Morphological patterns in high-intertidal gastropods: Adaptive strategies and their limitations

An analysis of shell form in several families of high-intertidal herbivorous gastropods has revealed the existence of morphological gradients. Among tropical littorinids, relative spire height and degree of development of external shell sculpture generally increase interspecifically, and often intraspecifically, from low to high shore levels. This gradient is also evident from temperate to tropical latitudes, and may be modified or overridden by exposure to wave action and other factors. Among acmaeid, patellid, and siphonariid limpets, there is a well-marked interspecific increase in relative shell height with increasing shore level, but latitudinal trends in morphology cannot at present be distinguished. Evidence from both tropical and temperate coasts indicates that limpets found in shaded places tend to have smoother shells than those often exposed to direct sunlight. In the Neritidae, relative shell globosity tends to increase interspecifically from low to high shore levels; most high-shore species have strong shell sculpture, with the base of the shell only slightly excavated. A decrease in the number of neritid species towards higher latitudes is accompanied by selective deletion of globose, strongly sculptured types living at high shore levels. Neritids and some limpets can employ evaporative cooling as a mechanism for temperature regulation. The morphological gradients in these groups reflect a relative increase in the volume of the extravisceral cavity (water reservoir) and a decrease in the area in contact with the substratum, as well as a relative decrease in the rate of water loss, in an upshore direction. Neritids are better adapted than limpets to high temperature and desiccation stresses because of their ability to regulate rate of evaporation and contact between soft parts and substratum. These differences are reflected in the complementary patterns of latitudinal diversity in the two groups. In Littorinidae, which attach to the substratum by a mucus film, the morphological gradients reflect a minimization of water loss and substratum contaet, and a maximization of reflective relative to absorptive surface area.This paper is an expanded version of part of a Ph. D. dissertation presented to Yale University.     

Conspecific brood parasitism in an upland-nesting bird: cues parasites use to select a nest

Behavioral Ecology and Sociobiology - In the animal kingdom, conspicuous colors are often used for inter- and intra-sexual communication. Even though primates are the most colorful mammalian taxon,...     

Genetic variability in the marine gastropods Patella coerulea and Patella aspera: patterns and problems

Allozymic variation encoded by 15 gene loci was compared and contrasted in naturally coexisting populations of two marine gastropods (Patella coerulea and Patella aspera) collected along the rocky beaches of the Mediterranean sea of Israel in June, 1979. The genetic variability did not differ consistently or significantly between the two species, but in all the populations studied, P. aspera was more heterozygous than P. coerulea. These results are discussed in terms of the potential ecological factors affecting this genetic differentiation.     

Hatching mode and latitude in marine gastropods: revisiting Thorson's paradigm in the southern hemisphere

For much of the last century the developmental mode of marine invertebrates, particularly of prosobranch gastropods, has been thought to reflect a latitudinal pattern known as Thorson's rule; the proportion of species with pelagic larvae decreases with increase in latitude. Although the predictions of this rule have been criticized recently, its validity along latitudinal transects in the world ocean still remains poorly tested. In the present work, we compare the frequency of occurrence of contrasting prosobranch hatching modes (pelagic versus benthic development) along two latitudinal gradients of the subequatorial coastline of South America: the southeast Pacific and the southwest Atlantic. The results are clearly contrasting. While the pattern observed along the Pacific (Chilean) coast fits the predictions of Thorson's rule very well, benthic development predominates all along the Atlantic coast, even at subtropical latitudes. This difference in observed patterns is attributable to the different compositions of the gastropod assemblages on each side of the South American continent, which are determined, in turn, by differences in the ecological conditions on the two sides of the continent. The scarcity of pelagic development among the Atlantic prosobranch gastropods reflects the near-continuous soft-bottom habitat there, and the consequent prevalence of predaceous, soft-bottom taxa that had acquired in their evolution more evolved, non-pelagic patterns of development. In contrast, the Pacific coast is much more heterogeneous, with a diverse mixture of benthic habitats, including rocky substrates that, in part, support grazing taxa. Our results indicate that two factors are important for Thorson's rule to be valid. First, the habitat needs to include rocky substrates, as soft-bottom habitats appear to favour non-pelagic development. Second, a diverse assemblage of taxa need to be compared to avoid the problem of phyletic constraints, which could limit the evolution of different developmental modes. Received: 23 May 2000 / Accepted: 13 October 2000     

