Hot Rocks and Not-So-Hot Rocks on the Seashore: Patterns and Body-Size Dependent Consequences of Microclimatic Variation in Intertidal Zone Boulder Habitat.

Microclimatic variation has emerged as an important driver of many ecological and evolutionary processes. Nonetheless, fine-scale temperature data are still rare in most habitats, limiting our ability to understand the consequences of microclimatic variation under current and future conditions. We measured fine-scale thermal variation in a common, species-rich, but rarely studied habitat with respect to temperature: the airspaces under rocks on intertidal zone boulder shores. The effects of thermal variation were investigated using physiological, behavioral, and demographic responses of the porcelain crab Petrolisthes cinctipes. Habitat temperatures were measured at fine spatial and temporal resolution over 18 months, producing 424,426 temperature records. Microclimatic variation increased with increasing intertidal elevation, particularly with respect to heat extremes. However, mean temperatures were similar across the entire intertidal zone. Overheating risk for P. cinctipes increases with intertidal elevation but is size dependent, as large animals are more heat sensitive than small animals. Still, microclimatic variation high in the intertidal zone provided thermal refugia even under the warmest conditions. Size-dependent thermal responses predicted that large crabs should be rare high in the intertidal zone, which was supported by demographic data. Furthermore, simulations parameterized by our microclimate and organismal data recapitulated demographic patterns. Therefore, interactions between microclimatic variation and size-dependent thermal responses may have significant ecological repercussions that warrant greater attention.

Rocas calientes y rocas no tan calientes en la orilla del mar: patrones y consecuencias dependientes del tamaño del cuerpo de la variación microclimática en el hábitat de rocas intermareales (Hot rocks and not-so-hot rocks on the seashore: patterns and body-size dependent consequences of microclimatic variation in intertidal zone boulder habitat) La variación microclimática se ha convertido en un importante impulsor de muchos procesos ecológicos y evolutivos. No obstante, los datos de temperatura a pequeña escala aún son raros en la mayoría de los hábitats, lo que limita nuestra capacidad de comprender las consecuencias de la variación microclimática en las condiciones actuales y futuras. Medimos la variación térmica a pequeña escala en un hábitat común, rico en especies, pero rara vez estudiado con respecto a la temperatura: los espacios aéreos debajo de las rocas en las costas de rocas de zonas intermareales. Los efectos de la variación térmica se investigaron utilizando respuestas fisiológicas, conductuales y demográficas de los cangrejos de porcelana Petrolisthes cinctipes. Las temperaturas del hábitat se midieron con una resolución espacial y temporal fina durante 18 meses, produciendo 424.426 registros de temperatura. La variación microclimática aumentó con el aumento de la elevación intermareal, particularmente con respecto a los extremos de calor. Sin embargo, las temperaturas medias fueron similares en toda la zona intermareal. El riesgo de sobrecalentamiento de P. cinctipes aumenta con la elevación intermareal, pero depende del tamaño, ya que los animales grandes son más sensibles al calor que los animales pequeños. Aún así, la variación microclimática alta en la zona intermareal proporcionó refugios térmicos incluso en las condiciones más cálidas. Las respuestas térmicas dependientes del tamaño predijeron que los cangrejos grandes deberían ser raramente altos en la zona intermareal, lo que fue respaldado por datos demográficos. Además, las simulaciones parametrizadas por nuestros microclimas y datos organismales recapitularon patrones demográficos. Por lo tanto, las interacciones entre la variación microclimática y las respuestas térmicas dependientes del tamaño pueden tener importantes repercusiones ecológicas que justifican una mayor atención. Translated to Spanish by YE Jimenez (yordano_jimenez@brown.edu).

Rochas quentes e rochas não-tão-quentes à beira-mar: padrões e conseqüências dependentes do tamanho coporal para a variação microclimática no habitat de rochas da zona entremarés (Hot rocks and not-so-hot rocks on the seashore: patterns and body-size dependent consequences of microclimatic variation in intertidal zone boulder habitat) A variação microclimática emergiu como um importante causador de muitos processos ecológicos e evolutivos. No entanto, dados de temperatura em escala fina ainda são raros para a maioria dos habitats, limitando nossa capacidade de compreender as conseqüências da variação microclimática sob condições atuais e futuras. Nós medimos a variação térmica numa escala fina em um habitat comum, rico em espécies, mas raramente estudado em relação à temperatura: os espaços aéreos sob as rochas nas margens dos blocos de pedra da zona entremarés. Os efeitos da variação térmica foram investigados usando respostas fisiológicas, comportamentais e demográficas do caranguejo Petrolisthes cinctipes. As temperaturas do habitat foram medidas em resolução espacial e temporal ao longo de 18 meses, produzindo 424.426 registros de temperatura. A variação microclimática aumentou com o aumento da elevação intertidal, particularmente com relação aos extremos de calor. No entanto, as temperaturas médias foram semelhantes em toda a zona entremarés. O risco de superaquecimento de P. cinctipes aumenta com a elevação intertidal, mas é dependente do seu tamanho, já que os animais grandes são mais sensíveis ao calor do que os pequenos. Ainda assim, a variação microclimática alta na zona entremarés forneceu refúgios térmicos mesmo sob as condições mais quentes. Respostas térmicas dependentes do tamanho previam que os grandes caranguejos deveriam ser raros na zona entremarés, o que era apoiado por dados demográficos. Além disso, as simulações parametrizadas pelos nossos dados de microclima e organismal recapitularam padrões demográficos. Portanto, as interações entre a variação microclimática e as respostas térmicas dependentes do tamanho corporal podem ter repercussões ecológicas significativas que merecem maior atenção. Translated to Portuguese by G Sobral (gabisobral@gmail.com).

