Patterns of adult abundance in Chthamalus stellatus (Poli) and C. montagui Southward (Crustacea: Cirripedia) emerge during late recruitment

The aim of this study was to investigate when adult distribution patterns are established in the barnacles Chthamalus stellatus and C. montagui. Adult ‘zones’ were identified by analysing field counts of both species at mid and upper shore heights. Monthly collections of cyprids, < 1 month old metamorphs and recruits (all metamorphosed individuals older than approximately 1 month) were made for C. stellatus and C. montagui in natural barnacle beds at six shores in SW Ireland. This was carried out over one year in 1996/1997, using a hierarchical sampling design. Abundance of total recruits (0-3 months old) was compared between adult zones after the main settlement season had ended. In addition, scales of variability in 0-3 month recruitment into adult zones were compared between the species at two scales: shores (1000s of metres) and sites within shores (10s of metres). Older recruits of each species, up to 11 months of age, were also compared between adult zones.The majority of settlement (measured as attached cyprids) occurred between August and October 1996. In October, there was no effect of adult zone on the abundance of total (0-3 month) recruitment up to that point in either species. Despite this homogeneity in recruitment between adult zones, significant spatial variation was found in 0-3 month recruits of both species at both of the scales examined. In C. stellatus the amount of variation associated with the larger scale (shore) was more than twice that of sites or of the residual variation (replicates within sites). 0-3 month recruitment in C. montagui was also most variable at the scale of shores but the residual variability (between replicates within site) was of similar magnitude to that of shores. Variability in 0-3 month C. montagui recruitment was relatively low at the scale of sites.There was a small but consistent input of recruits to adult zones over 9 months of the year, complicating the assessment of when adult patterns were set-up in these species. By June 1997, characteristic patterns of adult dominance had been established at all shores. Settlement had completely ceased by this time and individual barnacles were potentially 11 months old. Neither settlement nor early recruitment are significant in determining adult zonation patterns in these species. Instead, differential mortality patterns in individuals up to the age of 11 months are implicated in determining patterns of distribution of both species.     

Spatial analyses of intertidal assemblages on sheltered rocky shores

Abstract Understanding processes in complex assemblages depends on good understanding of spatial and temporal patterns of structure at various spatial scales. There has been little quantitative information about spatial patterns and natural temporal changes in intertidal assemblages on sheltered rocky shores in temperate Australia. Natural changes and responses to anthropogenic disturbances in these habitats cannot be accurately measured and assessed without quantitative data on patterns of natural variability in space and through time. This paper describes some suitable quantitative methods for examining spatial and temporal patterns of diversity and abundances of highshore, midshore and lowshore intertidal assemblages and the important component species for a number of shores in a bay that has not been severely altered by human disturbance. Despite a diverse flora and fauna on these shores, the midshore and lowshore assemblages on sheltered shores were characterized by a few species which were also the most important in discriminating among assemblages on a shore and, for each assemblage, among different shores. The same set of species was also important for measuring small-scale patchiness within each assemblage (i.e. between replicate sites on a shore). Therefore, these data provide a rationale for selecting species that are useful for measuring differences and changes in abundance among places and times at different scales and, hence, can be used in the more complex sampling designs necessary to detect environmental impacts. There was considerable spatial variability in all assemblages and all species (or taxa) examined at scales of metres, tens of metres and kilometres. There were no clear seasonal trends for most measures, with as much or more variability at intervals of 3 months as from year to year. Most interactions between spatial and temporal measures were at the smallest scale, with different sites on the same shore generally showing different changes from time to time. The cause(s) of this apparently idiosyncratic variability1 were not examined, but some potential causes are discussed. These data are appropriate for testing hypotheses about the applicability of these findings to other relatively undisturbed sheltered shores, about effects of different anthropogenic disturbances on sheltered intertidal assemblages and to test hypotheses about differences in intertidal assemblages on sheltered versus wave-exposed shores.     