Interplay of host morphology and symbiont microhabitat in coral aggregations

The Belizean reef coral Agaricia tenuifolia Dana forms aggregations in which rows of thin, upright blades line up behind each other. On average, the spacing between blades increases with depth and hence with decreasing ambient irradiance. We designed and built a small, inexpensive light meter and used it to quantify the effect of branch spacing on light levels within colonies at varying distances from branch tips. Concurrently, we measured photosynthetic pigment concentrations and population densities of symbiotic dinoflagellates (zooxanthellae) extracted from coral branches of colonies with tight (≤3 cm) vs wide (≥6 cm) branch spacing, collected at 15 to 17 m and from colonies with tight branch spacing collected at 1 to 2 m. Light levels decreased significantly with tighter branch spacing and with distance from the branch tips. Total cellular pigment concentrations (chlorophylls a, c ₂ and peridinin) as well as chlorophyll a:c ₂ and chlorophyll a: peridinin ratios all increased significantly with distance from the branch tip, indicating very localized differences in photoacclimation within individual branches. Zooxanthellae from colonies with widely-spaced branches displayed significantly lower chlorophyll a:c ₂ and chlorophyll a:peridinin ratios, and were present at significantly higher population densities than those from colonies with tightly-spaced branches collected at the same depth (15 m). Tightly-spaced colonies collected from shallow environments (1 to 2 m) displayed pigment ratios similar to those from widely-spaced colonies from deeper water (15 m), but maintained zooxanthellae populations at levels similar to those in tightly-branched colonies from deeper water. Thus, variation in colony morphology (branch spacing and distance from branch tip) can affect symbiont physiology in a manner comparable to an increase of over 15 m of water depth. These results show that a host's morphology can strongly determine the microhabitat of its symbionts over very small spatial scales, and that zooxanthellae can in turn display steep gradients in concordance with these altered physical conditions. Received: 12 June 1997 / Accepted: 24 June 1997     

Ecological and evolutionary significance of pelagic spermatophores of vermetid gastropods

Sperm are disseminated in vermetid gastropods by spermatophores released freely into the ocean. Spermatophores from 7 species of vermetids, including 3 genera, were obtained from reproductively active males or offshore plankton tows or both. Each vermetid spermatophore consists of a sperm mass containing highly ordered eupyrene and apyrene sperm enveloped concentrically by 3 transparent capsules. Generic and specific differences occur in the size, shape and composition of the sperm mass and capsule. The complexly layered spermatophore is assembled in the male reproductive tract, which includes and elaborate set of pallial reproductive glands for capsule production. Spermatophores are liberated by males into the seawater, and there dispersed by water movements. Some of these drifting spermatophores become entangled in the mucous feeding nets of female vermetids and are then activated by feeding movements. Activation of the complex ejaculatory apparatus appears to result from an interplay of osmotic and mechanical mechanisms. Spermatophores remain viable in the laboratory for 12 to 20 h after release from the male; after 20 h, the sperm mass breaks down and the sperm become immotile. Plankton tows capture some spermatophores with motile sperm and others in which sperm are no longer active. Evolutionary implications of the relationship between production of pelagic spermatophores, sessile mode of life and other aspects of vermetid biology are discussed.     