Overview on the European green crab Carcinus spp. (Portunidae, Decapoda), one of the most famous marine invaders and ecotoxicological models.

Green crabs (Carcinus, Portunidae) include two species native to Europe--Carcinus aestuarii (Mediterranean species) and Carcinus maenas (Atlantic species). These small shore crabs (maximal length carapace, approximately 10 cm) show rapid growth, high fecundity, and long planktonic larval stages that facilitate broad dispersion. Carcinus spp. have a high tolerance to fluctuations of environmental factors including oxygen, salinity, temperature, xenobiotic compounds, and others. Shipping of Carcinus spp. over the past centuries has resulted in its invasions of America, Asia, and Australia. Classified as one of the world's 100 worst invaders by the International Union for Conservation of Nature, Carcinus spp. are the most widely distributed intertidal crabs in the world. Their voracious predatory activity makes them strong interactors in local communities, and they are recognized as a model for invasiveness in marine systems as well as a sentinel species in ecotoxicology. This review shows an exhaustive analysis of the literature on the life cycle, diversity, physiological tolerance, genomic investigations, ecotoxicological use, historical invasion, control programs, and putative economical valorization of shore crabs.

Protein expression and genetic structure of the coral Porites lobata in an environmentally extreme Samoan back reef: does host genotype limit phenotypic plasticity?

The degree to which coral reef ecosystems will be impacted by global climate change depends on regional and local differences in corals' susceptibility and resilience to environmental stressors. Here, we present data from a reciprocal transplant experiment using the common reef building coral Porites lobata between a highly fluctuating back reef environment that reaches stressful daily extremes, and a more stable, neighbouring forereef. Protein biomarker analyses assessing physiological contributions to stress resistance showed evidence for both fixed and environmental influence on biomarker response. Fixed influences were strongest for ubiquitin-conjugated proteins with consistently higher levels found in back reef source colonies both pre and post-transplant when compared with their forereef conspecifics. Additionally, genetic comparisons of back reef and forereef populations revealed significant population structure of both the nuclear ribosomal and mitochondrial genomes of the coral host (F(ST) = 0.146 P < 0.0001, F(ST) = 0.335 P < 0.0001 for rDNA and mtDNA, respectively), whereas algal endosymbiont populations were genetically indistinguishable between the two sites. We propose that the genotype of the coral host may drive limitations to the physiological responses of these corals when faced with new environmental conditions. This result is important in understanding genotypic and environmental interactions in the coral algal symbiosis and how corals may respond to future environmental changes.

Differential gene expression during thermal stress and bleaching in the Caribbean coral Montastraea faveolata.

The declining health of coral reefs worldwide is likely to intensify in response to continued anthropogenic disturbance from coastal development, pollution, and climate change. In response to these stresses, reef-building corals may exhibit bleaching, which marks the breakdown in symbiosis between coral and zooxanthellae. Mass coral bleaching due to elevated water temperature can devastate coral reefs on a large geographical scale. In order to understand the molecular and cellular basis of bleaching in corals, we have measured gene expression changes associated with thermal stress and bleaching using a complementary DNA microarray containing 1310 genes of the Caribbean coral Montastraea faveolata. In a first experiment, we identified differentially expressed genes by comparing experimentally bleached M. faveolata fragments to control non-heat-stressed fragments. In a second experiment, we identified differentially expressed genes during a time course experiment with four time points across 9 days. Results suggest that thermal stress and bleaching in M. faveolata affect the following processes: oxidative stress, Ca(2+) homeostasis, cytoskeletal organization, cell death, calcification, metabolism, protein synthesis, heat shock protein activity, and transposon activity. These results represent the first medium-scale transcriptomic study focused on revealing the cellular foundation of thermal stress-induced coral bleaching. We postulate that oxidative stress in thermal-stressed corals causes a disruption of Ca(2+) homeostasis, which in turn leads to cytoskeletal and cell adhesion changes, decreased calcification, and the initiation of cell death via apoptosis and necrosis.

Molecular phylogeny of Eastern Pacific porcelain crabs, genera Petrolisthes and Pachycheles, based on the mtDNA 16S rDNA sequence: phylogeographic and systematic implications.

