Invasional meltdown: invader-invader mutualism facilitates a secondary invasion

In multiply invaded ecosystems, introduced species should interact with each other as well as with native species. Invader-invader interactions may affect the success of further invaders by altering attributes of recipient communities and propagule pressure. The invasional meltdown hypothesis (IMH) posits that positive interactions among invaders initiate positive population-level feedback that intensifies impacts and promotes secondary invasions. IMH remains controversial: few studies show feedback between invaders that amplifies their effects, and none yet demonstrate facilitation of entry and spread of secondary invaders. Our results show that supercolonies of an alien ant, promoted by mutualism with introduced honeydew-secreting scale insects, permitted invasion by an exotic land snail on Christmas Island, Indian Ocean. Modeling of land snail spread over 750 sites across 135 km2 over seven years showed that the probability of land snail invasion was facilitated 253-fold in ant supercolonies but impeded in intact forest where predaceous native land crabs remained abundant. Land snail occurrence at neighboring sites, a measure of propagule pressure, also promoted land snail spread. Site comparisons and experiments revealed that ant supercolonies, by killing land crabs but not land snails, disrupted biotic resistance and provided enemy-free space. Predation pressure on land snails was lower (28.6%), survival 115 times longer, and abundance 20-fold greater in supercolonies than in intact forest. Whole-ecosystem suppression of supercolonies reversed the probability of land snail invasion by allowing recolonization of land crabs; land snails were much less likely (0.79%) to invade sites where supercolonies were suppressed than where they remained intact. Our results provide strong empirical evidence for IMH by demonstrating that mutualism between invaders reconfigures key interactions in the recipient community. This facilitates entry of secondary invaders and elevates propagule pressure, propagating their spread at the whole-ecosystem level. We show that identification and management of key facilitative interactions in invaded ecosystems can be used to reverse impacts and restore resistance to further invasions.     

Effects of Buffer-Strip Retention and Clearcutting on Land Snails in Boreal Riparian Forests

Abstract:  We investigated the short-term effects of forest clearcutting on land snails (terrestrial gastropods) in 15 forest stands along small streams in Sweden. Two different silvicultural treatments were applied at each site: clearcutting across the stream channel and buffer strips 10 m wide on each side of the stream. Additionally, we studied 10 reference sites in unlogged riparian forests along similar-sized streams. All sites were studied before logging and then 2.5 years after logging. After clearcutting the number of individuals in a 0.5-m² sample from each site decreased on average from 107 to 87, and the mean number of species per sample decreased from 9.9 to 7.7. Most species were negatively affected, but there were also clear differences in sensitivity. There were correlations between species survival and ground moisture. At the wettest clearcut sites with an almost intact bryophyte cover, the land snails were unaffected by clearcutting. This result suggests that wet or moist forest floors can serve as refugia even at very small spatial scales (e.g., shallow hollows, crevices). If this is an important mechanism, the spatial distribution of small habitats could be important for the long-term survival of the snail fauna or other small, dispersal-limited organisms at clearcut sites. In the buffer strips, the number of individuals decreased but not the number of species, indicating that buffer-strip retention is a good practice for protecting land snails in riparian forests. The varying effectiveness of the buffer strip could partly be explained by the proportion of the remaining basal area, emphasizing that buffer strips could be even more effective if efforts are made to avoid heavy damage by windthrows.     

Individual- and Assemblage-Level Effects of Anthropogenic Sedimentation on Snails in Lake Tanganyika

Abstract:  Human impacts on aquatic biodiversity are often measured at the assemblage or community level, although it has been suggested that individual-level measures are more sensitive. We evaluated the effects of anthropogenic sedimentation on endemic snails in Lake Tanganyika, East Africa, by comparing assemblage-level (i.e., species richness, evenness, and abundance) and individual-level (i.e., frequencies of predation and parasitism, fecal organic content, life history) data between sediment-disturbed and reference sites. Previous studies have indicated that sedimentation kills snails and reduces mollusc diversity in this system, but we found little evidence of changes in species richness, evenness, or snail abundance at the levels of sedimentation recorded. In contrast, individual-level data revealed a variety of differences associated with sedimentation. Frequencies of shell scarring by predatory crabs and castration by parasitic trematodes were significantly lower at disturbed sites, indicating shifts in interspecific interactions. Snails ingested large amounts of inorganic sediments at disturbed sites, suggesting a reduction in food quality. In addition, sedimentation was associated with a large downward shift in size distribution within some species and reproduction at smaller size. These strong patterns in individual-level data contrast with the lack of effects at the assemblage level. We argue that incorporating individual-level measures will often enhance the sensitivity of impact surveys and may reveal effects of disturbance on important interspecific interactions.     

