A molecular phylogeny of the Patellogastropoda (Mollusca: Gastropoda)

Phylogenetic analyses of partial 18S rDNA sequences from species representing all living families of the order Patellogastropoda, most other major gastropod groups (Cocculiniformia, Neritopsina, Vetigastropoda, Caenogastropoda, Heterobranchia, but not Neomphalina), and two additional classes of the phylum Mollusca (Cephalopoda, Polyplacophora) confirm that Patellogastropoda comprises a robust clade with high statistical support. The sequences are characterized by the presence of several insertions and deletions that are unique to, and ubiquitous among, patellogastropods. However, this portion of the 18S gene is insufficiently informative to provide robust support for the monophyly of Gastropoda, or to address the division of the Gastropoda into the subclasses Eogastropoda (= Patellogastropoda + hypothetical coiled ancestors) and Orthogastropoda. These sequence data invariably group Patellogastropoda in a weakly supported clade with cocculiniform limpets, despite greater sequence divergences between Patellogastropoda and “Cocculiniformia” than between the Patellogastropoda and Orthogastropoda. Partial 18S sequences support the inclusion of the family Neolepetopsidae within the superfamily Acmaeoidea, and refute its previously hypothesized position as sister group to the remaining living Patellogastropoda. This region of the 18S rDNA gene diverges at widely differing rates, spanning an order of magnitude among patellogastropod lineages, and therefore does not provide meaningful resolution of the relationships among higher taxa of patellogastropods. Data from one or more genes that evolve more uniformly and more rapidly than the 18S rDNA gene (possibly one or more of the mitochondrial genes) seem more likely to be informative about relationships within Patellogastropoda. Received: 5 February 1999 / Accepted: 16 May 2000     

Chemical defense and evolution in the Sacoglossa (Mollusca: Gastropoda: Opisthobranchia)

Summary. Recent advances in both the systematics and the natural products chemistry of the order Sacoglossa (=Ascoglossa) in the gastropod subclass Opisthobranchia suggest a revised and improved historical account of the evolution of the group. Although the algal genus Caulerpa makes a suitable model for the ancestral food of the order, other siphonaceous algae are consistent with both morphological and chemical data. At an early evolutionary stage terpenoids are sequestered from the food, and used defensively, often with modification. With an evolutionary switch to different kinds of algal food, there is often a shift to other, related defensive chemicals. A switch to new food source sometimes leads to the abandonment of chemical defense, but in other cases there is de-novo synthesis of defensive metabolites. The synthesis of polypropionates, which are used defensively, occurs in some other gastropods, but otherwise is known only in fungi. The systematic distribution of the defensive polypropionates suggests that their defensive use has evolved several times among gastropods. Failure to detect them may mean that synthetic capacity has evolved more than once, or it may mean that they exist at low levels, perhaps having a non-defensive function. Received 9 February 1998; accepted 20 March 1998.     

The nature of heat coma in Littorina littorea (Mollusca: Gastropoda)

 The nature of heat coma was examined in the edible periwinkle Littorina littorea (L.). Duration of acclimation did not influence heat-coma temperature at 12 °C, although other acclimation temperatures were important in influencing thermal tolerance, with positive shifts in coma temperature observed in response to elevated temperatures. Previous thermal history also influenced heat-coma temperatures. Individuals subjected to repeat heat-coma events on a daily basis showed significant declines (P < 0.05) in coma-temperature; in contrast individuals exposed to repeat heat-coma events on a weekly basis showed no decline in thermal tolerance. Size-effects occurred at selected sites, where decreased heat-coma temperatures were recorded in large individuals. Received: 16 August 1999 / Accepted: 13 June 2000     

Intertidal distribution and long-term movements of Littorina irrorata (Mollusca: Gastropoda)

The distribution of Littorina irrorata Say on a low-energy barrier beach on the northern Gulf of Mexico is described, and correlated with the presence of Spartina alterniflora and other plants in the upper intertidal zone. The movements of 66 individually tagged snails were followed in the S. alterniflora zone for an average of 226 days, during which time an average of 10.6 positions were recorded per snail. The snails travelled an average total path distance of at least 995 cm, but due to contorted paths, ended up an average resultant distance of only 399 cm away from their original positions. They moved an estimated resultant distance of approximately 20 to 25 cm per activity period. Despite a slight offshore movement during the fall and winter, the snails moved more parallel to the shoreline than perpendicular to it.     