Organization of Squamata (Reptilia) assemblages in Mediterranean archipelagos

Mediterranean islands have complex reptile assemblages, but little is known about the factors that determine their organization. In this study, the structure of assemblages of Squamata was evaluated based on their species richness and two measures of phylogenetic diversity (variability and clustering). I evaluated the composition of the assemblages comparing distinct biogeographic subregions within the Mediterranean: Adriatic, Aegean, Balearic, Corsica–Sardinia, Crete, Gulf of Gabés, Ionian Sea, Ligurian Sea, Malta, Sicily, and Tyrrhenian Sea. The effect of island environments and geographical isolation on the diversity metrics was assessed using generalized linear models. The analyses indicated that species richness was mostly influenced by island area and geographical isolation. Assemblages on smaller islands were poorer in species and phylogenetically dispersed, possibly as an effect of interspecific competition. The species composition of the assemblages was determined by similar environmental drivers within the biogeographic subregions, including island area, island elevation, geographical isolation, and aridity. In several subregions, significant patterns of phylogenetic attraction were found in species co‐occurrences, caused by the limits imposed by the island size on large predatory species.     

Comparative genetic isolation patterns for multiple headwater fishes in three geographic regions

Headwater‐resident fishes may be prone to a high rate of isolation and a pronounced hierarchical genetic structure due to a combination of niche preference and dendritic effects of river networks. Genetic isolation patterns were compared using microsatellites in six headwater fishes, Fundulus olivaceus, Semotilus atromaculatus, Erimyzon claviformis, Etheostoma artesiae, Etheostoma whipplei and Etheostoma parvipinne, in three geographic regions that included drainages of small, medium and large sizes in the southern United States. All species showed hierarchical nesting of genetic populations and there were clear and mostly consistent differences between species and regions that were identified through summary statistics derived from two independent analyses. For species comparisons, a high isolation grouping (increased number of isolated genetic clusters or sites within regions) and a low‐isolation grouping (decreased number of clusters or sites) were identified. Species group placement was related to niche breadth along the river continuum and presumed abundance and variability of preferred microhabitats, with increased headwater specialization among species being associated with placement in the high‐isolation grouping. There was a weakly significant positive effect of drainage size on the number of isolated clusters or sites across all species. Regional patterns were shared in two species, with the region containing the smallest drainages having lower rates of isolation in both datasets. This study shows the effects of regional and species characteristics on genetic isolation for headwater species, which are especially prone to isolation due to spatial, dendritic effects of river networks.     

Genetic differentiation within and between isolated Algerian subpopulations of Barbary macaques (Macaca sylvanus): evidence from microsatellites

This study of wild-living Algerian Barbary macaques ( Macaca sylvanus ) was designed to examine genetic variability in subpopulations isolated in residual forest patches, in an attempt to obtain data on the effects of habitat fragmentation. The wild population of this species (estimated at a maximum of 15 000) is vulnerable and this study therefore has direct relevance for conservation measures. Data from five microsatellite loci were analysed for 159 individuals from nine different groups living in four isolates in Algeria. Genetic polymorphism was found to be relatively high (4–12 alleles per locus) compared with other genetic markers used in previous studies of this species; mean expected heterozygosity was 65%. The four isolates are all genetically distinct ( F _ST = 0; P < 0.001). Indeed, the results suggest that dispersal is limited even between some social groups within a single isolate. Genetic distances based on models not assuming stepwise mutation ( F _ST and chord distance) gave very similar results and are highly correlated with geographical distances within one isolate but not between isolates. This may indicate that isolation by distance exerts a significant influence within an isolate but that genetic drift prevails between the four isolates. After allowing for variation in sample size, we found no evidence of reduced allelic diversity within small isolates that may have been separated for 250 years or more. The surviving population of Algerian Barbary macaques taken as a whole still shows marked variability in microsatellite alleles, but maintenance of genetic variability over the long term will surely require effective protection of all isolates.     

Evaluating plant-soil feedback together with competition in a serpentine grassland

Plants can alter biotic and abiotic soil characteristics in ways that feedback to change the performance of that same plant species relative to co-occurring plants. Most evidence for this plant-soil feedback comes from greenhouse studies of potted plants, and consequently, little is known about the importance of feedback in relation to other biological processes known to structure plant communities, such as plant-plant competition. In a field experiment with three C4 grasses, negative feedback was expressed through reduced survival and shoot biomass when seedlings were planted within existing clumps of conspecifics compared with clumps of heterospecifics. However, the combined effects of feedback and competition were species-specific. Only Andropogon gerardii exhibited feedback when competition with the clumps was allowed. For Sorghastrum nutans, strong interspecific competition eliminated the feedback expressed in the absence of competition, and Schizachyrium scoparium showed no feedback at all. That arbuscular mycorrhizal (AM) fungi may play a role in the feedback was indicated by higher AM root colonization with conspecific plant neighbours. We suggest that feedback and competition should not be viewed as entirely separate processes and that their importance in structuring plant communities cannot be judged in isolation from each other.     