Among- and within-population variation in sperm quality in the simultaneously hermaphroditic land snail Arianta arbustorum

Sperm competition models on the evolution of sperm size assume associations with another sperm quality trait, sperm longevity. Sperm length can also provide an indication of possible mechanisms affecting motility and thus fertilization success. Despite their importance, however, detailed mechanisms of sperm competition at the gamete level are poorly understood. In simultaneously hermaphroditic land snails, sperm traits and cryptic female choice are assumed to be crucial in determining fertilization success. We examined the variation in sperm length and number among individuals from four natural populations of the land snail Arianta arbustorum, a species with multiple mating and long-term sperm storage. We also assessed variation in velocity, motility and longevity of sperm in snails from two of the four populations. Independent of shell size, sperm length differed among populations and, to a minor extent, even among individuals within populations. Mean sperm length of a snail was not correlated with the number of sperm delivered in a spermatophore. The mean sperm velocity (=VCL) did not differ between snails from two populations. However, VCL varied among snails. Percentage motility and longevity of sperm differed between snails from the two populations. No correlations were found between length, velocity, percentage motility and longevity of sperm. To conclude, individual snails differed in sperm quality, and this variation may partly explain the differential fertilization success between A. arbustorum snails. Moreover, our findings did not support the positive association between sperm length and longevity assumed by sperm competition models for internally fertilizing species.     

The chemosensory ability of the predatory leech Whitmania laevis (Arhynchobdellida: Haemopidae) for prey searching

Although prey-detecting and searching abilities of predatory leeches of rhynchobdellid or the Erpobdelliformes of arhynchobdellid species have been studied in the past, hirudiniformes leeches are rarely mentioned. In this study, we investigated the chemosensory ability for prey-detecting and searching in Whitmania laevis, a hirudiniformes species that mainly preys on freshwater snails, and examined if such ability aided in their prey selection. Five sympatric snail species, i.e., apple snail Pomacea canaliculata, thiarid snail Thiara tuberculata, viviparid snail Sinotaia quadrata, ear pond snail Radix auricularia swinhoei and tadpole snail Physa acuta were used as prey. Our results showed that W. laevis has the chemosensory ability to detect the waterborne odors of snails. However, they follow the snails by their mucus trails, and not by the odor that the snails leave in the water. Of these five snail species, W. laevis only followed the trails of the thiarid snails, ear pond snails and tadpole snails, and did not show a different response to the trails produced by snails of different sizes. Our results suggest that W. laevis can use waterborne odors to detect the existence of prey. They rely on mucus trails to follow their preferred prey, but do not distinguish between snails of a preferred size by their mucus trails. In addition, when following the trail of a preferred snail, W. laevis exhibits a newly described searching behavior, i.e., head tapping, and may use it to locate a snail trail and increase its probability of finding the trail-laying snail nearby.     

Maximum Plant-Community Endemism at Intermediate Intensities of Anthropogenic Disturbance in Bolivian Montane Forests

Abstract: I compared the endemism of four plant groups (Araceae, Bromeliaceae, Palmae, Pteridophyta) along gradients of increasing anthropogenic forest disturbance, from undisturbed mature forest to disturbed forest (logged, grazed, or burned), secondary forest, secondary scrub, and finally pasture, at 16 sites in the Bolivian Andes. I measured endemism as the mean inverse range size (number of 1° cells) of all species per study group encountered in each habitat and site. Overall, endemism was significantly higher in disturbed forest than in mature forest, but it declined in more strongly disturbed habitats. To explain the relationship of range size to habitat disturbance, I propose that endemic species are somewhat competitively inferior to other co-occurring taxa, limited in their ability to establish and maintain new populations following dispersal and thus to expand their ranges. Within their established ranges, endemic species depend on natural habitat disturbances to prevent their competitive exclusion by other species, so they profit from a certain level of anthropogenic disturbance. This pattern and the explanatory hypotheses should be subjected to critical evaluation. Although the pattern does not apply to every endemic tropical plant species, it indicates that conservation of part of the endemic tropical forest flora may be achieved in forest areas subject to sustainable forest use without the need to completely exclude human activities.     

