EFFECTS OF WOUNDING BY THE HERBIVOROUS SNAILS LITTORINA SITKANA AND L. SCUTULATA (MOLLUSCA) ON GROWTH AND REPRODUCTION OF THE INTERTIDAL ALGA FUCUS DISTICHUS (PHAEOPHYTA)1

Small (3–7 cm long) Fucus distichus ssp. edentatus (de la Pyl.) Pow. Plants were tagged at three sites in which densities of the herbivorous snails Littorina sitkana and L. scutulata ranged from 367–4690 animals · M^-2. From April–August 1986, the growth rate, degree of wounding, and reproductive status of individual thalli were monitored at 2–4 week intervals. Grazer-inflicted damage to the thalli varied within and among sites. Mean growth rates at the site with low densities of littorines were about twice those at the site with intermediate densities and about four times those at the high density site. At the site with high densities of littorines, F. distichus growth rates were negatively correlated with the degree to which the plants were wounded. There appeared to be no correlation of grazer density with F. distichus survivorship. Thalli at the site with few herbivores tended to reproduce earlier and at a larger size than did those at the other two sites. In all three areas, only thalli that had received little damage from herbivores became reproductive. By lowering growth rates and delaying reproduction in F. distichus, grazing by littorine snails can potentially cause variation in reproductive output among individual thalli.     

Predation induced changes in behavior and growth rate in three populations of the intertidal snail, Littorina sitkana (Philippi)

We investigated the sublethal effects of a predatory crab, Cancer productus (Randall), on the behavior and growth of its snail prey, Littorina sitkana, by setting up controlled rearing and prey-size selection experiments. L. sitkana were collected from three sites on San Juan Island, WA, USA. These sites varied in snail size, abundance, and vertical distribution, and in the abundance of the crab predator C. productus. Snails from all three populations were raised for 34 days under the following treatments: no-crab control, a non-feeding C. productus encased in mesh box, and an encased C. productus feeding on L. sitkana. The non-feeding crab treatment did not affect snail foraging behavior or growth rate in comparison with the no-crab control. In contrast, the presence of a feeding crab elicited escape behavior in the snails, halted grazing, and consequently reduced growth rates. A population difference in escape behavior was observed: upward migration in snails from rocky shores and hiding in crevices in snails from a mud flat. It thus appears that chemicals leaching from crushed conspecific snails, rather than the presence of the crab predator, act as the “alarm substance” to which L. sitkana react. The magnitude of the growth depression in the presence of feeding crabs was 85%, with no difference among the three populations. Once the feeding crab stimulus was removed, snails in all populations resumed normal growth, suggesting that this response to feeding predators is reversible with changing environmental conditions. Laboratory experiments were set up to determine if all size classes of L. sitkana are equally susceptible to C. productus predation. C. productus consistently selected the largest of three size classes of L. sitkana. These results suggest that slow growth rate and small size in L. sitkana may actually be an adaptation for coexisting with high C. productus abundance, rather than simply a cost of escape behavior.     

The maintenance of shore-level size gradients in an intertidal snail (Littorina sitkana)

Summary The size of many intertidal animals varies with tidal height. These size gradients could be produced by growth or survival varying with tidal height, or by animals moving to a preferred tidal level. The body size of the snail, Littorina sitkana, increases steadily with tidal height in rocky high intertidal habitats of British Columbia. To determine how size gradients were maintained in L. sitkana, I quantified how growth, survival, and snail movement varied with tidal height. I studied populations of L. sitkana found on sheltered pebble beach and exposed basaltic shelf habitats. Mark-recapture studies and experimental transplants showed that growth could not have produced the size gradients because snail growth rates in both habitats were as fast or faster at low tidal levels (where the snails were the smallest) than at high tidal levels. However, survival rates were lowest at low tidal levels. On pebble beaches, this was due to size selective predation on large snails by the pile perch, Rhacochilus vacca. On basaltic shelves, heavy wave action at low tidal levels may have caused the poor survival rates. Transplanted snails moved homeward on pebble beaches, but not on basaltic shelves. Reduced survival rates at low tidal levels cause size gradients in both habitats, and snail movement helps to maintain size gradients on pebble beaches.     