Body mass, habitat generality, and avian community composition in forest remnants

Aim Many theories of biodiversity and biogeography assume that species respond equally to variability in habitat area and isolation. This assumption does not allow for differential responses due to interspecific competition or other mechanisms, and therefore does not allow community composition to be predicted. As body size is relevant to area requirements and interspecific dominance, a natural experiment was conducted to quantify the differential responses of avian species abundance to variability in remnant area, isolation and forest cover based on average species body mass. Location Deciduous forest remnants of varying area and isolation throughout the State of Delaware, USA. Methods Forest remnants within stratified area and isolation classes were randomly selected for breeding bird surveys; total forest cover (ha) within 2 km of each survey point was subsequently determined as a covariate. Surveys were conducted within 100–150 m from the edge of each remnant and detected bird species were divided into five classes based on a log₂‐transformation of body mass (very small, small, medium, large and very large). Assuming a negative binomial distribution, the abundance of detected individuals in each mass class was analysed using generalized linear models with remnant area, isolation, local forest cover and two‐way interactions specified as independent variables. The same analyses were conducted for individual species where sample size allowed. Results Very small, small and very large bodied species decreased in abundance with decreasing local forest cover and remnant area and with increasing remnant isolation, while large species increased in abundance. Medium‐sized species decreased in abundance with increasing forest cover, did not respond to remnant isolation and showed a concave, curvilinear response to increasing remnant area. Large and medium‐sized species were the most abundant birds in small, isolated remnants despite occurring in the largest remnants with the more abundant very small and small species, suggesting that communities are not randomly organized. Main conclusions Regardless of presumed habitat associations, large and medium‐sized species are of the appropriate size to be dominant competitors when forest resources are limiting, and thus may be considered ‘generalists’. The smallest species may be excluded entirely from small, isolated remnants even though such remnants meet their ecological needs; the needs of very large species are not met in small remnants. The applicability of biodiversity theories to community composition, species abundance and, by extension, to conservation, can be improved by incorporating differential responses based on body mass into their assumptions.     

A striking example of the founder effect in the mollusc Littorina saxatilis

Two South African populations of Littorina saxatilis were examined by starch-gel electrophoresis at 16 enzyme loci and compared with 13 populations of North Atlantic saxatilis from both American and European coasts, and with six British populations of the closely related species Littorina arcana. The South African animals showed a severely reduced heterozygosity (– 0.052) compared with Atlantic populations of saxatilis ( = 0.181), and the mean genetic distance between the two areas was high ( = 0.203) compared with distances within the North Atlantic saxatilis populations (D = 0.034). In fact, the saxatilis from South Africa were genetically more distant from the North Atlantic samples of L. saxatilis than were the arcana from British shores. The reduced genetic heterozygosity and genetic divergence of the South African populations is attributed to founder effects following a postulated recent introduction by man.     

Contrasting spatial heterogeneity of sessile organisms within mussel (Perna perna L.) beds in relation to topographic variability

We examined the spatial heterogeneity in three sessile rocky shore organisms, the mussel Perna perna, the barnacle Octomeris angulosa (Sowerby) and the red alga Gelidium pristoides (Turn.) at a range of continuous local scales along horizontal transects within mid- and upper mussel beds of South African shores. We also examined the relationships between variability of organisms and topographic features (rock depressions, slope, aspect), and between mussel, barnacle and algal variability over the same scales. To estimate spatial heterogeneity, we analyzed scaling properties of semivariograms using a fractal approach. Relationships between different variables at the different scales were examined by cross-semivariograms. Spatial dependence of P. perna variability increased with spatial dependence of topographic variability, so that scaling regions of mussel and topographic distributions corresponded well. This relationship often improved with larger local scales (mussel cover increased with depressions, steeper slope and aspect towards waves), while at smaller spatial scales, variability in mussel cover was less well explained by variability in topography. The variability of the barnacle O. angulosa exhibited spatial dependence, even on topographically unstructured shores. In contrast, the distribution of the alga G. pristoides revealed high fractal dimensions, showing spatial independence on topographically unstructured shores. Algae also showed a very strong negative relationship with mussels at most local scales, and a negative relationship with barnacles in upper zones, especially at larger local scales. Barnacles may show clear spatial dependence because of hydrodynamics (at larger local scales) and the need to find a future mate in close proximity (at smaller local scales), while algae may show a strong negative relationship with mussels because of competition for space.     