Invasive Non‐Native Species’ Provision of Refugia for Endangered Native Species

Abstract: The influence of non‐native species on native ecosystems is not predicted easily when interspecific interactions are complex. Species removal can result in unexpected and undesired changes to other ecosystem components. I examined whether invasive non‐native species may both harm and provide refugia for endangered native species. The invasive non‐native plant Casuarina stricta has damaged the native flora and caused decline of the snail fauna on the Ogasawara Islands, Japan. On Anijima in 2006 and 2009, I examined endemic land snails in the genus Ogasawarana. I compared the density of live specimens and frequency of predation scars (from black rats[Rattus rattus]) on empty shells in native vegetation and Casuarina forests. The density of land snails was greater in native vegetation than in Casuarina forests in 2006. Nevertheless, radical declines in the density of land snails occurred in native vegetation since 2006 in association with increasing predation by black rats. In contrast, abundance of Ogasawarana did not decline in the Casuarina forest, where shells with predation scars from rats were rare. As a result, the density of snails was greater in the Casuarina forest than in native vegetation. Removal of Casuarina was associated with an increased proportion of shells with predation scars from rats and a decrease in the density of Ogasawarana. The thick and dense litter of Casuarina appears to provide refugia for native land snails by protecting them from predation by rats; thus, eradication of rats should precede eradication of Casuarina. Adaptive strategies, particularly those that consider the removal order of non‐native species, are crucial to minimizing the unintended effects of eradication on native species. In addition, my results suggested that in some cases a given non‐native species can be used to mitigate the impacts of other non‐native species on native species.     

Ecology of a corallivorous gastropod,<Emphasis Type="Italic"> Coralliophila abbreviata</Emphasis>, on two scleractinian hosts. I: Population structure of snails and corals

Despite their potential importance in structuring reef communities, invertebrate corallivores and their population structures are poorly understood. We found distinct differences in the population structures (length-frequency distribution and sex ratio) of the corallivorous gastropod Coralliophila abbreviata residing on two coral-host taxa, Montastraea spp. and Acropora palmata, in the Florida Keys. In each of two survey years, around 50% of the Montastraea spp. colonies were infested, with a mean snail density of eight snails per infested colony (range 1–45), while around 20% of A. palmata colonies harbored three snails per infested colony (range 1–23). Variation in patterns of snail occurrence was also observed within a host taxon. A. palmata occurred in low- and high-density stands (0.4 and 1 colony m^–2, respectively, at the initial survey) at different sites. Hurricane Georges struck the area in September 1998. When resurveyed in 1999, density of colonies in low-density stands had decreased by 75% to 0.1 colonies m^–2. This decrease was accompanied by a doubling in the proportion of colonies infested with snails (from 19% to 46%) and an increase in snail density per infested colony (from 3.7±3.3 SD to 5.4±4.6 SD) as snails apparently concentrated on surviving A. palmata. In contrast, sites with high density A. palmata stands (thickets) retained colony densities of about ~1 colony m^–2 among years, while snail infestation increased only from 9% to 14% of colonies surveyed and snail density essentially remained unchanged (from 2.7±1.8 to 2.9±1.9 snails per infested colony). Snails collected from Montastraea spp. were shorter than those from A. palmata in low-density stands and were longest on A. palmata in thickets. On both host taxa, female snails were longer than males. The sex ratio of snails on Montastraea spp. hosts was even (1:1), while that of snails on A. palmata was skewed (70% males). Factors that could explain observed differences in size structure and sex ratio between Coralliophila populations on the two coral host taxa include: differential susceptibility to predators, influence of host tissue nutritional quality and/or secondary metabolite content, and genetic differences (cryptic species). The host-specific characteristics of C. abbreviata populations imply that the impact of gastropods on reef communities will vary with the coral species composition.     

