Taxonomy,natural history and zoogeography of sand beach isopods from the coast of Southern Chile

Seven species of marine isopods were collected on sandy beaches of Southern Chile (from about 37° S until 42° S). From these, Excirolana hirsuticauda Menzies, Excirolana monodi Carvacho, Chaetilia paucidens Menzies, Macrochiridothea setifer Menzies and M. mehuinensis Jaramillo are known only from the Chilean coasts. Another species, Excirolana braziliensis Richardson, is known from other areas of American coasts. One species is identified here as Macrochiridothea aff. lilianae by its resemblance with M. lilianae Moreira from Southern Brazil. All these species are here briefly described and some aspects of their ecology added. Also Zoogeographic characterization of the studied area is discussed in relation to the distributional patterns of isopods living in them.     

Excirolana braziliensis, a Pan-American sand beach isopod: taxonomic status, zonation and distribution

Excirolana braziliensis Richardson is shown to be the senior synonym of Cirolana salvadorensis Schuster and C. koepckei Bott. The latter two species, incorrectly assigned to Cirolana , were formerly regarded as distinct and were separated on the basis of size-dependent characters. Sampling of sandy beaches along the eastern Pacific from Mexico to Chile (21 localities) and the western Atlantic from Mexico to Brazil, including the West Indies (20 localities), demonstrates that E. braziliensis is commonly the most abundant macroscopic organism present in this habitat. Marked differences in zonation of young and adults are illustrated from data in Panama.     

Isolation and characterization of 11 polymerase chain reaction primers for microsatellite loci for the Chilean marine isopod Excirolana hirsuticauda

The Chilean isopod Excirolana hirsuticauda is a marine benthic brooder with wide distributional range and low potential for long-distance dispersal. Eleven microsatellite markers were developed for E. hirsuticauda using enriched libraries. Characterization of those loci in 35 individuals from Playa Blanca beach showed high allelic diversity with a mean of 10.9 alleles per locus. The average expected and observed heterozygosities were 0.65 and 0.41. These microsatellite loci are the first published for any Excirolana species and should be useful to study the genetic structure of E. hirsuticauda.     

Sensitivity of a cirolanid isopod to human pressure

Cirolanid isopods are conspicuous members of the supralittoral and intertidal fringes of sandy beaches around the world, being dominant in terms of number or biomass. Excirolana braziliensis is one of the most abundant species on exposed sandy beaches, both urbanized and preserved, of Rio de Janeiro in southeast Brazil. Considering the negative effects of urbanization and human pressure on sandy beaches, this study aimed to analyze the population structure and reproductive aspects of E. braziliensis in different stretches of Barra da Tijuca beach (Rio de Janeiro, Brazil), which differ with respect to urbanization and occupation by bathers. Monthly samplings of E. braziliensis were conducted throughout 12 months within urbanized and preserved stretches, including measurements of beach parameters and human pressure. The anthropogenic effect seems to be a relevant factor in explaining the variability in the population structure of this species. Negative significant correlations were found between the species density and the number of visitors, who massively occupies the urbanized stretches. Similar life history strategies were observed for different populations of E. braziliensis. Although high fecundity rates were reported to all stretches, the probability of eggs/embryos survival under the adverse conditions provided by the Barra da Tijuca beach is not clearly known. According to the results of this study, it could be inferred that the human pressure over Barra da Tijuca beach affects the populations: (1) directly, through human trampling and/or natural habitat jeopardizing; or (2) indirectly, by the isolation of individuals in the preserved stretch, located between environments subjected to intense disturbance. In such case, the species strategy to thrive in a protected area of restricted size, within a highly urbanized and occupied area by bathers does not appear to be the best conservation measure for peracarid species, as in E. braziliensis. Nevertheless, E. braziliensis turned out to be a good monitoring species of impacts due to its high resistance to environmental stress, persisting in highly urbanized areas dominated by bathers.     

