An Unique Anchialine Pool in the Hawaiian Islands

The Sailor's Hat crater was artificially formed on the south coast of Kaho'olawe Island in 1965 with explosives. The explosion formed a crater about 50 m from the shoreline, which penetrates the watertable to a 5 m depth. The pool at the bottom of the crater meets the criteria of an anchialine pond because it shows tidal fluctuation, has measurable salinity, and lacks surface connections to the sea. The water chemistry of this pool is similar to the ocean except silica is elevated and salinity is slightly depressed suggesting a small groundwater influence. The fauna is dominated by waterboatmen, an endemic shrimp and tubeworm, polychaetes, amphipods, an ostracod, gastropod, solitary ectoproct. anemone, flatworm and sponge. The atyid shrimp, Halocaridina rubra, is a characteristic species of Hawaiian anchialine systems and probably colonized this 32-year old pool by active migration via the watertable. Colonization by the remaining fauna may have occurred by storm surf (for marine species) or with the wind. Most predators are unable to inhabit anchialine ponds because of difficult access due to physical barriers, or to unsuitable ecological conditions. The anchialine habitat and life history strategy of the atyid shrimp have probably been important influences on the adaptative success of H. rubra in the Hawaiian Islands, and may be important characteristics of hypogeal anchialine species elsewhere.     

Factors influencing fish distribution and community structure in a small coastal river in southwestern Costa Rica

We evaluated the relative importance of habitat size and diversity and distance from the ocean in explaining longitudinal patterns of fish distribution and community structure (species richness, evenness, and diversity) in the lower 5 km of the Rio Claro. The Rio Claro is a small coastal river in Corcovado National Park, southwestern Costa Rica, with a depauperate freshwater fish fauna. We observed 22 species in pools, 19 of which occured during quantitative sampling. Most of these species probably spent part of their lives in the Pacific Ocean. We observed no species in riffles, although these habitats were common and several taxa (e.g., Gobiesocidae, Gobiidae) were adapted for life in fast turbulent water. Fish abundance, and species richness, evenness, and diversity were highest near the ocean, where high tides influenced river levels and salinity. Eight species were not observed further than 2900 m from the ocean, whereas 6 species were not encountered in samples nearest the ocean; the remaining 8 species were present throughout the study area. Distance from the ocean was a better predictor of fish abundance and community structure than were pool width, pool depth, pool surface area, pool volume, amount of cover present, substrate diversity, or depth diversity. Based in qualitative comparisons of our data with previously published data from 1980–1986, the fish assemblage of the Rio Claro is persistent in both species composition and relative abundance.     

An island in the stream: Australia's place in the cosmopolitan world of Indo-West Pacific freshwater shrimp (Decapoda: Atyidae: Caridina)

Mitochondrial DNA sequences were used to investigate phylogenetic and biogeographic relationships among Australian freshwater shrimp from the genus Caridina H. Milne Edwards, 1837 (Atyidae) and congeners from potential source populations throughout the Indo-West Pacific region. Numerous Australian taxa have close evolutionary relationships with non-Australian taxa from locations throughout the region, indicating a diverse origin of the Australian freshwater fauna. This implies many colonisations to or from Australia over a long period, and thus highlights the surprising adeptness of freshwater shrimp in dispersal across ocean barriers and the unity of much of the region's freshwater biota. Interestingly, a study on Australia's other main genus of atyid shrimp, Paratya Miers, 1882, inferred only a single colonisation. A number of potential species radiations within Australia were also identified. This agrees with patterns detected for a large number of Australian freshwater taxa, and so implies a vicariant explanation due to the development of colder, dryer climates during the late Miocene/early Pliocene.     