Allozyme comparison of four littorinid species morphologically similar to Littorina sitkana

Eight species of the genus Littorina were hitherto recognised in the north-western region of the Pacific Ocean: L. sitkana, L. brevicula, L. mandshurica, L. squalida, L. aleutica, L. naticoides, L. kasatka and L. subrotundata. Using allozyme electrophoresis it has been demonstrated that, in the Kurile Islands, three of these species (L. sitkana, L. subrotundata and L. kasatka) co-occur, together with a fourth, still undescribed species (L. sp.). These four species were compared at 16 loci coding for 13 enzymes. All species were easily distinguished by diagnostic enzyme markers. The mean genetic distances and ranges between species pairs are: L. sitkana and L. sp. D = 0.622 (0.561–0.741), L. sitkana and L. subrotundata D=0.981 (0.821–1.110), L. subrotundata and L. sp. D=0.975 (0.955–0.995). The genetic distance between L. kasatka and each of the other three species was greater than 1 (range 1.123–2.087). These data suggest that L. sitkana, L. subrotundata and L. sp could be members of a species complex; according to current classifications these three belong to the subgenus Neritrema. However, the genetic distance between L. kasatka and L. sitkana is much greater than between L. sitkana and other Neritrema species, and thus supports the classification of L. kasatka in the subgenus Littorina.     

Local site differences in survival and parasitism of periwinkles (Littorina sitkana Philippi, 1846)

The periwinkle, Littorina sitkana, is found throughout the intertidal zone, often in isolated subpopulations. The majority of trematode parasites use snails as intermediate hosts, and decreased survivorship is often observed in snails infected with trematodes. Sampling L. sitkana from four sites in Barkley Sound, British Columbia, Canada, we test the effects of parasitic infection on snail survival using maximum likelihood and Bayesian approaches using the software MARK and WinBUGS. We found that survival of periwinkles and trematode community composition differed among sites, but survival and trematode prevalence were uncorrelated. WinBUGS performed better than MARK in two ways: (1) by allowing the use of information on known mortality, thus preventing survival overestimation; and (2) by giving more stable estimates while testing the effect of body size on snail survival. Our results suggest that snail survival depends heavily on local environmental factors that may vary greatly within a small geographical region. These findings are important because the majority of experimental studies on survival are done on snails from a single location.     

Life history of Littorina scutulata and L. plena, sibling gastropod species with planktotrophic larvae

Abstract. The intertidal, sibling species Littorina scutulata and L. plena (Gastropoda, Proso‐branchia) are sympatric throughout most of their ranges along the Pacific coast of North America. Both species release disc‐shaped, planktonic egg capsules from which planktotrophic veliger larvae hatch. Here I review existing data and present new observations on these species' life history, including age at first reproduction, spawning season, maximum fecundity rates, capsule morphology, egg size and number, pre‐hatching development, larval growth at three food concentrations, potential settlement cues, planktonic period, and protoconch size. Previous classification of egg capsule morphologies used to distinguish the species is inaccurate; instead, capsules can be categorized into three types of which each species may produce two. Females of L. scutulata produced capsules with either two rims of unequal diameter or one rim, while females of L. plena produced capsules with one rim or two rims of nearly equal diameter. Females of each species spawned sporadically from early spring to early fall in Puget Sound. Larvae of L. plena hatched one day earlier than those of L. scutulata, and both species grew fastest in the laboratory at intermediate food concentrations. Larvae metamorphosed in the presence of a variety of materials collected from their adult habitat, including conspecific adults, algae, rocks, and barnacle tests. This is the first report of planktotrophic larvae in this genus metamorphosing in the laboratory. The total planktonic period of 8 larvae of L. scutulata raised in the laboratory was 37–70 days, and a single larva of L. plena metamorphosed after 62 days. Protoconch diameter of shells collected from the field was 256–436 μm and did not differ significantly between the species. Previous allozyme and mitochondrial DNA work has suggested high levels of genetic variability in both species and greater genetic population structure in L. plena, despite the long spawning season and long‐lived larvae in both species. The interspecific life history differences described here appear insufficient to produce consistent differences in gene flow patterns.     