TEMPORAL VARIATION IN POPULATIONS OF THE MARINE ISOPOD EXCIROLANA: HOW STABLE ARE GENE FREQUENCIES AND MORPHOLOGY?

Excirolana braziliensis is a dioecious marine isopod that lives in the high intertidal zone on both sides of tropical America. It lacks a dispersal phase and displays a remarkable degree of genetic divergence even between localities less than 1 km apart. Nine populations of this nominal species from both sides of the Isthmus of Panama and one population of the closely allied species, Excirolana chamensis, from the eastern Pacific were studied for 2 yr for allozymic temporal variation in 13 loci and for 3 to 4 yr for morphological variation in nine characters. The genetic and morphological constitution of 9 out of 10 populations remained stable. Allele frequencies at two loci and overall morphology in a tenth beach occupied by E. braziliensis changed drastically and significantly between 1986 and 1988. The change in gene frequency is too great to explain by genetic drift occurring during a maximum of 14 generations regardless of assumed effective population size; drift is also unlikely to have caused observed changes in morphology. Selective survival of a previously rare genotype is more plausible but still not probable. The most credible explanation is that the resident population at this locality became extinct and that the beach was recolonized by immigrants from another locality. Such infrequent episodes of extinction and recolonization from a single source may account for the large amount of genetic divergence between local populations of E. braziliensis. However, the low probability of large temporal genetic change even in a species such as this, in which gene flow between local demes is limited and generation time is short, suggests that a single sample through time is usually adequate for reconstructing the genetic history of populations.     

Phylogeographic patterns of a lower intertidal isopod in the Gulf of California and the Caribbean and comparison with other intertidal isopods

A growing body of knowledge on the diversity and evolution of intertidal isopods across different regions worldwide has enhanced our understanding on biological diversification at the poorly studied, yet vast, sea–land interface. High genetic divergences among numerous allopatric lineages have been identified within presumed single broadly distributed species. Excirolana mayana is an intertidal isopod that is commonly found in sandy beaches throughout the Gulf of California. Its distribution in the Pacific extends from this basin to Colombia and in the Atlantic from Florida to Venezuela. Despite its broad distribution and ecological importance, its evolutionary history has been largely neglected. Herein, we examined phylogeographic patterns of E. mayana in the Gulf of California and the Caribbean, based on maximum‐likelihood and Bayesian phylogenetic analyses of DNA sequences from four mitochondrial genes (16S rDNA, 12S rDNA, cytochrome oxidase I gene, and cytochrome b gene). We compared the phylogeographic patterns of E. mayana with those of the coastal isopods Ligia and Excirolana braziliensis (Gulf of California and Caribbean) and Tylos (Gulf of California). We found highly divergent lineages in both, the Gulf of California and Caribbean, suggesting the presence of multiple species. We identified two instances of Atlantic–Pacific divergences. Some geographical structuring among the major clades found in the Caribbean is observed. Haplotypes from the Gulf of California form a monophyletic group sister to a lineage found in Venezuela. Phylogeographic patterns of E. mayana in the Gulf of California differ from those observed in Ligia and Tylos in this region. Nonetheless, several clades of E. mayana have similar distributions to clades of these two other isopod taxa. The high levels of cryptic diversity detected in E. mayana also pose challenges for the conservation of this isopod and its fragile environment, the sandy shores.     

Swimming ability and burrowing time of two cirolanid isopods from different levels of exposed sandy beaches