Life-history variations of the Lesser Sheathbill Chionis minor in contrasting habitats

From 1987 until 1995, life-history traits of the Lesser Sheathbill Chionis minor in the Kerguelen Archipelago were compared with those at other localities, where this species breeds in seabird colonies. At Kerguelen Island only, some pairs breed on shores free of penguin and cormorant colonies. Moreover, pairs, including nonbreeders, and solitary individuals maintain territories all the year round. Site and mate changes were not linked, and all divorces were permanent. Fidelity rates were similar in all localities and so were the annual survival rates of adults. The survival rate of immature individuals was highest at Kerguelen Island, where sheathbills laid smaller clutches, produced fewer fledglings and had lower breeding success than in the other islands. At Kerguelen Island, pairs breeding on shores had similar clutch sizes but fledged fewer offspring than those breeding in seabird colonies. Further, the proportion of vacant territories and nonbreeders was higher on shores. Kerguelen sheathbills devoted less time to food acquisition than those on Marion Island, being exceptionally kleptoparasitic, and spent a smaller part of their foraging time exploiting seabird colonies. Their diet was mainly algae. The differences on Kerguelen Island as compared with other localities were caused by the presence of an extensive intertidal zone on the former, which reduced competition, making many more sites suitable and the environment more predictable. These life-history and behavioural traits are discussed in relation to life-history theory. As differences were also found, although less extensive, within the Kerguelen Archipelago, we suggest that some traits represent an adaptive response to external constraints and that the life-history strategy of the Lesser Sheathbill is particularly opportunistic.     

Not as the crow flies: assessing effective isolation for island biogeographical analysis

Aim To evaluate the role of island isolation in explaining the distribution of vascular plant species in a dense freshwater archipelago, specifically comparing conventional measures of island isolation with landscape measures of island isolation. Location Data were collected from 35 islands within Massasauga Provincial Park on the eastern shores of the Georgian Bay, Ontario, Canada. Methods Sampled islands were located using stratified random selection based on location and size variation. The number of species was recorded along stratified random transects. Island isolation variables included distance to the mainland, distance to the nearest island, largest gap in a stepping‐stone sequence, distance to the closest upwind point of land, and a landscape measure of island isolation. The landscape measure of isolation was quantified as the percentage of the land area within 100, 250, 500, 1000, 1500 and 2000 m of each island’s perimeter. The isolation variables were calculated within a geographical information system (GIS). Dependent variables in the regression analyses included species richness, the logarithm of species richness and residuals of the species–area relationship. Independent variables included island isolation variables and their logarithmic transformations. Results Isolation plays a role, albeit small, in explaining species richness in the study area. In the regression analyses, the landscape measure of isolation provided a better fit than conventional measures of island isolation. Islands with less land than water within a 250‐m buffer were more effectively isolated and had fewer species present than islands surrounded by a greater proportion of water. Main conclusions Consistent with the species–isolation relationship, fewer species were present on more isolated islands within the Massasauga study area, as elucidated using a series of island buffers in a GIS. Applying a landscape measure of isolation to similar dense, freshwater archipelagos may elucidate species–isolation patterns not evident through conventional, straight‐line distance measurements of island isolation. The low value of the regression coefficients as well as the isolation history and high density of the Massasauga islands suggests caution in extending the results, especially to dissimilar archipelagos.     