Geographic range limitation of the intertidal gastropods Littorina sitkana and L. planaxis

Geographic ranges of marine intertidal organisms are not entirely governed by discontinuities in temperature gradients. This is especially true in the eastern north Pacific Ocean, where temperature gradation is less steep than in the western north Atlantic Ocean. The southern limit of distribution of Littorina sitkana (Philippi, 1845) and the northern limit of L. planaxis (Philippi, 1847) occur at Charleston, Oregon (Latitude 43.4°N). No discontinuity in air or water temperature can be correlated with these distributional boundaries. A third species, L. scutulata (Gould, 1849), has the widest range, overlapping with L. planaxis in the south and L. sitkana in the north (Latitude 27.5° to 58°N). L. planaxis transplanted to northern Washington survived for 4 years, grew and produced planktonic larvae. Either larval mortality or south-flowing currents during planktonic life prevent this species from dispersing northward. L. sitkana has direct development and occurs only in habitats where adults are sheltered from waves and egg masses from desiccation. Lack of suitable habitat, coupled with the presence of the predatory crab Pachygrapsus crassipes (Randall, 1839) prevent the southward establishment of L. sitkana.     

Seasonality and the evolutionary divergence of plant parasites

The coexistence of closely related plant parasites is widespread. Yet, understanding the ecological determinants of evolutionary divergence in plant parasites remains an issue. Niche differentiation through resource specialization has been widely researched, but it hardly explains the coexistence of parasites exploiting the same host plant. Time-partitioning has so far received less attention, although in temperate climates, parasites may specialize on either the early or the late season. Accordingly, we investigated whether seasonality can also promote phenotypic divergence. For plant parasites, seasonality generally engenders periodic host absence. To account for abrupt seasonal events, we made use of an epidemic model that combines continuous and discrete dynamics. Based on the assumption of a trade-off between in-season transmission and inter-season survival, we found through an "evolutionary invasion analysis" that evolutionary divergence of the parasite phenotype can occur. Since such a trade-off has been reported, this study provides further ecological bases for the coexistence of closely related plant parasites. Moreover, this study provides original insights into the coexistence of sibling plant pathogens which perform either a single or several infection cycles within a season (mono- and polycyclic diseases, or uni- and multivoltine life cycles).     

Measuring terrestrial movement behavior using passive integrated transponder (PIT) tags: effects of tag size on detection, movement, survival, and growth

Movement behaviors have broad ecological and evolutionary implications, affecting individual fitness, metapopulation dynamics, the distribution and abundance of species, as well as gene flow and thus adaptation and speciation. However, movement behaviors such as dispersal, station keeping, and ranging are poorly understood in many taxa due to the incompatibility of traditional tracking methods with long-term observations. This is particularly true for small-bodied life history stages and species. While the introduction of smaller passive integrated transponder (PIT) tags and the development of PIT telemetry have removed some barriers, the trade-offs between different tag sizes are unknown. Through a series of experiments, we tested for effects of PIT tag size on detection, movement, tag retention, growth, and survival of a juvenile amphibian. We found no effect of PIT tag size on initial movement distance, survival, or growth; and all individuals retained their tag for the course of the experiment. Detection and recapture rates, however, were increased with PIT tag size. The orientation of the tag relative to the vertical axis of the antenna also affected the size of the detection field, which was 15.78–43.90 % smaller when the antenna was moved perpendicular rather than parallel to the long axis of the tag. We conclude that PIT telemetry is a suitable technique for marking previously untraceable species or life history stages and may offer insight into the behaviors of these individuals. Investigations using multiple PIT tag sizes should include this in statistical analyses to account for tag size biased detection differences.     