Most of the macroinfauna from sandy beaches is highly mobile, emerging out of the sediment when the tide rises, and using the swash to migrate up and down the beach face or feed (searching for prey or carrion). After swash excursions, they usually burrow back into the sediment, maintaining zonation at low tide. Therefore, the different species abilities to emerge, move around and burrow under different swash climates and sediment conditions are expected to influence observed distribution patterns. Nonetheless, few attempts have been made to understand behavioral mechanisms of these organisms in moving fluids.In this study, we used a flume tunnel to investigate the orientation, swimming ability and burrowing time of two similar species of cirolanid isopods, Excirolana armata Dana and Excirolana braziliensis Richardson, under current velocities ranging from 5 to 30 cm·s⁻¹. E. armata inhabits middle levels of dissipative to intermediate beaches, while E. braziliensis is found towards the upper level of a wider range of beach states. Both species oriented countercurrent above a threshold velocity, which turned out to be significantly lower for E. armata than for E. braziliensis. E. armata proved to be a stronger swimmer as shown by the higher velocities surmounted, and the less drags experienced at the highest current velocity. E. armata also burrowed faster than E. braziliensis. Burrowing time was affected by sediment grain size and water content, but not by water flow. Once organisms managed to begin burrowing under different flow conditions, they were not affected by current velocity. Nonsaturated sand precluded burial, while coarse sand retarded it. Differences in the observed patterns of across-beach distribution may thus be the result of species-specific behavioral responses to swash climate, manifested in swimming ability, burying and orientation in directional flows.     

Natural history of the Pan-American sand beach isopod Excirolana braziliensis (Crustacea: Malacostraca)

A population of the intertidal isopod Excirolana braziliensis was followed over a 10-month period at Naos Island, Panama. Average intertidal density was 2147/m² with maximum abundance occurring in the highest level. Adults were present only in the highest tidal level. Juveniles were distributed throughout all tidal levels and comprised at least 70% of the population at all times. Mean size of individuals decreased from the highest level seaward. Reproductive females were present the year round. The number of young ranged from four to 17 per brood and was dependent on the length of the female. Adults were very tolerant of changes in temperature and salinity.     

Effects of intensive rice production practices on avian biodiversity in Southeast Asian managed wetlands

Rice fields are common throughout the agricultural landscape of Southeast Asia and sustain various bird species. These birds can provide ecosystem services, such as pest control, that improve agricultural yields whilst minimising the use of agrochemicals. This study quantified avian biodiversity in rice production landscapes during three farming stages in Peninsular Malaysia. In Malaysia, rice fields can be an important habitat for migrating birds due to Malaysia’s position on the East-Asian–Australasian Flyway. We determined bird abundance, species richness, and composition in rice field landscapes and compared these during different stages of rice growing. Wetland and terrestrial birds were counted in rice fields using the point-count method. Sixty sampling points were randomly established in three locations, from which 3447 individual birds of 46 species and 26 families were recorded. There was a significant difference in total bird abundance and species richness between the three different stages of rice growing. The growing stage supported greater bird abundance and species richness compared to the pre-harvest and post-harvest stages. Rice-growing provides temporary habitats to different bird species in this managed aquatic landscape. This implies the presence of abundant food, such as small fish and amphibians. The evidence from this study suggests that biodiversity-friendly agricultural practices should be implemented to improve habitat quality for birds in rice production landscapes.     

Some aspects of overwintering in southern England of the mosquitoes Anopheles atroparvus and Culiseta annulata (Diptera: Culicidae)

Abstract. 1. Overwintering of the mosquitoes Anopheles atroparvus van Thiel and Culiseta annulata Schrank, both of which feed on blood throughout the year, is compared.
2 The aquatic stages, males and gonoactive females of A.atroparvus disappear before winter, leaving a generation of inseminated, nulliparous, diapausing females to overwinter. Blood meals taken by diapausing females merely serve to maintain fat reserves. After re-activation in March–April, gonoactive life expectancy appears to be rather short. Aquatic development at this season is slow and eggs laid in the spring do not significantly affect adult densities until June.
3. Aquatic stages, males and gonoactive females of C.annulata are present throughout the year. This species appears to be able to overwinter without recourse to diapause, females passing the winter in alternating, or lasting, states of cold-induced quiescence or normal gonoactivity.     