Competition, sub-lethal mortality and diversity on Southern Ocean coastal rock communities

Overgrowth competition, sub-lethal mortality (some zooids killed but colony survives) and subsequent growth from fragments, and diversity of communities encrusting rocks (pebble through to boulder size) were examined from five Antarctic localities, along a latitudinal gradient. There were distinct gradients in the ecology of both assemblages and individual species with latitude within the Southern Ocean. Compared with warm-water equivalents, the polar assemblages had many fewer species, considerably less variability in species richness, highly transitive interactions and lower incidences of interspecific encounters. There is no gradual transition but a dramatic alteration of assemblage and species level ecology around the margins of the Southern Ocean, which may primarily be due to ice scour. Sub-lethal mortality was common and peaked on different boulder sizes along the Antarctic Peninsula. This is indicative of the disturbance cline along the region.     

Biodiversity in extreme aquatic environments: Lakes, ponds and streams of the Ross Sea sector, Antarctica

The Ross Sea Sector (RSS) of Antarcticallies between the lines of longitude 150°E and 150°W and contains diverse landscapes with a variety of lakes, ponds and streams. Neither insects nor crustacean species have been recorded in these ecosystems but most contain planktonic and/or benthic communities that are composed exclusively of microscopic organisms. Microbial brodiversity is low with a small number of species (e.g. filamentous cyanobacteria of the family Oscillatoriaceae) occurring under a broad range of environmental conditions throughout the region. There is no evidence to date of microbial endemism in the RSS; however, there is a need to apply molecular and cellular techniques to compare biodiversity and genetic characteristics with assemblages elsewhere in Antarctica and with comparable communities in the north polar zone. A series of hypotheses are advanced to help guide further work. These derive from the conclusion that environmental extremes plus biogeographical isolation control the biodiversity of RSS communities, and that biological interactions (competition, grazing, predation, parasitism) are weak and play a minor role by comparison with temperate latitude ecosystems.     

Isozyme variability of the wetland specialist Swertia perennis (Gentianaceae) in relation to habitat size, isolation, and plant fitness

We examined the effects of size and spatial isolation of fens on the isozyme variability of 17 populations of Swertia perennis. This long-lived perennial is a locally abundant fen specialist in Switzerland, where wetlands have been strongly fragmented. Isozyme variability was comparable to other outcrossing plants (A = 1.53, AP(p) = 2.01, P(p) = 42.5, H(o) = 0.113, H(e) = 0.139). F statistics indicated both inbreeding within and differentiation between populations (F(IS) = 0.076, F(IT) = 0.194, F(ST) = 0.128), with moderate gene flow between populations (N(e)m = 1.703). Populations in small, isolated fens had reduced genetic variability and the highest within-population inbreeding coefficients (F(IS)). Isozyme variability was significantly associated with vegetative fitness traits (MANOVA), and the magnitude of leaf herbivory decreased as the percentage of polymorphic loci increased. These data suggest that the reduced genetic variability of S. perennis in small, isolated populations may reduce plant fitness, thereby increasing susceptibility to herbivore damage. Our study also shows that habitat fragmentation can reduce the genetic variability of populations of fairly common habitat specialists, which so far have attracted less conservation attention than rare species.     

Actinoplanetes in soil and on plant litter from freshwater habitats

An isolation method for actinoplanetes is described which uses the ability of the sporangium to withstand desiccation and release motile spores when subsequently rehydrated. The desiccation stage reduces the number of associated Gram negative bacteria which cause problems when attempts are made to isolate species on agar plates from natural substrates. The method has enabled the isolation of actinoplanetes directly from soil and plant litter and indirectly from soil with the aid of baits such as pollen and hair. Actinoplanetes appear to be common in dry soils collected from arid areas and sand dune systems. They are normally present on leaves submerged for a time in rivers, streams and lakes and subsequently deposited in heaps on shores or as debris on overhanging vegetation. An actinomycete resembling the 'spore-dome' organism recovered from leaves and twigs by Willoughby (1969) was frequently isolated during the course of these studies. Preliminary work suggests that different species can be recognized and could constitute a new genus within the family Streptomycetaceae.     

Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina

A phylogenetic approach to the origin and maintenance of species diversity ideally requires the sampling of all species within a clade, confirmation that they are evolutionarily distinct entities, and knowledge of their geographical distributions. In the marine tropics such studies have mostly been of fish and reef-associated organisms, usually with high dispersal. In contrast, snails of the genus Echinolittorina (Littorinidae) are restricted to rocky shores, have a four-week pelagic development (and recorded dispersal up to 1400 km), and show different evolutionary patterns. We present a complete molecular phylogeny of Echinolittorina, derived from Bayesian analysis of sequences from nuclear 28S rRNA and mitochondrial 12S rRNA and COI genes (nodal support indicated by posterior probabilities, maximum likelihood, and neighbor-joining bootstrap). This consists of 59 evolutionarily significant units (ESUs), including all 50 known taxonomic species. The 26 ESUs found in the Indo-West Pacific region form a single clade, whereas the eastern Pacific and Atlantic species are basal. The earliest fossil occurred in the Tethys during the middle Eocene and we suggest that the Indo-West Pacific clade has been isolated since closure of the Tethyan seaway in the early Miocene. The geographical distributions of all species (based on more than 3700 locality records) appear to be circumscribed by barriers of low temperature, unsuitable sedimentary habitat, stretches of open water exceeding about 1400 km, and differences in oceanographic conditions on the continuum between oceanic and continental. The geographical ranges of sister species show little or no overlap, indicating that the speciation mode is predominantly allopatric. Furthermore, range expansion following speciation appears to have been limited, because a high degree of allopatry is maintained through three to five branching points of the phylogeny. This may be explained by infrequent long-distance colonization, habitat specialization on the oceanic/continental gradient, and perhaps by interspecific competition. In the eastern Pacific plus Atlantic we identify five cases of divergence on either side of the Isthmus of Panama, but our estimates of their ages pre-date the emergence of the Isthmus. There are three examples of sister relationships between species in the western Atlantic and eastern Atlantic, all resulting from dispersal to the east. Within the Indo-West Pacific, we find no geographical pattern of speciation events; narrowly endemic species of recent origin are present in both peripheral and central parts of the region. Evidence from estimated divergence times of sister species, and from a plot of the number of lineages over time, suggest that there has been no acceleration of diversification during the glacio-eustatic cycles of the Plio-Pleistocene. In comparison with reefal organisms, species of Echinolittorina on rocky shores may be less susceptible to extinction or isolation during sea-level fluctuations. The species richness of Echinolittorina in the classical biogeographic provinces conforms to the common pattern of highest diversity (11 species) in the central "East Indies Triangle" of the Indo-West Pacific, with a subsidiary focus in the eastern Pacific and western Atlantic, and lowest diversity in the eastern Atlantic. The diversity focus in the East Indies Triangle is produced by a mosaic of restricted allopatric species and overlap of a few widespread ones, and is the result of habitat specialization rather than historical vicariance. This study emphasizes the plurality of biogeographic histories and speciation patterns in the marine tropics.     

Scales of spatial patterns of distribution of intertidal invertebrates

Few comparative studies of spatial patterns at different scales have examined several species in the same habitat or the same species over a range of habitats. Therefore, variability in patterns among species or among habitats has seldom been documented. This study quantifies spatial patterns of a suite of intertidal snails and a species of barnacle using a range of statistical techniques. Variability in densities was quantified from the scale of adjacent quadrats (over a distance of centimeters) to tens of kilometers. Significant differences in abundances occurred primarily at two spatial scales. Small-scale differences were found at the scales of centimeters or 1–2 m and, for many species on many shores, these accounted for most of the variability in abundances from place to place. These are likely to be determined by behavioural responses to small-scale patches of microhabitat. Large-scale differences in abundance were also found in most species at the scale of hundreds of meters alongshore. These are likely to be due to variation in recruitment (and/or mortality) because of limited dispersal by adults of these species. There was little or no additional variation among shores, separated by tens of kilometers, than was shown among patches of shore separated by hundreds of meters. Identification of the scale(s) at which significant differences in abundance are found focus attention on the processes (and the scales at which these processes operate) that influence patterns of distribution and abundance. Some of the advantages and disadvantages of various procedures are discussed.     