Genetic exchange between anchialine cave populations by means of larval dispersal: the case of a new gastropod species Neritilia cavernicola

Despite being limited to caves, many anchialine taxa have disjunct insular distributions, which raises questions about their origins and colonization history. This study deals with the new gastropod Neritilia cavernicola sp. n. (Neritopsina: Neritiliidae) from anchialine caves on two islands in the Philippines that are separated by the deep Bohol Strait and situated 200 km apart along the coastline of Cebu Island. Neritilia cavernicola is an obligate stygobiont and most closely resembles Neritilia littoralis , which lives in interstitial waters of the Nansei-shoto Islands, Japan. Its eggs and larval shells are identical to those of other Neritilia species, despite their different adult habitats. This suggests a marine planktotrophic phase (as occurs in amphidromous riverine species of Neritilia ), and consequent migration between islands via ocean currents. Here we present the first genetic structure for anchialine cave organisms; comparisons of 1276 bp sequences from mitochondrial cytochrome oxidase I show no evidence of genetic isolation between the islands. All individuals evidently are part of a panmictic population and the low vagility of adults and their seemingly isolated cave habitats do not limit gene flow in N. cavernicola . This migration model, based on marine larval dispersal, may be widely applicable to anchialine stygobites with insular distributions, as many such organisms (including shrimps, crabs and fishes) are phylogenetically allied to amphidromous species.     

Rocks and clocks: linking geologic history and rates of genetic differentiation in anchialine organisms

The geologic history of a region can significantly impact the development of its flora and fauna, with past events shaping community patterns and evolutionary trajectories of species. In this context, islands are excellent “natural laboratories” for studying the fundamental processes of evolution due to their discrete geographical nature and dynamic geologic histories. An island system meeting these criteria is the Hawaiian Archipelago, which is ideal for testing how island geologic history influences the processes leading to population genetic variation and differentiation. One Hawaiian endemic whose evolutionary history is closely tied to the geology of the islands is the anchialine atyid shrimp Halocaridina, whose mitochondrial cytochrome oxidase I (COI) gene is hypothesized to be evolving at the rate of 20% per million years. To validate this rapid evolutionary rate, time since divergence estimates between geographically close, yet genetically distinct, populations were calculated for Halocaridina from anchialine habitats on the islands of Hawai’i, Maui, and O’ahu. On the younger (i.e., <1.5 million years) islands of Hawai’i and Maui, where all anchialine habitats occur in basalt, application of the Halocaridina molecular clock identified a strong correlation between levels of genetic divergence and the geologic age of the region inhabited by those populations. In contrast, this relationship weakened when similar analyses were conducted for Halocaridina from limestone anchialine habitats on the older (i.e., >2.75 million years) island of O’ahu. These results suggest geologic age, basin origin and/or composition are important factors that should be taken into consideration when conducting molecular clock analyses on anchialine flora and fauna as well as island populations in general.     

Responses of Euglossine Bees (Hymenoptera,Apidae, Euglossina) to an Edge-Forest Gradient in a Large Tabuleiro Forest Remnant in Eastern Brazil

Euglossine fauna of a large remnant of Brazilian Atlantic forest in eastern Brazil (Reserva Natural Vale) was assessed along an edge-forest gradient towards the interior of the fragment. To test the hypotheses that the structure of assemblages of orchid bees varies along this gradient, the following predictions were evaluated: (i) species richness is positively related to distance from the forest edge, (ii) species diversity is positively related to distance from the edge, (iii) the relative abundance of species associated with forest edge and/or open areas is inversely related to the distance from edge, and (iv) relative abundance of forest-related species is positively related to distance from the edge. A total of 2264 bees of 25 species was assessed at five distances from the edge: 0 m (the edge itself), 100 m, 500 m, 1000 m and 1500 m. Data suggested the existence of an edge-interior gradient for euglossine bees regarding species diversity and composition (considering the relative abundance of edge and forest-related species as a proxy for species composition) but not species richness.     

Patterns of Ant (Hymenoptera: Formicidae) Richness and Relative Abundance along an Aridity Gradient in Western Venezuela