Evolution of mating isolation between populations of Drosophila ananassae

Prezygotic mating isolation has been a major interest of evolutionary biologists during the past several decades because it is likely to represent one of the first stages in the transition from populations to species. Mate discrimination is one of the most commonly measured forms of prezygotic isolation and appears to be relatively common among closely related species. In some cases, it has been used as a measure to distinguish populations from subspecies, races, and sister species, yet the influences of various evolutionary mechanisms that may generate mate discrimination are largely unknown. In this study, we measured the level and pattern of mate discrimination among 18 populations of a cosmopolitan drosophilid species, Drosophila ananassae , from throughout its geographical range and its sister species, Drosophila pallidosa, which has a restricted geographical distribution in the South Pacific Islands. In addition, we measured genetic differentiation between all 18 populations using mitochondrial DNA polymorphism data. Mate discrimination varies considerably throughout the species range, being higher among populations outside the ancestral Indonesian range, and highest in the South Pacific. Our results suggest that colonization and genetic differentiation may have an influence on the evolutionary origin of mate discrimination. Our phylogeographical approach clarifies the ancestral relationships of several populations from the South Pacific that show particularly strong mate discrimination and suggests that they may be in the early stages of speciation. Furthermore, both the genetic and behavioral results cast doubt on the status of D. pallidosa as a good species.     

Review The ecology of ticks transmitting Lyme borreliosis

The main vectors of Borrelia burgdorferi sensu lato, the cause of Lyme borreliosis, are ixodid ticks of the Ixodes persulcatus species complex. These ticks, which occur throughout the northern temperate zone, have very similar life cycles and ecological requirements. All are three-host ticks, with the immature stages mainly parasitizing small to medium-sized mammals and birds and the adult females parasitizing large mammals such as deer, cattle, sheep and hares. The host-seeking stages show a distinct seasonality, which is regulated by diapause mechanisms and there appear to be major differences in this respect between the Old World and New World species. Most cases of human borreliosis are transmitted in the summer by the nymphal stages, with the exception of the Eurasian species, I. persulcatus, in which the adult females are mainly responsible. The ticks acquire the spirochaetes from a wide variety of mammals and birds but large mammals do not seem to be infective, so that t icks that feed almost exclusively on large mammals, for example in some agricultural habitats, are rarely infected. The greatest tick infection prevalences occur in deciduous woodland harbouring a diverse mix of host species and the diversity of the different genospecies of B. burgdorferi s.l. is also greatest in such habitats. There is evidence that these genospecies have different host predilections but, apart from the fact that I. persulcatus does not seem to be infected by B. burgdorferi sensu stricto, they do not seem to be adapted to different tick strains or species. © Rapid Science Ltd. 1998     

The estrous cycle in the dog: a review

Clinical, morphologic and hormonal changes throughout the estrous cycle in the dog are discussed in relation to the well established stages in this species — proestrus, estrus, metestrus and anestrus. Although most efforts in the past have been devoted to the period of reproduction, events throughout the remaining months of the cycle are presented to characterize the dog as a monocyclic animal in contrast to polycyclic species.     

Bird communities in habitats along a successional gradient: Divergent patterns of species richness,specialization and threat

The long history of human influence on northern temperate landscapes has created a mosaic of successional stages, from closed forest to open grassland. Various species thus adapted to different habitats and it is interesting to explore how these differences in species composition among particular successional stages translate into differences at the community level. For this purpose, we surveyed breeding birds in 233 patches of five different habitats covering a gradient from bare ground to forest in 29 abandoned military training sites scattered throughout the Czech Republic. Linear mixed effects modelling revealed that late-successional habitats (dense scrubland and forest) were the most species-rich, whereas early-successional stages hosted bird communities with the highest habitat specialization and threat level. These results suggest that the habitats of late-successional stages are important for the maintenance of high bird species richness, but that early-successional habitats are essential for highly specialized and threatened bird species. Given the highly adverse impacts of agricultural intensification and land abandonment on open habitats, it is necessary to promote factors creating initial successional stages suitable for specialized and threatened species.     

Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.     