The effects of parasites on male ornaments and female choice in the lek-breeding black grouse (Tetrao tetrix)

Summary We describe the results of two studies of parasitic infection in the black grouse (Tetrao tetrix). The first deals with our own observations of lekking black grouse in which the parasite levels of two blood parasites, the protozoan Leucocytozoon lovati and microfilaria, probably produced by a nematode worm Splendidofilaria tuvensis, were scored. We also obtained measures of age, survival, number of copulations, body mass and length of the ornamental tail feathers (the lyre) of the lekking males. The second study analysed the data from Lund (1954) which involved eight gut parasites obtained from birds which were killed. In the first study we found higher levels of infection of Leucocytozoon in adults relative to young birds. Parasites had no effect on male survival and there was no correlation in infection between the two types of parasites. Birds infected with microfilaria had shorter tail ornaments. There was no relationship between parasitic infection and mating success. However, the data indicated that such a trend indeed may exist for Leucocytozoon and the most successful males on the leks were less often infected by Leucocytozoon than other males. Results of the second study showed a negative relationship between parasite load (a combined measure of all parasites) and both ornamental tail feather length and body mass. These observations are compatible with, but not conclusive evidence for, the hypothesis of Hamilton and Zuk (1982) on the evolution of secondary sexual characters, where females choose to mate with genetically resistant males which show their resistance by expressing larger and more showy secondary sexual characters. Alternative explanations for the observed patterns are: females avoid infected males for some immediate benefit; and/or parasite loads are indicators of general stress rather than genetical resistance. Under the latter hypothesis females could mate with more vigorous males for reasons unrelated to parasite resistance.     

Satin bowerbird parasites: a test of the bright male hypothesis

Summary The number of a common parasite (Cuclotogaster sp.) on male satin bowerbirds was related to male mating success in a test of Hamilton and Zuk's (1982) bright male hypothesis. The data do not show the expected inverse correlation between female mating preferences and the level of parasitic infection of males predicted by that model. Nearly all matings are accomplished by bower-holding males (Borgia 1985a), but the vast majority of these males were uninfected. There were large differences in mating success among the uninfected bower holders, but this could not be explained by between male differences in the level of parasitic infection. From this I conclude that levels of parasitic infection are not now an important direct cause of intermale variation in mating success. The results are, however, consistent with a hypothesis that a low level of infection is indicative of the overall healthy condition of a male. If this is true, it supports the hypothesis that the ability to hold a bower may be an indicator of male condition to females.     

Size-related selection of two intertidal gastropods by the reef crab Ozius truncatus

Reef crabs, Ozius truncatus H. Milne-Edwards 1834, and gastropods, Bembicium nanum (Lamarck, 1882) and Nerita atramentosa Reeve, 1855, were collected from a South Australian rocky intertidal platform in April 1981. Laboratory experiments were conducted to determine (1) the size of prey chosen, and (2) the energy gained by the reef crabs feeding on B. nanum and N. atramentosa. The time taken to break open and consume prey increased exponentially with prey shell height. Crabs with larger master claws had shorter handling times for a given prey size. When offered a range of prey sizes, crabs selected prey of a certain size, well below the maximum prey size that they could handle. The mean prey size selected by crabs did not concide with the size predicted to yield the maximum rate of energy gain. For example, one reef crab preferred B. nanum 5 to 8 mm in size, but the size class predicted to yield the greatest energy gain was 4 to 4.9 mm. The results of the laboratory prey-choice experiments are discussed in relation to the shore-level size gradients of B. nanum and N. atramentosa.     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Dissecting the drivers of population cycles: Interactions between parasites and mountain hare demography

There is a growing awareness that cyclic population dynamics in vertebrate species are driven by a complex set of interactions rather than a single causal factor. While theory suggests that direct host-parasite interactions may destabilise population dynamics, the interaction between host and parasite may also influence population dynamics through indirect effects that result in delayed responses to either density or to life-history traits. Using empirical data on mountain hares (Lepus timidus) infected with a nematode parasite (Trichostrongylus retortaeformis), we developed an individual-based model (IBM) that incorporated direct effects and delayed life-history effects (DLHEs) of a macroparasite, alternative transmission mechanisms and seasonality in host population dynamics. The full model describes mean characteristics of observed mountain hare time series and parasite abundance, but by systematically removing model structure we dissect out dynamic influences of DLHEs. The DLHEs were weakly destabilising, increasing the propensity for cyclic dynamics and suggesting DLHEs could be important processes in host-parasite systems. Further, by modifying model structure we identify a strong influence of parasite transmission mechanism on host population stability, and discuss the implications for parasite aggregation mechanisms, host movement and natural geographical variation in host population dynamics. The effect of T. retortaeformis on mountain hares likely forms part of a complex set of interactions that lead to population cycles.     