Morphologie,Entwicklung und Ökologie digener Trematoden-Larven der StrandschneckeLittorina littorea

More than 6000 individuals of the snailLittorina littorea L., from 6 different localities on the coast of the German Bay (North Sea), were examined for larval trematodes by crushing their shells, an additional 6000 by isolation in small quantities of sea water. Six species of larval trematodes were found inL. littorea and described: Notocotylid larva —Cercaria lebouri Stunkard, 1932; Cercaria ofHimasthla elongata (Mehlis, 1831); Cercaria ofPodocotyle atomon (Rudolphi, 1802); Cercaria ofRenicola roscovita (Stunkard, 1932) n. comb.; Metacercaria ofMicrophallus pygmaeus (Levinsen, 1881); Cercaria ofCryptocotyle lingua (Creplin, 1825). Considering the large number of investigated snails, the trematode fauna ofL. littorea in this area is assumed to be completely known. The life cycles ofHimasthla elongata andRenicola roscovita are described. Larvae of the different trematode species show differential preferences for distinct size groups of snail hosts. Juvenile snails are not infected. No correlation exists between infection rates and sex of the hosts. Quantitative aspects of multiple infections are calculated and discussed. The incidence of larval trematodes in the investigated localities differs. Information is presented on seasonal variations of infestations and their possible causes.     

Host preference of a symbiotic flatworm in relation to the ecology of littoral snails

Some polyclads associate with other marine invertebrates such as gastropods, but how these associations evolve is not well understood. This study examined the relationships between a polyclad, Stylochoplana pusilla, and six species of snails in 10 eulittoral sites in Mutsu Bay, northern Japan, in 2010 and 2011. Of the six potential host snail species, Monodonta labio was present at all sites and the prevalence of S. pusilla in the snails’ mantle cavities overall was 82.6 %. Four other snail species occurred at five or six sites in comparable densities, but of these, only Chlorostoma turbinatum and Omphalius rusticus had S. pusilla in their mantle cavities (prevalence of 42.0 and 15.4 %). A congener of M. labio, Monodonta neritoides, occurred at only one site and had no S. pusilla. The mantle, which provides habitat space for S. pusilla, was proportionally the longest in M. labio. In laboratory experiments, the flatworms preferentially entered M. labio over other snail species with the same mantle length. Observed vertical distributions of snails in the littoral zone indicate that M. labio is present in the eulittoral at both high and low tides, while C. turbinatum and O. rusticus are usually found in the sublittoral. In an evolutionary sense, the relationship between S. pusilla and its host may have developed as a mechanism for the flatworm to avoid predation by living in the eulittoral and to avoid desiccation by establishing commensal relationships with certain snails.     

Effects of Dam‐Induced Landscape Fragmentation on Amazonian Ant–Plant Mutualistic Networks

Mutualistic networks are critical to biological diversity maintenance; however, their structures and functionality may be threatened by a swiftly changing world. In the Amazon, the increasing number of dams poses a large threat to biological diversity because they greatly alter and fragment the surrounding landscape. Tight coevolutionary interactions typical of tropical forests, such as the ant–myrmecophyte mutualism, where the myrmecophyte plants provide domatia nesting space to their symbiotic ants, may be jeopardized by the landscape changes caused by dams. We analyzed 31 ant–myrmecophyte mutualistic networks in undisturbed and disturbed sites surrounding Balbina, the largest Central Amazonian dam. We tested how ant–myrmecophyte networks differ among dam‐induced islands, lake edges, and undisturbed forests in terms of species richness, composition, structure, and robustness (number of species remaining in the network after partner extinctions). We also tested how landscape configuration in terms of area, isolation, shape, and neighborhood alters the structure of the ant–myrmecophyte networks on islands. Ant–myrmecophytic networks were highly compartmentalized in undisturbed forests, and the compartments had few strongly connected mutualistic partners. In contrast, networks at lake edges and on islands were not compartmentalized and were negatively affected by island area and isolation in terms of species richness, density, and composition. Habitat loss and fragmentation led to coextinction cascades that contributed to the elimination of entire ant–plant compartments. Furthermore, many myrmecophytic plants in disturbed sites lost their mutualistic ant partners or were colonized by opportunistic, nonspecialized ants. Robustness of ant–myrmecophyte networks on islands was lower than robustness near lake edges and in undisturbed forest and was particularly susceptible to the extinction of plants. Beyond the immediate habitat loss caused by the building of large dams in Amazonia, persistent edge effects and habitat fragmentation associated with dams had large negative effects on animal–plant mutualistic networks. Efectos de la Fragmentación del Paisaje Inducida por Presas sobre Redes Mutualistas Hormiga‐Planta Amazónicas     