Regional extinction of a conspicuous dorid nudibranch (Mollusca: Gastropoda) in California

Felimare californiensis (=Hypselodoris californiensis) was once common throughout the Southern California Bight (SCB) and California Channel Islands. This well-known shallow-water nudibranch, which specializes on dysideid sponges, has persisted for decades in Mexico, but in California disappeared from its entire range by 1984. Since reappearing in 2003, it has been found only at Santa Catalina Island, plus sightings of single individuals in 2011 at Santa Cruz Island and San Diego. The decline of F. californiensis in California was documented using published historical records, museum collections, unpublished field accounts, and images posted online. The loss of this emblematic species is unique among Californian nudibranchs, including (1) its congener Felimare porterae (=Mexichromis porterae), with which it appears to overlap in diet, and (2) opisthobranch species with similar historical geographic ranges and mode of development. The decline in F. californiensis is not predicted by warming trends and climate variation over the past 40 years, including the strong El Niño events of 1983 and 1998. Coastal pollution from the large human population in southern California may have impacted Dysidea amblia, the primary reported prey of F. californiensis. Historical overcollecting of the nudibranch and habitat loss through the development of major ports likely also contributed to its decline. Sightings since 2003 are consistent with a nascent recovery, as elements of water quality have improved in the SCB in recent decades.     

Spawning behaviour and egg capsules of Concholepas concholepas (Mollusca: Gastropoda: Muricidae)

Observations on the spawning behaviour of the commercially important muricid snail Concholepas concholepas are reported. The sequence of events relating to external transportation, molding and hardening, and attachment of the egg capsules to a substratum were timed and divided into 6 major events. The snails studied in the laboratory spawned between April and Septemer. These results and data from the literature reveal an all-year-round spawning activity for the species. The frequency, size and time of spawning were studied over 1 year. Spawning activity always began at night, suggesting that the species retains its spawning rhythm in the laboratory. These was a direct relationship between length of the snail and size of the capsules spawned. No nutritive eggs were found; the number per capsule varied between 668 and 14,250 according to capsule size. Hatching time varied between 69 and 128 days at 13.5° to 14.5°C.     

Biodiversity in Littorina species (Mollusca: Gastropoda): a physiological approach using heat-coma

 The upper critical thermal limits were measured as heat-coma temperatures in Littorina species as a tool for examining physiological diversity. Thermal tolerance was found to be stable within a population, but extensive variation occurred between populations and species. All species examined displayed heat-coma values of ∼30 °C and did not show a positive correlation with shore height. Eulittoral-fringe species tended to have higher coma temperatures than eulittoral species. Coma temperatures varied both seasonally and geographically. Lethal thermal limits were also investigated; these were ∼10 C° higher than coma temperatures. Received: 16 August 1999 / Accepted: 13 June 2000     

Evolution and systematics in Haliotidae (Mollusca: Gastropoda): inferences from DNA sequences of sperm lysin

Abalone taxonomy and systematics have remained unresolved: neither stable species-level nomenclature nor a cladistic hypothesis of relationships among species have been established. To infer the phylogeny of the genus Haliotis and to identify species using molecular data, we compared complementary DNA (cDNA) sequences of sperm lysin from 27 species-group taxa from California, Japan, Australia, New Zealand, Taiwan, Borneo, Madagascar, South Africa, Greece, France, Italy and the Azores. The lysin cDNA sequences reveal that 22 of the 27 taxa are clearly distinguishable by >20 nucleotide differences. Of the remaining 5, H. coccinea from the Azores may be a subspecies of H. tuberculata, if not a sibling species (10 nucleotide differences). The other four taxa are most probably the same species as one of the 22 taxa: the lysin sequences are almost identical between H. madaka and H. discus hannai, H. conicopora and H. rubra, H. diversicolor supertexta and H. diversicolor aquatilis, and H. tuberculata lamellosa and H. tuberculata tuberculata. The phylogeny of lysin cDNA suggests that there are three groups among the 27 species-group taxa: (1) all California species and 3 Japanese species (H. gigantea, H. discus hannai, and H. madaka): (2) 1 New Zealand species (H. iris); (3) 1 Japanese species (H. diversicolor aquatilis), Indo-West Pacific species and European species. These groups can be assigned to three previously recognized subgenera (Nordotis, Paua and Padollus) in the genus Haliotis. Two historical hypotheses are proposed to explain the biogeography and evolution within these abalone: (1) Tethyan distribution of the ancestral abalone, during the Cretaceous, followed by extinction in most of the habitat, but radiation in California and Southeast Asia which later spread to the other areas; (2) North Pacific rim distribution of the ancestral abalone, followed by dispersal to the other areas during the Paleogene.     