A holomorph approach to xiphosuran evolution—a case study on the ontogeny of Euproops

Specimens of Euproops sp. (Xiphosura, Chelicerata) from the Carboniferous Piesberg quarry near Osnabrück, Germany, represent a relatively complete growth series of 10 stages. Based on this growth sequence, morphological changes throughout the ontogeny can be identified. The major change affects the shape of the epimera of the opisthosoma. In earlier stages, they appear very spine-like, whereas in later stages the bases of these spine-like structures become broader; the broadened bases are then successively drawn out distally. In the most mature stage known, the epimera are of trapezoidal shape and approach each other closely to form a complete flange around the thoracetron (=fused tergites of the opisthosoma). These ontogenetic changes question the taxonomic status of different species of Euproops, as the latter appear to correspond to different stages of the ontogenetic series reconstructed from the Piesberg specimens. This means that supposed separate species could, in fact, represent different growth stages of a single species. It could alternatively indicate that heterochrony (=evolutionary change of developmental timing) plays an important role in the evolution of Xiphosura. We propose a holomorph approach, i.e., reconstructing ontogenetic sequences for fossil and extant species as a sound basis for a taxonomic, phylogenetic, and evolutionary discussion of Xiphosura.     

Interspecific facilitation favors rare species establishment and reduces performance disparities among adults

Questions

A variety of mechanisms sustain diversity in natural communities as a result of ecological interactions between organisms. Competition has been studied extensively in the context of species maintenance, but facilitation is often conceptualized as simply reducing competition between functionally different species, which tends to decline throughout the plants' life span. Here we explore how interspecific facilitation may sustain diversity throughout the species' life by avoiding the extinction of locally rare species at juvenile stages and reducing performance disparities between neighbors of differing species at mature stages.

Methods

To do so, we measured whether rarer species relied more on facilitation than abundant ones in semiarid shrubland in southeast Spain. A mechanistic explanation of this relationship was subsequently tested by correlating rarity with the species' affinity to a particularly edaphic stressful environment. Finally, we assessed whether growing associated with neighbors in vegetation patches shaped by facilitation could balance performance disparities between species when they become adults.

Results

We show that facilitation (i) favors the rare species, which, in addition, tend to be those with low affinity to the stressful environment, and (ii) reduces the performance dissimilarities among plants growing associated within multispecific vegetation patches compared to plants growing alone.

Conclusions

These facilitative effects, beyond the reduction of competition between functionally similar species, might ensure positive and long-lasting effects of biotic interactions, implying a more critical role for facilitation in preserving biodiversity than previously thought.     

Potential of MALDI-TOF MS-based proteomic fingerprinting for species identification of Cnidaria across classes,species, regions and developmental stages

Morphological identification of cnidarian species can be difficult throughout all life stages due to the lack of distinct morphological characters. Moreover, in some cnidarian taxa genetic markers are not fully informative, and in these cases combinations of different markers or additional morphological verifications may be required. Proteomic fingerprinting based on MALDI-TOF mass spectra was previously shown to provide reliable species identification in different metazoans including some cnidarian taxa. For the first time, we tested the method across four cnidarian classes (Staurozoa, Scyphozoa, Anthozoa, Hydrozoa) and included different scyphozoan life-history stages (polyp, ephyra, medusa) in our dataset. Our results revealed reliable species identification based on MALDI-TOF mass spectra across all taxa with species-specific clusters for all 23 analysed species. In addition, proteomic fingerprinting was successful for distinguishing developmental stages, still by retaining a species specific signal. Furthermore, we identified the impact of different salinities in different regions (North Sea and Baltic Sea) on proteomic fingerprints to be negligible. In conclusion, the effects of environmental factors and developmental stages on proteomic fingerprints seem to be low in cnidarians. This would allow using reference libraries built up entirely of adult or cultured cnidarian specimens for the identification of their juvenile stages or specimens from different geographic regions in future biodiversity assessment studies.