Intra-island patterns of cryptic diversity within an oceanic island: Insights from the differentiation of Schizidium Verhoeff, 1901 (Oniscidea, Armadillidiidae) within Cyprus, with descriptions of two new species.

Cyprus is a Mediterranean island of volcanic origin isolated for at least 5.3 Myr from surrounding continental areas. The present study focuses on the diversification of the isopod genus Schizidium within the island, including also specimens from surrounding continental areas. The genus Schizidium sensu lato is probably non monophyletic, comprising 26 species distributed from Greece to Iran. Up to date the only representative of the genus reported from Cyprus was Schizidium fissum. Aiming to investigate the patterns of genetic diversity within the focal island, to evaluate the morphology-based taxonomy of the species in the genus, and at the same time to explore phylogenetic relationships with mainland populations, we applied genome-wide ddRADseq as well as Sanger sequencing targeting three mitochondrial (16S, COI and 12S) and the nuclear NaK loci. Results of phylogenetic analyses support the existence of two distinct epigean Schizidium clades with well-defined geographic boundaries that conform to the known paleogeography of Cyprus, plus one endogean clade with restricted distribution within the island. Genetic data and morphology corroborate the assignment of this latter endogean clade to a new species, Schizidium myrrae n. sp. The two epigean clades are also considered as distinct species, one corresponding to the known S. fissum at the eastern part of the island (Pentadaktylos massif) and the other to the newly described S. christosi n. sp. distributed along the western part of the island (Troodos massif). Even though detailed examination of many specimens could not retrieve any morphological differences among representatives of these two clades, clado-chronological analysis indicates a long isolation between them, estimated at âˆ¼ 9 Mya, as well as the sharing of a common ancestor with S. tiberianum from Israel at âˆ¼ 15 Mya. Hence, we can consider these epigean Schizidium species as one more case of cryptic diversity on Cyprus, exhibiting similar patterns with the recently described case in the genus Armadillo.

Diversification within an oceanic Mediterranean island: Insights from a terrestrial isopod.

Understanding intra-island patterns of evolutionary divergence, including cases of cryptic diversity, is a crucial step towards deciphering speciation processes. Cyprus is an oceanic island isolated for at least 5.3 Mya from surrounding continental regions, while it remains unclear whether it was ever connected to the mainland, even during the Messinian Salinity Crisis. The terrestrial isopod species Armadillo officinalis, that is widespread across the Mediterranean, offers the opportunity to explore intra-island divergence patterns that might exhibit geographical structure related also to the region's known paleogeography. Genome-wide ddRADseq, as well as Sanger sequencing for four mitochondrial and three nuclear loci data were generated for this purpose. In total, 71 populations from Cyprus, neighbouring continental sites, i.e., Israel, Lebanon and Turkey, and other Mediterranean regions, i.e. Greece, Italy, and Tunisia, were included in the analysis. Phylogenetic reconstructions and population structure analyses support the existence of at least six genetically discrete groups across the study area. Five of these distinct genetic clades occur on Cyprus, four of which are endemic to the island and one is widely distributed along the circum-Mediterranean countries. The sixth clade is distributed in Israel. The closest evolutionary relationship of endemic Cypriot populations is with those from Israel, while the evolutionary clade that is present in countries all around the Mediterranean is very shallow. Cladochronological analyses date the origin of the species on the island at ∼6 Mya. Estimated f4 and D statistics as well as FST values indicate the genetic isolation between the populations sampled from Cyprus and surrounding continental areas, while there is evident gene flow among populations within the island. Species delimitation and population genetic metrics support the existence of three distinct taxonomic units across the study area, two of which occur on the island and correspond to the endemic clade and the widespread circum-Mediterranean one, respectively, while the third corresponds to Israel's clade. The islands' paleogeographic history and recent human activities seem to have shaped current patterns of genetic diversity in this group of species.

Are terrestrial isopods able to use stridulation and vibrational communication as forms of intra and interspecific signaling and defense strategies as insects do? A preliminary study in Armadillo officinalis.

The capability of producing sounds and vibrations is well known in insects and is thought to be a form of intra- and interspecific communication. Sounds and vibrations are used and modulated for several aims such as interacting with conspecifics, getting information from the environment, and defending against predators. This phenomenon is less known but also present in other arthropods, including a few roller-type terrestrial isopods. In this study, we used a Y-shape test apparatus to investigate the behavior of adult individuals of Armadillo officinalis Duméril, 1816 (Crustacea: Isopoda: Oniscidea) when exposed to two particular vibrational stimuli, namely species-specific stridulations and non-specific substrate-borne vibrations. Our results showed that adults of A. officinalis significantly react to the presence of both types of vibrational stimuli, by moving away from the vibrational source as if they experienced these vibrations as a sign of danger or disturbance. A. officinalis can produce stridulations only when it rolls into a ball during the so-called conglobation, a possible defense mechanism against predators. Stridulation might thus be a secondary form of defense used during conglobation to deter a predator following contact with it and might be experienced as an alert by conspecifics nearby. The high sensitivity to non-specific substrate-borne vibrations might provide A. officinalis with the possibility to anticipate dangers and adverse conditions, giving it a better chance of survival.