Seasonal changes in the biochemical composition of the chiton Chiton iatricus (Polyplacophora: Mollusca) and the marine pulmonate Onchidium verruculatum (Gastropoda: Mollusca) in relation to their reproductive cycles

Seasonal changes in the biochemical composition of two molluses, gonochoric Chiton iatricus and a marine pulmonate, Ochnidium verruculatum inhabiting the same tropical area, but having different modes of reproduction, were studied. Protein is the major biochemical component stored in C. iatricus, whereas lipid is predominantly stored in O. verruculatum. The correlation between these quantifications and the annual reproductive cycles of the two species investigated suggest that nutrients stored in the somatic body parts are channeled towards the gonad during the gametogenesis. In C. iatricus sexual differences efrist in the biochemical needs of the ovary and the testis. The ovary always has a higher protein content than the testis. In the hermaphrodite pulmonate O. verruculatum, biochemical expenditure on gamete production is relatively low because this species has to spend energy on the reproductive propagation viz. copulation and egg-laying. On the contrary, in C. iatricus the biochemical energy is mainly utilized for the production of gametes which have to be produced in great numbers in order to counter-balance the loss of gametes caused by external fertilization. The collected data are discussed in the light of comparative physiology of reproduction.     

Chemical defense and evolutionary trends in biosynthetic capacity among dorid nudibranchs (Mollusca: Gastropoda: Opisthobranchia)

Summary. An evolutionary scenario incorporating recent advances in phylogenetic research begins with an opisthobranch-pulmonate common ancestor that was herbivorous and had some diet-derived chemical defense. The Nudibranchia and their closest relatives, the Notaspidea, form a lineage the ancestors of which had switched to feeding upon sponges and deriving protection from metabolites contained in them. Subsequently there have been repeated shifts in food and defensive metabolites, and trends are evident in the ability to detoxify, sequester and utilize metabolites from food, as well as to synthesize defensive compounds de novo. The Notaspidea display a minor adaptive radiation that foreshadows a more extensive one in the various lineages of nudibranchs. This review emphasizes changes that have occurred within the Holohepatica, or dorid nudibranchs (order Doridacea). Their sister-group, the Cladohepatica, consists of three other orders, Dendronotacea, Arminacea, and Aeolidiacea, in which there has been a shift from sponges to Cnidaria as food. The Dendronotacea often feed upon Octocorallia, which combine spicules, chemical defense, and stinging capsules and thereby suggest a transition from feeding on sponges. A previous diet of Octocorallia is suggested by the defensive use of prostaglandins in the dendronotacean Tethys fimbria, which eats crustaceans. A shift to bryozoans in some Arminacea is accompanied by use of different metabolites. Dorid nudibranchs evidently began as sponge-feeders, but some lineages have shifted to a variety of other food organisms, and others have specialized in the kind of sponges they feed on and how they do it. There have been shifts to bryozoans (Ectoprocta) and ascidians (Chordata: Urochordata) that track metabolites rather than the taxonomy of the food. There is a crude correlation between the genealogy and the defensive metabolites of the sponge-feeding dorids. De novo synthesis is well documented in this order and the metabolites are appropriately positioned so as to have an adaptive effect. The hypothesis that the capacity for de novo synthesis was acquired by gene transfer across lineages is rejected, partly on the basis of different chirality of metabolites in the nudibranchs and their food organisms. Instead it is proposed that there has been a preadaptive phase followed by evolution in a retrosynthetic mode, with selection favoring enzymes that enhance the yield of end products that are already present in the food. Received 5 February 1999; accepted 26 July 1999     

Competition and interaction between Neosarmatium smithi (Crustacea: Grapsidae) and Terebralia palustris (Mollusca: Gastropoda) in a Kenyan mangrove

 In a Kenyan mangrove, we studied the interactions between the gastropod Terebralia palustris and the crab Neosarmatium smithi when foraging on decaying mangrove leaves. Interactions are considerable on account of their wide overlap in zonation (Rhizophora mucronata belt), food items (mangrove leaves) and activity window (diurnal low water). The snails find a leaf by a systematic transecting of the platform and eat the leaf after crawling on to it, usually en masse. The crabs rarely venture beyond 80 cm from their burrows, and once they find a leaf they quickly drag it back into their burrow, probably to reduce the strong intra-specific competition. A crab is able to drag a leaf away even if a number of snails are already feeding on it, by strongly pulling the leaf away or by pushing the snails off the leaf. The success of the “thief ” depends on both the crab's size and the number of snails on the leaf. Received: 25 July 1999 / Accepted: 27 April 2000     