In xeric ecosystems, ant diversity response to aridity varies with rainfall magnitude and gradient extension. At a local scale and with low precipitation regimes, increased aridity leads to a reduction of species richness and an increased relative abundance for some ant species. In order to test this pattern in tropical environments, ant richness and relative abundance variation were evaluated along 35 km of an aridity gradient in the Araya Peninsula, state of Sucre, Venezuela. Three sampling stations comprising five transects each were set up. Pitfall traps and direct collecting from vegetation were assessed per transect. Overall, 52 species, 23 genera, and 7 subfamilies of ants were recorded in the peninsula. The total number of species and genera recorded by both sampling stations and transects decreased linearly with increasing aridity. Total relative abundance was highest in the most arid portion of the peninsula, with Crematogaster rochai (Forel) and Camponotus conspicuus zonatus (Emery) (Hymenoptera: Formicidae) being the numerically dominant species. Spatial and multivariate analyses revealed significant changes in ant composition every 11 km of distance, and showed a decrease of ant diversity with the increase of harsh conditions in the gradient. Here, we discuss how local geographic and topographic features of Araya originate the aridity gradient and so affect the microhabitat conditions for the ant fauna.     

Void development on carbonate coasts: creation of anchialine habitats

Anchialine caves in coastal locations develop in two ways: by pseudokarst processes that form talus caves, sea caves, tafoni, fissure caves and lava tubes, and by karst dissolutional processes that form stream caves, flank margin caves, and blue holes. Pseudokarst caves are of minor importance in anchialine cave habitat development, with some lava tubes being notable exceptions. Dissolution caves provide the most extensive, variable, and long-term environments for anchialine habitats. The Carbonate Island Karst Model (CIKM) allows dissolutional cave development in carbonate coasts to be understood as the interplay between freshwater and marine water mixing, sea-level change, rock maturity, and interaction with adjacent non-carbonate rocks. Glacioeustatic sea-level changes of the Quaternary have moved all coastal anchialine cave environments repeatedly through a vertical range of over 100 m, and modern anchialine environments could not develop at their current elevations until ~4,000 years ago when sea level reached its present position. Blue holes form by a variety of mechanisms, but the most common is upward stoping and collapse from deep dissolutional voids. As a result, they provide vertical connection between different levels of horizontal cave development produced by a variety of earlier sea-level positions. Blue holes are overprinted by successive sea-level fluctuations; each sea-level event adds complexity to the habitats within blue holes and the cave systems they connect. Blue holes can reach depths below the deepest glacioeustatic sea-level lowstand, and thereby provide a refugia for anchialine species when cave passages above are drained by Quaternary sea-level fall. Blue holes represent the most significant anchialine cave environment in the world, and may provide clues to anchialine cave species colonization and speciation events.     

Atyid shrimps (Decapoda: Atyidae) influence the spatial heterogeneity of algal communities over different scales in tropical montane streams, Puerto Rico

1. Atyid (Decapoda: Atyidae) shrimps influence the distribution of algal communities over different scales in tropical montane streams of Puerto Rico. Within pools of an atyid-dominated stream, atyid shrimps enhanced patchiness in algal communities along the depth gradient. Algal bands occurred in shallow pool margins where atyids did not forage (< 3 cm below water surface), with significantly greater standing crop, taxon richness, and structural complexity than deeper areas. In deeper water, atyids reduced small-scale patchiness in algal community composition and maintained a low-growing understorey turf dominated by sessile diatoms (Bacillariophyta) and, sometimes, closely cropped, filamentous blue-greens (Cyanophyta).
2. Among pools of the atyid-dominated stream, atyids interacted with light to determine algal patchiness between stream margins and deeper areas. In sunny pools, algal standing crop was 140-fold greater in pool margins than in deeper areas where atyids foraged. In shaded pools, however, standing crop in pool margins was only 5-fold greater than in deeper areas. Effects of light on algal standing crop were greater outside atyid foraging areas than within, indicating that shrimp grazing overrides the positive effects of light.
3. In contrast to the atyid-dominated stream, algal communities in an atyid-poor stream were characterized by a high biomass of loosely attached epipelic diatoms and no depth zonation. Interstream rock and shrimp transplant experiments indicated that atyids significantly reduced algal standing crop and altered community composition on rocks from atyid-poor streams within 24 h. Results support the hypothesis that atyid shrimps play a major role in determining observed interstream differences in algal communities.     