Patterns of growth and energy allocation in northern California populations of Littorina (Gastropoda: Prosobranchia)

Seasonal patterns in shell growth rates, tissue weights, and female reproductive efforts were monitored for three species of western North American rocky shore gastropods, Littorina keenae Rosewater (supralittoral), L. scutulata Gould (high intertidal), and L. plena Gould (high intertidal). Growth rates conformed to the Von Bertalanffy growth model. Relative to male snails, female L. keenae appeared to sacrifice shell growth to expend more energy on the production of large quantities of gametes during the spring; growth rates of females and large males were highest during autumn when microalgal food supplies were seasonally abundant. In contrast, spring tended to be the best season for shell growth of L. scutulata and L. plena; however, reproductive output was poor and sporadic for these two species, and growth rates in the field during all seasons were low compared to growth rates of snails held under laboratory conditions with abundant food. High population densities of snails at shore levels occupied by L. scutulata and L. plena may greatly reduce food levels upon which individual snails rely for growth and reproduction.     

Mechanisms of species coexistence: a field test of theoretical models using intertidal snails

Competitor coexistence is often facilitated by spatial segregation. Traditionally, spatial segregation is predicted to occur when species differ in the habitat in which they are either superior at competing for resources or less susceptible to predation. However, predictions from a behavioural model demonstrate that spatial segregation and coexistence can also occur in the absence of such interspecific trade‐offs in competitive ability and vulnerability to predation. Unlike other models of competitor coexistence this model predicts that when species rank both habitat productivity and ‘riskinesses’ similarly, but differ slightly in their habitat‐specific vulnerabilities to predators, they will tend to segregate across habitats, with the species experiencing the higher ratio of mortality risk across the habitats occurring primarily in the safer habitat. Here, we investigate the hypothesis that intraspecific trade‐offs between resource availability and mortality risk can lead to spatial segregation of competing species by (1) documenting the spatial (i.e. intertidal) distribution of two marine snails, Littorina sitkana and L. subrotundata and (2) performing field experiments to quantify growth and mortality rates of each species at ‘low’ and ‘high’ intertidal heights. Our results indicate that both species agree on the rankings of habitat riskiness and productivity, experiencing higher predation and higher growth in low‐ than in high‐intertidal habitats. However, L. sitkana and L. subrotundata experienced differences in their habitat‐specific mortality risks and growth rates. Despite both species being similarly at risk of predation in high‐intertidal habitats (where mortality was lower), L. subrotundata was subject to significantly higher mortality than L. sitkana at the low‐intertidal height. In contrast, growth rate differences between habitats were greater for L. sitkana than for L. subrotundata. Whereas both species grew at the same rate at the high‐intertidal level (where growth was lower), L. sitkana individuals grew more rapidly than L. subrotundata snails at the low‐intertidal level. As predicted by the behavioural model, the species that experienced the higher ratio of mortality across habitats (i.e. L. subrotundata) occurred exclusively in the safer, high‐intertidal habitat. Taken together, these results provide support for the hypothesis that spatial segregation, and potentially competitor coexistence, can occur in the absence of interspecific trade‐offs in resource acquisition ability or vulnerability to predation.     

Microsatellite Loci from Lanyu Scops Owl (Otus elegans botelensis) and their Cross-Species Application in Four Species of Strigidae

We developed six new microsatellite markers containing tetranucleotide repeat motifs (GATA/CTAT) for Lanyu scops owl (Otus elegans botelensis) from an enriched partial library. All these loci are polymorphic and conform to Hardy–Weinberg equilibrium. We cross-species tested these and 12 other microsatellite primer pairs previously developed from O. elegans on four other species of owls (O. lettia, O. spilocephalus, O. scops, and Ninox scutulata). Results showed that the degree of polymorphism decreased with increasing phylogenetic distance to O. elegans. Most loci (66.7, 83.3, and 100%) were polymorphic in the three Otus owls but only five (27.8%) were polymorphic in N. scutulata. These microsatellites should be very useful genetic markers in studying the mating system, population genetics, and conservation of other little studied Old World Otus owls.     