Factors affecting variability of foraging excursions in a population of the limpet Patella vulgata (Mollusca,Gastropoda)

Paths moved by the limpet Patella vulgata L. were monitored on a sheltered vertical rocky shore in North Wales using time-lapse photography throughout nocturnal low tides in April and November 1992, roughly corresponding to periods of minimum and maximum gonad ripeness, respectively. Various motion parameters, including total duration, total length, and maximum distance reached from home were computed from 124 complete foraging routes obtained from 18 higher-zoned and 18 lower-zoned limpets. P. vulgata typically performed a single loop per night (average total length, ca. 70 cm), moving from the individual home scar to graze the surrounding algal grounds (average maximum distance, ca. 25 cm), then following its own trail back home. On the average, limpets moved from the home scar during three of four available nocturnal low tides and exploited about half the emersion time. Evidence for size-related variability in behaviour was found, as a positive correlation was assessed between both total duration and legth of the excursions, and shell length. In addition, zonation was proved to affect the limpets' foraging behaviour, since low-shore limpets moved faster and covered greater distances than high-shore ones. Moreover, consistent seasonal variation in foraging behaviour emerged, since in April the excursions were longer and longer-lasting than in November, but limpets exploited a larger fraction of potential activity phases in November than in April. The within-population variability in the temporal and spatial characteristics of the foraging excursions is discussed in relation to the available data on zonal and seasonal variation of food resources and in relation to physiological changes due to reproductive cycle.     

Veränderungen der Eikapselvolumina während der entwicklung verschiedener Opisthobranchier-Arten (Mollusca,Gastropoda)

An attempt has been made to estimate volumes of eggs and egg-capsules employing the formula of the rotationellipsoid. Measurements of the following stages have been made: immediately after oviposition; blastula; gastrula; appearance of velumanlage; velarcilia, foot and velum small; velarcilia, foot and velum distinct; 1 to 2 days before hatching; hatching. Calculations permitt analysis of: (1) relationship between egg-case volume and egg-volume; (2) capsule volumes during development until hatching in comparison to those after oviposition (enlargement factor). The sacoglossan nudibranch Elysia viridis shows no increase in capsule volume during development. The nudibranchs Catriona aurantia, Tergipes despectus, Eubranchus exiguus, Eubranchus cingulatus, Eubranchus pallidus, Eubranchus farrani and Janolus cristatus reveal a distinct increase in capsule volume when velarcilia, foot, and velum appear. The degree of eggcapsule enlargement and its intensity during development exhibit interspecific differences. The results of this and of a previous paper (Kress, 1971) are compared and discussed with respect to possible explanations for the enlargements, (e.g. excretion of larval kidneys, composition of capsular fluid, influence of salinity).     

Factors affecting fertilization success in two species of patellid limpet (Mollusca: Gastropoda) and development of fertilization kinetics models

Laboratory experiments were undertaken to examine fertilization success in the intertidal prosobranch limpets Patella ulyssiponensis and Patella vulgata. Alkalization of eggs (10 min in pH 9.0 sea water) prior to fertilization trials improved fertilization rates greatly. Fertilization success was found to be a function of sperm concentration, gamete age and contact time. Sperm concentration needed for optimum fertilization success in vivo ranged between 10⁵ and 10⁷ sperm ml⁻¹ for both species although at higher concentrations the number of normally developing trochophore larvae decreased. For P. vulgata, sperm longevity (at a concentration of 10⁶ sperm ml⁻¹) did not exceed 6 h, whereas eggs of both species were fertilizable for up to 12 h. Maximum fertilization success occurred after 15–30 min gamete contact time. The Vogel et al. (Math Biosci 58:189–216, 1982) fertilization kinetics model is developed to allow for non-complete fertilizations under optimal sperm concentrations, and a new parameter fitting technique is developed to improve estimates of fertilization success for short gamete contact times.     