Diurnal dynamics of the microbial loop in peatlands: structure, function and relationship to environmental parameters

Globally, introductions of alien species are increasingly common, with invasive predators potentially having detrimental effects via predation on native species. However, native prey may avoid predation by adopting new behaviors. To determine whether invasive fish populations consume endemic shrimp in invaded Hawaiian anchialine habitats or if adopted patterns of diel migration prevents predation as previously hypothesized, a total of 183 invasive poeciliids (158 Gambusia affinis and 25 Poecilia reticulata) were collected for gut content analyses from four anchialine sites during wet and dry seasons on the islands of Hawai‘i and Maui. Predation on shrimp was not detected in habitats where they retreat exclusively into the underlying aquifer diurnally and only emerge nocturnally. However, low levels of predation were detected (7/65 fishes, only by Gambusia affinis) at Waianapanapa Cave, Maui, where shrimp retreat into both the aquifer and a cave during the day. Thus, adopted behavioral responses to invasive fishes generally, though not universally, prevent predation on endemic Hawaiian anchialine shrimps. However, non-consumptive effects resulting from behavioral modification of shrimps may have appreciable impacts on the Hawaiian anchialine ecosystem and warrant further study.     

Patterns of genetic connectivity among anchialine habitats: a case study of the endemic Hawaiian shrimp Halocaridina rubra on the island of Hawaii

Anchialine habitats, landlocked bodies of mixohaline water that fluctuate with the tides but have no surface connection to the sea, are known from around the world. Many anchialine organisms have widespread distributions and it has been hypothesized that high levels of gene flow and low levels of genetic differentiation are characteristic of populations from these habitats. However, the generality of this hypothesis requires further assessment, particularly in light of the significant negative impact these habitats and their biota have experienced from anthropogenic causes. This study investigated the population structure and demography of an endemic Hawaiian anchialine species, the atyid shrimp Halocaridina rubra, using mitochondrial cytochrome c oxidase subunit I (COI) gene sequences. A survey of 305 individuals from 16 populations collected on the island of Hawaii revealed 135 haplotypes. These haplotypes belonged to one of two divergent (2.7-4.9%) lineages; notably, no haplotypes were shared between the two coasts of the island. Along each coast, strong subdivision and little to no gene flow occurs between populations separated by > 30 km. The population structure and demography of H. rubra on Hawaii are influenced by regional hydrology, geology, volcanism and two distinct colonization events of the island. Thus, H. rubra on Hawaii demonstrates that populations of endemic anchialine organisms may exhibit significant levels of genetic structure and restricted levels of gene flow over limited geographic scales. This report brings novel insight into the biology of anchialine organisms and has important implications for the future management of these habitats and their biota.     

Atyid shrimps (Decapoda: Atyidae) influence the spatial heterogeneity of algal communities over different scales in tropical montane streams, Puerto Rico

1. Atyid (Decapoda: Atyidae) shrimps influence the distribution of algal communities over different scales in tropical montane streams of Puerto Rico. Within pools of an atyid-dominated stream, atyid shrimps enhanced patchiness in algal communities along the depth gradient. Algal bands occurred in shallow pool margins where atyids did not forage (< 3 cm below water surface), with significantly greater standing crop, taxon richness, and structural complexity than deeper areas. In deeper water, atyids reduced small-scale patchiness in algal community composition and maintained a low-growing understorey turf dominated by sessile diatoms (Bacillariophyta) and, sometimes, closely cropped, filamentous blue-greens (Cyanophyta).
2. Among pools of the atyid-dominated stream, atyids interacted with light to determine algal patchiness between stream margins and deeper areas. In sunny pools, algal standing crop was 140-fold greater in pool margins than in deeper areas where atyids foraged. In shaded pools, however, standing crop in pool margins was only 5-fold greater than in deeper areas. Effects of light on algal standing crop were greater outside atyid foraging areas than within, indicating that shrimp grazing overrides the positive effects of light.
3. In contrast to the atyid-dominated stream, algal communities in an atyid-poor stream were characterized by a high biomass of loosely attached epipelic diatoms and no depth zonation. Interstream rock and shrimp transplant experiments indicated that atyids significantly reduced algal standing crop and altered community composition on rocks from atyid-poor streams within 24 h. Results support the hypothesis that atyid shrimps play a major role in determining observed interstream differences in algal communities.     