Macrogeographic genetic variability in the Gastropod Mollusk <Emphasis Type="Italic">Littorina sitkana</Emphasis> from the Northwest Pacific

Variation at four highly polymorphic allozyme loci (inorganic pyrophosphatase, peptidase, and two esterase loci) was examined in 25 settlements of the marine snail Littorina sitkana (Mollusca, Gastropoda). The sampling localities covered a wide part of the species range: from the Peter the Great Bay (the Sea of Japan) at the southwest to the Mednyi Island (Commander Islands) at the northeast. Like other littorines lacking the pelagic stage, L. sitkana was characterized by significant genetic differentiation (G _ST for the pooled sample was 0.310). Cluster analysis and nonmetric multidimensional scaling conducted on a matrix of pairwise genetic distances between all of the settlements studied revealed four genetically different groups: southern Primorye, northern Prymorye, Sakhalin, and Kuril-Commanders. The population-genetic structure of the L. sitkana settlements is similar to that described by the isolation-by-distance and stepping-stone models: the geographic and the genetic distances between the most settlements examined are distinctly correlated.Translated from Genetika, Vol. 41, No. 3, 2005, pp. 374–384.Original Russian Text Copyright © 2005 by Zaslavskaya, Pudovkin.     

Fine structural observations of sperm resorption in the seminal vesicle of a marine snail,Littorina scutulata (Gould, 1849)

Summary Waste sperm and spermatozeugmata in the seminal vesicle of Littorina scutulata are phagocytised either by cell buds (large vesicles given off from the epithelial cells) or by the epithelial cells themselves. Cell buds containing sperm, are in turn engulfed by epithelial cells. In both cases, heterophagic vacuoles are formed inside the cell and subsequently the vacuoles are fused with primary lysosomes or lysosomal derivatives to become secondary lysosomes. Throughout this process the sperm are being digested. The second lysosome transforms further to telolysosome and finally to residual body when the sperm is completely digested.     

Phylogeography of <Emphasis Type="Italic">Littorina sitkana</Emphasis> in the northwestern Pacific Ocean: evidence of eastward trans-Pacific colonization after the Last Glacial Maximum

We investigated genetic diversity and population structure of the Sitka periwinkle Littorina sitkana along the coastlines of the northwestern Pacific (NWP) to evaluate the possibility of trans-Pacific colonization of this species from the NWP to the northeastern Pacific (NEP) after the Last Glacial Maximum. We sampled L. sitkana from 32 populations in the NWP, and sequenced a region of the mitochondrial cytochrome b oxidase gene for population genetic analyses. The results were compared with those of previous reports from the NEP. The genetic diversity of L. sitkana was much higher in the NWP than in the NEP. Genetic connectivity between the NWP and NEP populations was indicated by an extremely abundant haplotype in the NEP that was also present in eastern Hokkaido and the Kuril Islands. To confirm these results, we compared sequences of the longest intron of the aminopeptidase N gene (APN54) in the nuclear genome in four populations of L. sitkana in the NWP with previous results from the NEP. Again, much higher genetic diversity was found in the NWP than in the NEP and genetic connectivity was supported between the Kuril Islands and the NEP. These results imply postglacial colonization of this species from the NWP to the NEP, probably along the Kuril and Aleutian Island chains. This study is the first report of possible trans-Pacific postglacial colonization of a direct-developing gastropod, inferred from genetic data.     