Spread ofCyclope neritea (Mollusca: Gastropoda) along the north-eastern Atlantic coasts in relation to oyster culture and to climatic fluctuations

The geographic distribution ofCyclope neritea (Linné) (Mollusca: Gastropoda) along the Atlantic coasts of Portugal, Spain and France is reported. Analysis of information published over the last 90 yr indicates that, since 1983,C. neritea has extended its range northwards from Arcachon Bay along the coast of the Bay of Biscay. The spread of this small gastropod is directly related to the commercial transport of the Pacific cup oyster (Crassostrea gigas Thunberg) to the Atlantic from Mediterranean oyster-culture sites, and to its encountering climatic conditions suitable for its settlement and reproduction in its new habitat. Its transport by human intervention explains the rapid spread ofCyclope neritea during 1983 and 1984, despite its lack of a planktonic stage or of effective natural migration by adults. Climatic conditions in the southern parts of the Bay of Biscay are similar to those of the French Mediterranean lagoons, explaining the easy acclimatization of this eurythermic gastropod. Also, coastal air and sea-surface temperature records over the last two decades (1971–1988) have revealed a slight warming of the southern parts of the Bay of Biscay. This has resulted in a more strongly marked latitudinal thermal gradient along the Bay of Biscay and has furthered survival and reproduction ofCyclope neritea emigrants. A detailed analysis of commercial supplies of cup oysters to the Bay of Biscay from the Mediterranean coast indicates that theC. neritea emigrants undoubtedly originated from the salt-water lagoons of Salse-Leucate and Thau (French Mediterranean coast).     

Aspects of the ecology and growth of an intertidal juvenile population of Concholepas concholepas (Mollusca: Gastropoda: Muricidae) at Las Cruces,Chile

The habitat, density and growth rate for an intertidal population of Concholepas concholepas (Bruguière, 1789) were studied at Las Cruces, Chile (Lat. 33°30′S; Long. 71°38′W) during 1977–1978. The growth rate (3.67 mm month^-1) was determined in a newly settled group, whose average length was 11.3 mm, and whose age was estimated at 3 months. The densities found ranged between 1.1 and 107.3 “locos” m^-2. Based on these results, the time of settlement of C. concholepas was calculated; capsule deposition, maximum maturity and recruitment reported by other authors for different localities are discussed.     

Ingestion of a medusa (Aegina citrea) by the nematocyst-containing ctenophore Haeckelia rubra (formerly Euchlora rubra): phylogenetic implications

The rare ctenophore Haeckelia rubra (formerly Euchlora rubra) has long been known to have nematocysts rather than colloblasts in its tentacles. Five specimens were collected in the San Juan Archipelago, Washington State, USA in 1980 and 1981, and their feeding behavior was observed in the laboratory. We found that H. rubra readily eats the tentacles of a medusa, Aegina citrea, whose nematocysts (apotrichous isorhizas) are nearly identical in morphology to the nematocysts of the ctenophore. When H. rubra was offered 16 other species of hydromedusae and 1 siphonophore in the laboratory, the ctenophores showed little or no tendency to ingest these potential prey items. In addition to its routinely positive response to A. citrea, the ctenophore could be induced by manipulation and starvation to accept and ingest bits of the bodies of 4 additional species of hydromedusae and 1 siphonophore. These results, combined with the histological and rearing experiments of other investigators, leave little doubt that the nematocysts in H. rubra are not endogenous, but are kleptocnidae similar to those nematocysts retained and subsequently used by some species of nudibranchs that feed on Cnidaria. A close phylogenetic link between the Cnidaria and the Ctenophora is most unlikely.     

Laboratory rearing of Sepiolinae (Mollusca: Cephalopoda)

Five species of Sepiola and Sepictta were reared in the laboratory from egg to adult size. Spawning was achieved in 3 species of Sepiola after 5 to 7 months. The growth rate of the species reared did not depend upon temperature, which ranged from 12.5° to 20°C. A fairly constant size increase (2.5 mm mantle length/month) was observed in Sepiola during the 5 months after hatching. In Sepietta, the same growth rate was observed until the fourth month after hatching, when it increased to the rate of 5 mm mantle length/month.Laboratoire associé au C.N.R.S.     

Growth,reproduction and mortality of the sea hare Dolabella auricularia (Gastropoda: Aplysiidae) in the Central Visayas,Philippines

The growth parameters W and K of the von Bertalanffy growth equation were estimated from size-frequency data of the sea hare Dolabella auricularia using an objective, computer-based method. The results obtained from two locations in the central Philippines were comparable, although based on only a few individuals; the means for both sites were W=493 g and K=0.9 (yearly basis). A value of Z=3.66, corresponding to an annual survival rate of 2.6%, was estimated for the juveniles and adults from a length-converted catch curve. Spawning and recruitment were found to occur throughout the year with peaks in May to July and September to October.ICLARM Contribution No. 170