Purification of multiple vitellogenins in grey mullet (<Emphasis Type="Italic">Mugil cephalus</Emphasis>)

Coralliophila abbreviata (Lamarck) is a corallivorous gastropod that lives and feeds on several species of scleractinian coral in the Western Atlantic and Caribbean. Previous studies of C. abbreviata have revealed that snails on branching acroporid corals are larger and consume more tissue than those on massive and plating corals. To ascertain whether snail life-history and fitness are differentially affected by the coral host, an analysis of the age structure and female reproductive output of snail populations on three coral host taxa (Acropora palmata, Diploria spp., and Montastraea spp.) was conducted at four shallow (2–7 m depth) reef sites off Key Largo, Florida in June through August, 2004. Snails were, on average, almost twice as large on A. palmata than on Diploria spp. and Montastraea spp., averaging 30.3 mm shell length, compared to 17.2 and 17.6 mm, respectively. Brood size increased as a power function with female shell length. Females on A. palmata were significantly larger than females on the other two hosts and, therefore, produced more offspring per female. The number of growth striae on the inner surface of the operculum was used to estimate snail age. Estimates of growth rate were obtained by fitting the Gompertz growth function to size-at-age plots and mortality was estimated using growth parameters and size-frequency data. The data suggest that C. abbreviata inhabiting A. palmata are larger than on alternative hosts due to a combination of a faster growth rate and longer life-span. The species is believed to be a protandrous hermaphrodite. The timing of sex change varied among hosts; snails on A. palmata changed sex later at larger sizes relative to those on the other two hosts. Based on these results, it seems probable that C. abbreviata has developed reaction norms for life-history traits, allowing snails to adjust and maximize fitness in the different environments associated with various coral hosts.     

Species richness patterns along environmental gradients in island land molluscan fauna

The relationship between species richness and environmental variables may change depending on habitat structure, dispersal ability, species mixing, and community adaptation to the environment. It is crucial to know how these factors regulate the environment-diversity relationship. The land molluscan fauna of the Ogasawara Islands in the West Pacific is an excellent model system to address this question because of the high species endemicity (> 90%), small area, and simple habitat structure of the islands. I examined relationships among indigenous species composition, richness, and habitat condition, and especially productivity and forest moisture on the island of Anijima. Two major communities of snails could be distinguished by detrended correspondence analysis (DCA): one group dominated in a moist habitat with high productivity, and the other group dominated in a dry habitat with low productivity. However, species richness became highest at the intermediate condition between the habitats in which the two snail communities were dominant, so that species richness showed a hump-shaped relationship with moisture and productivity. In contrast, the species richness of the snail community in the moist habitat showed a monotonically positive correlation, and that in the dry habitat showed a monotonically negative correlation with moisture and productivity. Thus, the greater species richness in intermediate moisture and productivity resulted from the ecotone effect or community overlap at the transitional areas, where faunas with different ecologies can meet in a single site. These findings suggest that hump-shaped productivity-diversity relationships in land Mollusca would reflect the ecotone effect as a result of the mixing of species adapted to either fertile habitats or sterile habitats.     