Porcelain crabs, genera Petrolisthes and Pachycheles, are diverse and abundant members of the eastern Pacific near-shore decapod crustacean community. Morphology-based taxonomic analyses of these crabs have determined groupings of affiliated species, but phylogenetic relationships remain unknown. We used sequence data from the mitochondrial 16S rRNA gene of 46 species of eastern Pacific porcelain crabs to perform phylogenetic analyses by distance and parsimony methods. Our results are used to compare the taxonomic significance of morphological and molecular characters, to examine sequence divergence rates of crab 16S rRNA genes, and to analyze the phylogeographic history of these crabs. Our phylogenetic trees indicate that the genus Petrolisthes is divided into two main clades, reflecting morphological features. One clade contains primarily tropical species, and the other contains species from throughout the eastern Pacific, as well as species in the genera Allopetrolisthes and Liopetrolisthes. Phylogenetic trees of Pachycheles suggest an antitropical distribution; north and south temperate species form one clade and tropical species form a second clade. Sequence divergence rates of the 16S rRNA gene from three pairs of geminate species can be used to date divergence times, and we discuss porcelain crab phylogeographic patterns in relation to paleogeographic events.

A comparative analysis of the evolutionary patterning and mechanistic bases of lactate dehydrogenase thermal stability in porcelain crabs, genus Petrolisthes.

The kinetic properties of orthologous homologs (orthologs) of enzymes are typically correlated with environmental temperatures in species adapted to different thermal regimes, but correlations between adaptation temperature and enzyme thermal stability are less clear. Although the thermal stability of a protein is related chiefly to its primary structure (including post-translational modification), thermal stability can also be altered by extrinsic factors present in the intracellular milieu. Here, we present a comparative analysis of the thermal stability of lactate dehydrogenase (LDH) orthologs from 22 congeneric species of porcelain crab (genera Petrolisthes and Allopetrolisthes) from a broad range of thermal habitats. Interspecific diversity of LDH stability is high: temperatures required for a 50 % loss of activity in 10 min ranged from 65 to 75.5 degrees C, corresponding to half-lives of less than 1 min to more than 3 h at 70 degrees C. Although stability is positively correlated with maximal habitat temperature in some sister taxa, phylogenetic comparative analysis incorporating all 22 species does not indicate that the interspecific diversity of LDH stability represents an adaptive response to current thermal habitats. Examination of the mechanistic bases of LDH stabilization indicates that differences in stability are related both to properties of the LDH molecule itself (intrinsic stability) and to the effects of extrinsic protein(s). Intrinsic differences were shown by the unfolding of structure during heating, as measured by circular dichroism spectroscopy. Stabilizing effects of extrinsic proteins are implied by the results of cellular fractionation experiments that removed low-molecular-mass solutes and proteins from the muscle homogenates. We conclude that the overall structural stability and functional properties of proteins can evolve independently and that in vivo protein-protein interactions can provide another means to regulate protein stability selectively.

A comparative analysis of the upper thermal tolerance limits of eastern Pacific porcelain crabs, genus Petrolisthes: influences of latitude, vertical zonation, acclimation, and phylogeny.

Marine intertidal organisms are subjected to a variety of abiotic stresses, including aerial exposure and wide ranges of temperature. Intertidal species generally have higher thermal tolerance limits than do subtidal species, and tropical species have higher thermal tolerance limits than do temperate species. The adaptive significance of upper thermal tolerance limits of intertidal organisms, however, has not been examined within a comparative context. Here, we present a comparative analysis of the adaptive significance of upper thermal tolerance limits in 20 congeneric species of porcelain crabs, genus Petrolisthes, from intertidal and subtidal habitats throughout the eastern Pacific. Upper thermal tolerance limits are positively correlated with surface water temperatures and with maximal microhabitat temperatures. Analysis of phylogenetically independent contrasts (from a phylogenetic tree on the basis of the 16s rDNA gene sequence) suggests that upper thermal tolerance limits have evolved in response to maximal microhabitat temperatures. Upper thermal tolerance limits increased during thermal acclimation at elevated temperatures, the amount of increase being greater for subtidal than for intertidal species. This result suggests that the upper thermal tolerance limits of some intertidal species may be near current habitat temperature maxima, and global warming thus may affect the distribution limits of intertidal species to a greater extent than for subtidal species.

Evolutionary history and adaptive significance of respiratory structures on the legs of intertidal porcelain crabs, genus Petrolisthes.

Semiterrestrial and terrestrial crabs have evolved multiple strategies for aerial respiration. An uncommon strategy for aerial respiration is seen in porcelain crabs, genus Petrolisthes, where decalcified areas on the meral segments of the walking legs are used as respiratory structures. Here, the evolutionary history and adaptive significance of these structures in porcelain crabs is examined. Interspecific variation in leg membrane size is from 0% to 60% of the surface area of the meral segment. Leg membrane relative size is positively correlated with body size across species but not within one species, Petrolisthes cinctipes. Phylogenetic analyses suggest that leg membranes are ancestral to one of two eastern Pacific Petrolisthes clades. Comparative analyses using phylogenetic independent contrasts indicate a relationship between leg membrane relative size and body size that is phylogenetically independent. In large-bodied intertidal species, whole-animal lactate accumulation during aerial incubation is 200%-300% higher when the leg membranes are obscured, indicating that the leg membranes are functional respiratory structures in these species. Thus, it is possible that leg membranes have facilitated the evolution of larger body sizes by providing additional respiratory surfaces to accommodate the associated higher metabolic demands.