Edaphic beetle (Insecta: Coleoptera) diversity over a forest-matrix gradient in a tropical rainforest

Species from natural communities show different capacities for moving across contrasting habitats, and they may gradually respond along the transition between forests and anthropogenic areas. Because beetles are effective bioindicators, we analyzed the Coleoptera assemblage structure in the transition between sugarcane matrices and forest fragment. The study was conducted in an Atlantic rainforest fragment and sugarcane matrices after 2 and 6 months of replantation. Beetles were sampled in linear transects that were 50, 100, and 200 m from the forest edge and toward both forest core and matrices. We analyzed beetle abundance, species richness and composition, and diversity numbers. The abundance and species richness were higher in the forest, and species richness was higher at the sampling site 100 m inside the forest than at the gradients within the sugarcane matrices. The species composition differed between the forest and matrices, but not between the matrices with different replantation ages. Alpha diversity based on the number of species was higher in the forest, and alpha diversity based on the Shannon index was higher in the forest and matrix after 6 months of replantation. Our results suggest that the sugarcane matrices, chiefly the matrix after 2 months of replantation, sustain an impoverished beetle assemblage when compared with the forest fragment. Despite the strong habitat distinction, the beetle fauna in the sugarcane matrices was not affected by the horizontal distance from the forest. Here, we have highlighted the importance of forest fragments embedded in harsh matrices for the maintenance of biodiversity.     

Phylogeny and historical biogeography of the cave-adapted shrimp genus Typhlatya (Atyidae) in the Caribbean Sea and western Atlantic

Aim To infer phylogenetic relationships among five species of the cave‐adapted shrimp genus Typhlatya in order to test competing hypotheses of dispersal and colonization of the disjunct cave localities occupied by these five species. Location Typhlatya species are found in caves and anchialine ponds across the northern margin of the Caribbean Sea, along the Mediterranean and Adriatic coasts and on oceanic islands in the Atlantic and eastern Pacific oceans. This study focuses on five species, one from Bermuda, one from the Caicos Islands and three from the Yucatan Peninsula of Mexico. Methods Partial sequences (c. 1400 bp) from the mitochondrial cytochrome b, 16S rDNA and COI genes were obtained from representative samples of the five species. Phylogenetic inference was carried out with maximum parsimony and maximum likelihood analyses. Parsimony networks were constructed for the Bermudian species Typhlatya iliffei and one Yucatan species Typhlatya mitchelli, to determine the degree of connectivity among populations inhabiting different cave systems. Results All three land masses were recovered as monophyletic. The two insular marine species from Bermuda and the Caicos Islands formed a clade, while the three continental freshwater species from the Yucatan Peninsula formed another. Within both Bermuda and the Yucatan, shared haplotypes were found in different cave systems, suggesting recent or ongoing gene flow among populations in both locales. Main conclusions The two insular marine Typhlatya species originated from an ancestral marine population, possibly already cave‐adapted, that is suggested to have colonized the Caicos Islands and subsequently dispersed to Bermuda via the Gulf Stream. Divergence estimates suggest that colonization occurred before the formation of present‐day anchialine cave habitat, which did not form on either island until the late Pliocene to early Pleistocene. Divergence estimates also indicate that the Yucatan freshwater species split before the formation of freshwater cave habitat in the Yucatan. These species could have inhabited crevicular marine habitats before the late Pliocene/early Pleistocene in the Yucatan or elsewhere in the Caribbean, and subsequently migrated to freshwater caves once they formed.     

Global biodiversity and phylogenetic evaluation of remipedia (crustacea)

Remipedia is one of the most recently discovered classes of crustaceans, first described in 1981 from anchialine caves in the Bahamas Archipelago. The class is divided into the order Enantiopoda, represented by two fossil species, and Nectiopoda, which contains all known extant remipedes. Since their discovery, the number of nectiopodan species has increased to 24, half of which were described during the last decade. Nectiopoda exhibit a disjunct global distribution pattern, with the highest abundance and diversity in the Caribbean region, and isolated species in the Canary Islands and in Western Australia. Our review of Remipedia provides an overview of their ecological characteristics, including a detailed list of all anchialine marine caves, from which species have been recorded. We discuss alternative hypotheses of the phylogenetic position of Remipedia within Arthropoda, and present first results of an ongoing molecular-phylogenetic analysis that do not support the monophyly of several nectiopodan taxa. We believe that a taxonomic revision of Remipedia is absolutely essential, and that a comprehensive revision should include a reappraisal of the fossil record.     