Road Facilitation of Trematode Infections in Snails of Northern Alaska

Abstract:  Road disturbances can influence wildlife health by spreading disease agents and hosts or by generating environmental conditions that sustain these agent and host populations. I evaluated field patterns of trematode infections in snails inhabiting ponds at varying distances from the Dalton Highway, a wilderness road that intersects northern Alaska. I also assessed the relationships between trematode infections and snail densities and six environmental variables: calcium concentration, aquatic vegetative cover, canopy cover, temperature, pond size, and community structure. Presence of trematode infections and snail density were negatively correlated with distance from the highway. Of the pond characteristics measured, only calcium concentration and vegetation density declined with distance from road. However, neither variable was positively associated with snail density or trematode presence. One potential explanation for observed patterns is that vehicles, road maintenance, or vertebrate vectors attracted to the highway facilitate colonization of snails or trematodes. Emerging disease threats to biological diversity in northern ecosystems highlight the importance of understanding how roads affect disease transmission.     

Human disturbance and shifts in vertebrate community composition in a biodiversity hotspot

Understanding how human modification of the landscape shapes vertebrate community composition is vital to understanding the current status and future trajectory of wildlife. Using a participatory approach, we deployed the largest camera-trap network in Mesoamerica to date to investigate how anthropogenic disturbance shapes the occupancy and co-occurrence of terrestrial vertebrate species in a tropical biodiversity hotspot: the Osa Peninsula, Costa Rica. We estimated species richness in different categories of land protection with rarefaction analysis and estimated the expected occupancy with a joint species distribution model that included covariates for anthropogenic disturbance, land protection, habitat quality, and habitat availability. Areas with the most stringent land-use protections (e.g., Corcovado National Park, 24 species [95% CI 23–25]) harbored significantly more species than unprotected areas (20 species [19.7–20.3]), mainly due to a reduced presence of large-bodied species of conservation concern in unprotected areas (e.g., jaguar Panthera onca and white-lipped peccary Tayassu pecari). Small-bodied generalist species, such as opossums (Didelphidae) and armadillos (Dasypus novemcinctus), in contrast, were more common at disturbed sites, resulting in a significant difference in vertebrate community composition between sites with low and high disturbance. Co-occurrence of species was also mainly associated with response to disturbance. Similar responses to disturbance create two groups of species, those whose site-level occupancy usually increased as anthropogenic disturbance increased and those whose estimated occupancy decreased. The absence of large-bodied species entails an important loss of ecological function in disturbed areas and can hinder forest development and maintenance. Efforts to protect and restore forested landscapes are likely having a positive effect on the abundance of some threatened species. These efforts, however, must be sustained and expanded to increase connectivity and ensure the long-term viability of the wildlife community.     

Utilization of partially predated snail shells by the hermit crab Pagurus longicarpus Say, 1817

The hermit crab Pagurus longicarpus was shown to inhabit shells that were partially predated from intertidal areas of Long Island, New York. Among field collections of P. longicarpus, 2.13% of the hermit crabs (46 of 2155) were found with shells with snail tissue present. Over 90% of these partially predated snail shells were occupied by male hermit crabs. Although hermit crabs were in 8 species of snail shells, only Littorina littorea and Nassarius obsoletus were found occupied by hermit crabs and containing snail tissue. In the laboratory, we found that specimens of the spider crab Libinia emarginata were able to pull off the operculum of snails, leaving damage as found in field collections. In contrast, specimens of P. longicarpus were not able to prey on live, healthy snails. When specimens of P. longicarpus were placed in communal tanks, hermit crabs preferred partially predated snail shells to empty and original shells. However, original shells and empty shells were occupied with more frequency than partially predated shells when crabs were isolated. These findings indicate P. longicarpus actively seeks shells soon after attack and abandonment by snail predators, especially in the presence of competitors.     

Effects of Polinices duplicatus (Gastropoda: Naticidae) on infaunal community structure at Barnstable Harbor,Massachusetts, USA