ALLOZYME COMPARISON OF LITTORINA SPECIES FROM THE NORTHWESTERN PACIFIC

Three unknown Littorina species were revealed by allozyme studyof snails from the northwestern Pacific. One of them was foundin Kasatka Bay (Pacific coast of Iturup Island, south KurileIslands) together with L. sitkana, which it resembled in anatomicalfeatures, but the genetic distance between these species wasD=0.622. The second one was found in Tauiskaya Inlet (northernpart of Okhotsk Sea) and was again similar to L. sitkana inmorphological and anatomical features, but D=1.160. The thirdspecies was found in Egvekinot Inlet (Anadyrskiy Gulf, BeringSea) in sympatry with L. natica, to which was close genetically(D=0.256) although it differed in shell shape. (Received 4 April 2005; accepted 26 August 2005)     

Separation of Leydigia louisi Jenkin, 1934 from L. leydigi (Schoedler, 1863) (Chydoridae, Anomopoda, Cladocera)

The aim of the present article was to contribute to the systematics of the leydigi-like species of Leydigia consisting of a few (probably 3) formal species with: (1) a large basal spine on the postabdominal claw (as long as claw thickness at the base); (2) a short setulation at anterior margin of labral keel; (3) no longitudinal striation on the female valves; (4) at least three lateral setae in each fascicle on the postabdomen; (5) three large lateral setae on exopodite III. The morphology of L. leydigi (Schoedler, 1863) and L. louisi Jenkin, 1934 are redescribed, and type material of L. macrodonta Sars, 1916 is studied. In contrast to previous suggestions (Jenkin, 1934; Smirnov, 1971), I found that: (1) L. louisi is a valid species, not a subspecies of L. macrodonta; (2) L. macrodonta is not a member of the L. leydigi-group. The third member of leydigi-group, Leydigia macrodonta longiseta Chen Shou-zhong, 1992, was described from China. It is not a subspecies of L. macrodonta, but a relative of L. leydigi; most probably, it is a valid species, but this opinion must to be confirmed by examination of original Asian material. In this article, the presence of L. leydigi in Palearctic only, and that of L. louisi in only Africa was confirmed. A new subspecies of L. louisi, found in Mexico, will be reported separately.     

Invasion by Limnoperna fortunei (Dunker, 1857) (Bivalvia, Mytilidae) of the Pantanal Wetland, Brazil

Limnoperna fortunei (Bivalvia, Mitylidae) was introduced into South America in 1991 in the La Plata River (Argentina). It arrived in the ballast water of ships coming from Asia, where this species is native. It was first observed in 1998 in the Paraguay River. Limnoperna was introduced into the Pantanal region as hull fouling of vessels using the Paraguay–Parana waterway. This study describes how L. fortunei came to the Pantanal region, and provides details of its occurrence, density, and impacts. From 1999 to 2002, observations and sampling on natural and artificial substrates in the Paraguay River were made. Some aspects of the spread and impacts, based on local community information, were also analyzed. On artificial substrate the density reached 523.8 individuals m⁻² and on natural substrate (rocks), up to 10,000 individuals m⁻² were found. The densities observed were quite low compared to those found in Southern Brazil, where values up to 100,000 individuals m⁻² have been recorded in the last 3 years. In the Paraguay River, the population density of L. fortunei can be negatively impacted by periodic low levels of dissolved oxygen and decreases in pH to between 5 and 6. Such conditions are frequently present during the periodic flooding or inundation of this area. Under these conditions, a high mortality of L. fortunei was recorded in March of 2002, on both natural and artificial substrates. Despite low densities, L. fortunei can colonize water cooling systems of boats, obstructing water circulation and causing motor overheating. Accumulation in water supply equipment, such as pumps and pipes has also been observed. An erratum to this article is available at .     