Evolutionary Dynamics in the Southwest Indian Ocean Marine Biodiversity Hotspot: A Perspective from the Rocky Shore Gastropod Genus Nerita

The Southwest Indian Ocean (SWIO) is a striking marine biodiversity hotspot. Coral reefs in this region host a high proportion of endemics compared to total species richness and they are particularly threatened by human activities. The island archipelagos with their diverse marine habitats constitute a natural laboratory for studying diversification processes. Rocky shores in the SWIO region have remained understudied. This habitat presents a high diversity of molluscs, in particular gastropods. To explore the role of climatic and geological factors in lineage diversification within the genus Nerita, we constructed a new phylogeny with an associated chronogram from two mitochondrial genes [cytochrome oxidase sub-unit 1 and 16S rRNA], combining previously published and new data from eight species sampled throughout the region. All species from the SWIO originated less than 20 Ma ago, their closest extant relatives living in the Indo-Australian Archipelago (IAA). Furthermore, the SWIO clades within species with Indo-Pacific distribution ranges are quite recent, less than 5 Ma. These results suggest that the regional diversification of Nerita is closely linked to tectonic events in the SWIO region. The Reunion mantle plume head reached Earth’s surface 67 Ma and has been stable and active since then, generating island archipelagos, some of which are partly below sea level today. Since the Miocene, sea-level fluctuations have intermittently created new rocky shore habitats. These represent ephemeral stepping-stones, which have likely facilitated repeated colonization by intertidal gastropods, like Nerita populations from the IAA, leading to allopatric speciation. This highlights the importance of taking into account past climatic and geological factors when studying diversification of highly dispersive tropical marine species. It also underlines the unique history of the marine biodiversity of the SWIO region.     

Variation in rocky shore assemblages in the northwestern Mediterranean: contrasts between islands and the mainland

Insular assemblages of species are often considered unique because they are exposed to unpredictable patterns of colonization/extinction that depend on distance from other sources of colonists and on size of islands. An alternative explanation is that islands provide fundamentally different habitats of those of the mainland, regardless of any possible effect of size and isolation. These alternatives were examined by comparing assemblages of rocky shores on islands of the Tuscany Archipelago with those of the mainland in the same geographical region. Sandy beaches created a pattern of discrete areas of rock along the mainland with spatial discontinuities and extents comparable to those of the insular environment. Possible effects of isolation and size were therefore controlled in this study, so that one would expect no difference between islands and the mainland if only size and isolation matter. In contrast, differences are expected if historical events or other processes have distinct influences on assemblages in these environments. These hypotheses were tested by comparing assemblages of midshore and lowshore habitats of two islands with those of two similarly distributed locations on the mainland over a period of 2 years, using a hierarchical sampling design. Multivariate and univariate analyses revealed various patterns in the data. There were differences between islands and the mainland in structure of assemblages, in mean abundance of common taxa and in the magnitude of spatial and temporal variance in abundance in both habitats. Collectively, these findings support the model that islands in the Tuscany Archipelago have distinct assemblages from the mainland, thereby contributing to the regional diversity and complexity of assemblages of rocky shores over and above any possible effect of size or isolation.     

Resource control by territorial male cichlid fish in Lake Malawi

1. The rocky habitat communities of Lake Malawi contain a high diversity of ecologically similar, predominantly herbivorous, cichlid fish species. How so many species can coexist is still unknown.
2. Adult males of the majority of these species hold permanent territories which form dense multispecies mosaics across the shores.
3. The study examined six coexisting species of cichlids from the rocky habitat where males are known to control access to the algal food resources within their territorial space. These included four sibling species of the Pseudotropheus ( Tropheops ) species complex, which are specialized epilithic algal herbivores, and two sibling species of the trophically more generalist P . ( Maylandia ) species complex which feed upon both epilithic algae and plankton.
4. The hypotheses that both intraspecific and interspecific territoriality occur in these communities and that interspecific aggression initiated by territorial males is preferentially directed at species with the most similar diets to their own were tested.
5. It was found that males preferred to exclude conspecific intruders, while they tolerated intruders with different diets to their own. Also, male P . ( Tropheops ) preferentially excluded similar heterospecific dietary specialists, while male P. ( Maylandia ) tolerated similar heterospecific dietary generalists.
6. Based on this study, it is proposed that interspecific territoriality may be reducing competition between species with different diets and promoting their coexistence, while it may be enhancing competition between species with the same diet. Furthermore, it suggests that ecological generalism may be reducing the intensity of interspecific competition, while specializations for the same resources may be increasing competition.