Effects of the predatory naticid snail Polinices duplicatus (Say) on species composition, diversity and density, of the infauna of intertidal sand-flats at Barnstable Harbor, Massachusetts, USA, were studied using field experiments. Responses of molluscs (prey of P. duplicatus) and nonmolluscs (nonprey) were considered separately to distinguish between the effects of feeding and sediment disturbance during foraging. The fauna of 0.25 m² predator-exclusion cages, coarse-mesh cages, sieved areas, and controls was followed for 1 yr. Species associations within cages from which predators were excluded were denser, more diverse, and richer in molluscs than those in other treatments. Larger areas (3×3 m) of natural bottom were fenced and maintained as snail exclosures and enclosures during two feeding seasons. For both molluscs and nonmolluscs, diversity (H'), number of species, evenness (SD), and density all decreased with increasing snail density. Intense predation pressure on molluscs and selective feeding on thin-shelled bivalves, which were rare, removed individuals and species of molluscs from the community. Comparison of samples taken inside and outside trails made by snails showed that disturbance of the surface sediment layers by snails decreased the abundance of spionid polychaetes and total nonmolluscs. The sipunculan Phascolopsis gouldi and the bivalve Gemma gemma dominated the community in all experimental treatments and were little affected by the activity of Polinices duplicatus. Predation and disturbance by snails lowered community diversity by removing individuals of the less abundant species, and generally maintained population densities below the level where strong competition would occur.     

Asymmetric competition via induced resistance: specialist herbivores indirectly suppress generalist preference and populations

Species may compete indirectly by altering the traits of a shared resource. For example, herbivore-induced responses in plants may make plants more resistant or susceptible to additional herbivorous insect species. Herbivore-induced plant responses can significantly affect interspecific competition and herbivore population dynamics. These herbivore-herbivore indirect interactions have been overlooked in aquatic ecosystems where previous studies used the same herbivore species to induce changes and to assess the effects of these changes. We asked whether seaweed grazing by one of two herbivorous, congeneric snail species (Littorina obtusata or Littorina littorea) with different feeding strategies and preferences would affect subsequent feeding preferences of three herbivore species (both snails and the isopod Idotea baltica) and population densities of three herbivore species (both snails and a third periwinkle snail, Lacuna vincta). In addition, we measured phlorotannin concentrations to test the hypothesis that these metabolites function as induced defenses in the Phaeophyceae. Snail herbivory induced cue-specific responses in apical tissues of the seaweed Fucus vesiculosus that affected the three herbivore species similarly. When compared to ungrazed controls, direct grazing by Littorina obtusata reduced seaweed palatability by at least 52% for both snail species and the isopod species. In contrast, direct grazing by L. littorea did not decrease seaweed palatability for any herbivore, indicating herbivore-specific responses. Previous grazing by L. obtusata reduced populations of L. littorea on outplanted seaweeds by 46% but had no effect on L. obtusata populations. Phlorotannins, a potential class of inducible chemicals in brown algae, were not more concentrated in grazed seaweed tissues, suggesting that some other trait was responsible for the induced resistance. Our results indicate that marine herbivores may compete via inducible responses in shared seaweeds. These plant-mediated interactions were asymmetric with a specialist (L. obtusata) competitively superior to a generalist (L. littorea).     

Using Parasitic Trematode Larvae to Quantify an Elusive Vertebrate Host

Abstract: Digenean trematode parasites require multiple host species to complete their life cycles, and their abundance can often be strongly correlated with the abundance of their host species. Species richness and abundance of parasites in easily sampled host species may yield an accurate estimate of the species richness and abundance of other hosts in a parasite's life cycle that are difficult to survey directly. Accordingly, we investigated whether prevalence and mean abundance of trematodes could be used to estimate the abundance of one of their host species, diamondback terrapins (Malaclemys terrapin), which are difficult to sample and are designated as near threatened (by the International Union for Conservation of Nature [IUCN Red List]) along some U.S. coasts. As an adult the trematode Pleurogonius malaclemys is specific to terrapins. Its larval stages live first inside mud snails (Ilyanassa obsoleta) and are subsequently shed into the environment where they form external metacercarial cysts on hard surfaces such as snail opercula. The life cycle of P. malaclemys is completed when terrapins ingest these cysts. At 12 sites along the coast of Georgia (U.S.A.), we determined the prevalence of internal P. malaclemys larvae in mud snails (proportion of infected snails in a population) and the prevalence and mean abundance of external trematode cysts. We examined whether these data were correlated with terrapin abundance, which we estimated with mark‐recapture methods. The abundance of external cysts and salinity explained ≥59% of the variability in terrapin abundance. We suggest that dependent linkages between the life stages of multihost parasites make them reliable predictors of host species’ abundance, including hosts with abundances that are challenging to quantify directly.