitats utilized differed markedly between the two forms. Adults with typical C. ebraeus DNA and radular teeth preyed primarily on errant polychaetes (Eunicidae); those with anomalous DNA and teeth ate mainly sedentary capitellids. Juveniles (shell length <13 mm) had more similar teeth and ate primarily syllids. Radular teeth of the anomalous form agreed with those of Conus judaeus, distinguished from C. ebraeus by Rudolph Bergh in 1895 solely on tooth characters of one specimen from the Philippines. Samples from other widely scattered Pacific localities revealed only typical C. ebraeus gene sequences. Both forms occurred in Seychelles (western Indian Ocean), where their radular teeth and diets were consistent with the data from Okinawa, but DNA of available material was degraded. Although C. judaeus was long dismissed as an aberrant specimen and junior synonym of C. ebraeus, our results support its validity as a distinct species. These results highlight the importance of molecular and radular tooth characters relative to those of the shell. Moreover, cryptic species could well be important components of species richness in Conus specifically and marine molluscan biodiversity more generally.

Imposex in Australian Conus

Previously unknown in Conus (Mollusca: Gastropoda) and poorly documented in the Australian region, imposex is demonstrated in 80% of females of six Conus species from Rottnest Island, Western Australia. In no case were the effects sufficiently severe to cause sterility, and reproduction appeared normal. Specimens from the same location preserved in 1975 were unaffected. With more than 500 species, Conus is the largest genus of marine molluscs. Because many species are conspicuous, widely distributed, and occur in populations whose past densities are known, they could serve as monitors of artificially induced pseudohermaphroditism and an early warning of future population responses to pollutants in marine environments at low latitudes.