Stable isotope profiles from subtropical marine gastropods of the family Fasciolariidae: growth histories and relationships to local environmental conditions

Oxygen and carbon stable isotope profiles were constructed for two species of large subtropical gastropods of the family Fasciolariidae—Triplofusus giganteus and Fasciolaria tulipa—from the Florida Keys and the Bahamas, to evaluate their life history and to assess their potential as paleoenvironmental proxies. Oxygen isotope profiles revealed T. giganteus and F. tulipa grew their shells for 6 and 3 years, respectively. Both mollusks show faster growth rates during the first half of their lifespan. Mean annual temperatures (MAT) derived from oxygen isotopes for T. giganteus were 26.5 °C and for F. tulipa were 26.7 °C, both matching instrumental MATs of 26.7 and 26.5 °C for the Florida Keys. Both shells, however, failed to record entire mean annual temperature ranges (MART). Fasciolaria tulipa yielded a calculated MART of 5.6 °C compared with a measured MART of 9.3 °C, and T. giganteus showed a calculated MART of 6.9 °C compared with a measured MART of 9.4 °C. Carbon isotopes of T. giganteus were ambiguous and reveal no significant relationships with trends in nutrient concentrations (N and P), dissolved oxygen, and dissolved organic carbon, although they did exhibit more negative values concomitant with landfall of Hurricane Irene and trended to increasing values with ontogeny that could reflect migration. Carbon isotopes in F. tulipa were lower during winters, possibly reflecting seasonal upwelling or seagrass-mediated carbon cycling.     

Genetic variability in littoral gastropods: Phosphoglucose isomerase and phosphoglucomutase in Patella vulgata and P. aspera

Genetic variability of the phosphoglucose isomerase (PGI) and phosphoglucomutase (PGM) loci are compared in Patella vulgata L. and P. aspera Lamark from two sites on the west and south coasts of Ireland. P. vulgata was less variable than P. aspera at these 2 loci, and in only 1 of 7 other loci briefly examined was P. vulgata more variable. Evidence is also presented that the PGI enzyme in P. vulgata is more stable at high temperature than that of P aspera. The significance of these observations is discussed in the light of, the different zonal distributions of these species on the shore.     

Signal use by collared lizards, Crotaphytus collaris: the effects of familiarity and threat

Many lizard species use lateral compressions of the body during agonistic encounters. I investigated the signal value of the frequency at which these displays are presented and how that rate is affected by familiarity and threat. The response of resident collared lizards (Crotaphytus collaris) to tethered strangers, neighbors on the correct boundary, and neighbors displaced to the opposite boundary, was recorded by counting the number of lateral displays directed at the intruder and the number of subsequent acts of escalated aggression. There was no relationship between size asymmetry of the opponent and the rate of lateral display or aggression, nor was there a relationship between body size and the rate of lateral display or aggression. However, there was a high positive correlation between lateral display rate and aggression, suggesting that the rate of lateral displays is a conventional signal of motivation to attack. The highest rates of display and aggression were directed toward displaced neighbors, somewhat less toward strangers, and the least toward neighbors at the correct boundary. The ratio of aggressive acts to lateral displays followed the same pattern, presumably because the perceived threat to the resident decreases in the same order. Taken together these data suggest that collared lizards are able to assess the threat of an opponent and signal motivation to respond aggressively towards that opponent.Communicated by S. Downes