Projected latitudinal and regional changes in vascular plant diversity through climate change: short-term gains and longer-term losses

A simple approach is suggested to project potential changes in the diversity of vascular plants. We use the current (recent past) relationship between plant diversity and geographic variation in the climate, as well as elevation range, to project changes in regional species richness (at 100 × 100 km resolution), concentrating on six climate scenarios for 2020, 2050 and 2080. The results show an overall trend towards increased vascular plant species richness. Increases in richness by 2050 and 2080 are expected over approximately three-quarters of the land surface, but decreases are expected in other regions. The magnitudes of richness gains and losses both increase over time, as the level of warming grows. The latitudinal pattern of change suggests that richness increases will be greatest at high latitudes, where plant productivity and diversity are largely limited by temperature. Richness decreases are not projected consistently in any latitudinal band, but are most likely to be observed at 5–40ºN, where declines in precipitation drive most projected decreases in richness.     

Shrimps down under: evolutionary relationships of subterranean crustaceans from Western Australia (Decapoda: Atyidae: Stygiocaris)

Background

We investigated the large and small scale evolutionary relationships of the endemic Western Australian subterranean shrimp genus Stygiocaris (Atyidae) using nuclear and mitochondrial genes. Stygiocaris is part of the unique cave biota of the coastal, anchialine, limestones of the Cape Range and Barrow Island, most of whose nearest evolutionary relations are found in coastal caves of the distant North Atlantic. The dominance of atyids in tropical waters and their food resources suggest they are pivotal in understanding these groundwater ecosystems.

Methodology/Principle Findings

Our nuclear and mitochondrial analyses all recovered the Mexican cave genus Typhlatya as the sister taxon of Stygiocaris, rather than any of the numerous surface and cave atyids from Australia or the Indo-Pacific region. The two described Stygiocaris species were recovered as monophyletic, and a third, cryptic, species was discovered at a single site, which has very different physiochemical properties from the sites hosting the two described species.

Conclusions/Significance

Our findings suggest that Stygiocaris and Typhlatya may descend from a common ancestor that lived in the coastal marine habitat of the ancient Tethys Sea, and were subsequently separated by plate tectonic movements. This vicariant process is commonly thought to explain the many disjunct anchialine faunas, but has rarely been demonstrated using phylogenetic techniques. The Cape Range''s geological dynamism, which is probably responsible for the speciation of the various Stygiocaris species, has also led to geographic population structure within species. In particular, Stygiocaris lancifera is split into northern and southern groups, which correspond to population splits within other sympatric subterranean taxa.     

Distribution patterns of macrobenthic biodiversity in the intertidal seagrass beds of an estuarine system, and their conservation significance

Intertidal stands of seagrass are important elements in the ecology of many estuaries yet the manner in which their associated macrobenthic biodiversity is distributed throughout any single estuary has never been determined. This has now been attempted for the Knysna estuarine bay in the Garden Route National Park, South Africa, an important site for one vulnerable and declining seagrass, Cape dwarf-eelgrass (Nanozostera capensis), and for several associated animals. Although spanning a salinity range of <5–35, the seagrass beds of this estuary contained a unitary fauna with local variation in frequency and abundance of the various species. Faunal biodiversity was highest along the main estuarine channel in the marine-influenced region and declined both upstream and into the sheltered peripheral zones, overall faunal abundance being inversely correlated with species diversity (though not with species richness). This pattern results in the beds around the 4 km shoreline of a single island near the mouth supporting 91 % of the total macrobenthic invertebrate species present in the system. This situation is discussed in relation to such data as are available from other comparable systems and to the conservation of estuarine seagrass faunal biodiversity. It is concluded that in general and with caveats for some individual species of concern where conservation resources are limited attention would most profitably be focused on the seagrass meadows of downstream estuarine areas.     