Labiobaetis species of Japan, Taiwan, and Korea, with a new synonym of L. atrebatinus (Eaton 1870) and reerection of the subspecies L. atrebatinus orientalis (Kluge 1983) (Ephemeroptera, Baetidae)

We associated nymphs of Labiobaetis sp. G and Labiobaetis sp. Q from Japan with imagoes reared from nymphs in the field. Labiobaetis sp. G was identified with L. atrebatinus (Eaton 1870) based on characters of the reared male and female imagoes, nymphs, and eggs. We also synonymized a Taiwanese species, L. morus (Chang and Yang 1994), with L. atrebatinus. After further examination of the characters of male imagoes from Japan and Korea and nymphs from Japan and Taiwan, we found them to be correspondent to subspecies L. atrebatinus orientalis (Kluge 1983). Thus, we reerected the subspecific status of L. a. orientalis, although it had been considered not distinguishable from the nominotypical subspecies L. a. atrebatinus. Labiobaetis a. orientalis is distributed in the Russian Far East, Japan, Korea, and Taiwan. We identified Labiobaetis sp. Q with L. tricolor (Tshernova 1928) based on characters of the reared male and female imagoes, nymphs, and eggs. Labiobaetis tricolor was recorded from Japan for the first time.     

A new species of the genus Laboulbenia (Laboulbeniales) on Ophionea indica (Coleoptera, Carabidae, Odacanthini) from Taiwan

Laboulbenia ophioneae is described as a new species with illustrations. It is closely related to Laboulbenia celestialis and Laboulbenia asiatica because of some similarities in the morphology of appendages and perithecia. The present species from the latter two species can be distinguished by the shorter, inflated perithecia, the shorter receptacles, and the appendages consisting of more or less darkly colored, broader branches. This new species was found on elytra of Ophionea indica from Taiwan, which has been also known as a host of Laboulbenia polymorpha. A mature thallus noticed by Terada (2004) on the slide 673b (M. Ishikawa collection) and a young thallus illustrated by Sugiyama (1978, fig. 1-G as L. polymorpha) on 673d (M. Ishikawa collection) are both identified as L. ophioneae.     

Phylogeny and bionomics of Lasius austriacus (Hymenoptera,Formicidae)

Summary Phylogenetic analysis based on sequence data of the mitochondrial COI gene confirms the species status of the recently described Lasius austriacus. The five haplotypes of L. austriacus do not cluster according to their geographic origin, indicating a recent gene flow among the populations. The molecular data corroborate the morphology based hypothesis that L. austriacus belongs to the Lasius (Lasius s.str.) brunneus group. The invasive species Lasius neglectus forms a sister taxon with L. turcicus, both next related to L. austriacus. Other phylogenetic relationships within the genus Lasius are in accordance with morphological data.First data on the bionomics of L. austriacus are discussed in context with its phylogenetic position. Based on gyne and male morphology, excavations of nests, pitfall trapping and observations in formicaries, we hypothesize that L. austriacus is a mainly hypogaeic, monogynous species with nuptial flight. These are characters of the Lasius brunneus group in general, except the polygynous-polycalic, intranidally copulating L. neglectus. Aggression tests, however, revealed non-aggressive behaviour (antennation) between separated L. austriacus populations, but pronounced interspecific aggression against L. neglectus. This confirms the species status of L. austriacus and indicates a reduced level of intraspecific aggression, similar to L. neglectus.The status of L. austriacus as a native species in Central Europe is confirmed.Received 10 February 2003; revised 24 June 2003; accepted 23 July 2003.     

Culture-independent analysis for determination of yeast diversity during solid substrate fermentation of grated cassava for gari production

Yeast diversity during fermentation of grated cassava for gari production in Nigeria was studied using culture independent methods based on Eukarya18S rDNA and the PCR-Denaturing Gradient Gel Electrophoresis (PCR-DGGE). DNA extracted directly from the grated cassava at different fermentation time regimes, as well as bulk cells generated from different viable count agar plates were used as template for amplification of 18S rDNA gene in a PCR experiment. Analysis of the 18S rDNA gene by sequencing of the PCR-DGGE band fragments revealed closest relative of Issatchenkia scutulata, Candida rugopelliculosa, Candida maritime, Zygosaccharomyces rouxii and Galactomyces geotricum as member of yeast community involved with the fermentation. PCR-DGGE can be useful for yeast in situ profiling during fermentation; this will contribute to safety of traditional fermented foods and optimization during large scale production of gari.