Effects of omnivorous shrimp in a montane tropical stream: sediment removal,disturbance of sessile invertebrates and enhancement of understory algal biomass

Freshwater shrimp dominate the faunal biomass of many headwater tropical streams: however, their role in community organization is unclear. Enclosure/exclosure experiments in a montane Puerto Rican stream examined direct and indirect effects of two dominant taxa of atyid (Atyidae) shrimp, Atya lanipes Holthuis and Xiphocaris elongata Guerin-Meneville. Both shrimp taxa caused significant reductions in sediment cover on rock substrata, reducing sedimentation and enhancing algal biovolume on clay tiles in cages. When tiles incubated in shrimp exclosures for 2 wks were placed outside of cages, atyid shrimp removed 100% of the sediment cover within a 30 min observation period. Atyid shrimp appear to play an important role in stream recovery after high discharge events by rapidly removing sediments and detritus deposited on benthic substrata in pools. We evaluated the mechanism by which A. lanipes influences algae and benthic insects by comparing patterns of algal biomass, taxonomic composition, and insect abundance between shrimp-exclusion and shrimp-presence treatments both with and without manual sediment removal. The shrimp exclusion treatment without manual sediment removal bad significantly lower algal biomass and greater sedimentation than all other treatments. The treatment in which shrimp were excluded but sediment was manually removed, however, accrued almost the same algal biovolume as the shrimp enclosure treatment, supporting the hypothesis that sediment removal enhances the biovolume of understory algal taxa. Algal community composition was similar between stream bottom bedrock exposed to natural densities of shrimp and all experimental treatments for both Atya and Xiphocaris: a diatom community strongly dominated (78–95%) by the adnate taxon, Achnanthes lanceolata Breb ex. Kutz. Atyid shrimp are important in determining the distribution and abundance of benthic insects through both direct and indirect effects. Sessile, retreat-building chironomid larvae (Chironomidae: Diptera) are negatively affected by both A. lanipes and X. elongata, through direct removal by foraging activities and/or indirectly through depression of sediment resources available to larvae for the construction of retreats. In constrast, the mobile grazer, Cloeodes maculipes (Baetidae: Ephemeroptera) was not adversely affected and atyid shrimp have the potential to exert positive indirect effects on this taxon by facilitating its exploitation of algal resources and/or through enhancement of understory algal food resources through sediment removal.     

Seashore in the mountain: limestone‐associated land snail fauna on the oceanic Hahajima Island (Ogasawara Islands,Western Pacific)

Because land snails inhabiting the seashore are most likely to be carried by ocean currents or by attaching to seabirds, land snail fauna on oceanic islands include species derived from the mainland ancestors inhabiting the seashore. If habitat use of the island descendants is constrained by the ecology of the mainland ancestor, the island species that moved from the coastal habitat to the inland habitat may still be restricted to relatively exposed microhabitats with high pH, calcium carbonate‐rich substrates, and poor litter cover. We tested this hypothesis by investigating the association between environmental conditions and species diversity of seashore‐derived species of the endemic land snails on the oceanic Hahajima Island (Ogasawara Islands). Seashore‐derived species showed higher species richness on limestone outcrops than non‐limestone areas, whereas the other species showed no significant increase in species richness in limestone outcrops. There was a higher proportion of seashore‐derived species on the limestone ridges than on the soil of dolines, even in the limestone area. Accordingly, the species derived from the seashore of the mainland are restricted to microhabitats with poor vegetation cover, poor litter cover, high pH, and calcium carbonate‐rich substrates, which supports the hypothesis that the inland species on an island derived from the mainland seashore still prefer environments similar to the seashore. In addition, the seashore‐derived species on the limestone outcrop include cave‐dwellers lacking functional eyes. This suggests that the probability of colonizing a cave environment is restricted to seashore‐derived species. The findings obtained in the present study suggest that habitat use of the ancestral lineages can constrain habitat use of the descendants, even in the oceanic islands with depauperate fauna. This bias in the species composition on the limestone outcrop constrains lineages that can colonize and adapt to the inside of caves, and therefore, habitat use of the ancestral lineages affects the ability of descendant lineages to colonize novel habitats. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 686–693.