Tail autotomy affects bipedalism but not sprint performance in a cursorial Mediterranean lizard.

Running is essential in all terrestrial animals mainly for finding food and mates and escaping from predators. Lizards employ running in all their everyday functions, among which defense stands out. Besides flight, tail autotomy is another very common antipredatory strategy within most lizard families. The impact of tail loss to sprint performance seems to be species dependent. In some lizard species, tail shedding reduces sprint speed, in other species, increases it, and, in a few species, speed is not affected at all. Here, we aimed to clarify the effect of tail autotomy on the sprint performance of a cursorial lizard with particular adaptations for running, such as bipedalism and spike-like protruding scales (fringes) on the toepads that allow high speed on sandy substrates. We hypothesized that individuals that performed bipedalism, and have more and larger fringes, would achieve higher sprint performance. We also anticipated that tail shedding would affect sprint speed (though we were not able to define in what way because of the unpredictable effects that tail loss has on different species). According to our results, individuals that ran bipedally were faster; limb length and fringe size had limited effects on sprint performance whereas tail autotomy affected quadrupedal running only in females. Nonetheless, tail loss significantly affected bipedalism: the ability for running on hindlimbs was completely lost in all adult individuals and in 72.3% of juveniles.

Seasonal Variation in the Thermoregulation Pattern of an Insular Agamid Lizard.

Ectotherms, including lizards, rely on behavioral thermoregulation to maintain their body temperature within an optimal range. The benign climate of islands is expected to favor the thermoregulation efficiency of reptiles throughout their activity period. In this study, we investigated the seasonal variation in thermoregulation in an insular population of the roughtail rock agama (Laudakia stellio) on Naxos Island, Greece. We measured body, operative, and preferred temperatures across three seasons (spring, summer, and autumn), and we evaluated the effectiveness of thermoregulation, using the Hertz index (E). Our results revealed that the effectiveness of thermoregulation was significantly influenced by seasonality. E was quite high in summer (0.97) and spring (0.92), and lowest in autumn (0.81). Accordingly, the quality of the thermal environment was significantly low during autumn, and maximum during summer. However, despite the environmental temperature fluctuations, lizards exhibited remarkable stability in body temperatures. They also adjusted their preferred temperatures seasonally and doubled the thermal niche breadth they occupied during summer, thus enhancing thermoregulation efficiency. Whether or not these adjustments are plastic or fixed local adaptations remains to be explored in further research across multiple years and seasons, including additional insular populations.

The genus Armadillo Latreille, 1802 (Oniscidea, Armadillidae) from Cyprus, with descriptions of two new species.

The present work aims to morphologically characterize the species of Armadillo from Cyprus. Moreover, two new species of the genus are described, A. konstantinoui sp. nov. from several places along the island, and A. karametae sp. nov. from Pafos region. The circum-Mediterranean species A. officinalis is also redescribed.

An all-inclusive approach: A universal protocol for the successful amplification of four genetic loci of all Onscidea.

Accounting more than 3,700 described species, Oniscidea is the largest and at the same time the only terrestrial isopod suborder inhabiting almost all terrestrial biomes. Despite the great effort dedicated on describing taxonomic diversity of Oniscidea, mainly employing morphology, there is still a considerable number of species/genera of uncertain generic/familiar assignment. Based on different morphological characters, alternative evolutionary relationships have been proposed to describe the diversity of Oniscidea at different phylogenetic levels. Accumulating morphological and genetic data are repeatedly challenging the monophyly of established taxa, undermining the validity of several morphological characters traditionally used in terrestrial isopod taxonomy, leading to often revisions of the current taxonomy of the Oniscidea . The use of genetic data facilitates the efforts to reconstruct the complex evolutionary history of the focal group by providing important data for the identification, delimitation, and description of species. The proposed protocol with universal PCR conditions and primers was used to successfully amplify COI, 16S, 28S and NAK loci in diverse Oniscidea taxa. The application of this protocol is anticipated to facilitate the generation of new genetic data and hence promote scientific research in Isopoda taxonomy, evolution, ecology, and other related fields.

Island species richness increases with habitat diversity.

Species richness is commonly thought to increase with habitat diversity. However, a recent theoretical model aiming to unify niche and island biogeography theories predicted a hump-shaped relationship between richness and habitat diversity. Given the contradiction between model results and previous knowledge, we examine whether the relationship between species richness and habitat diversity is consistently monotonically increasing and under which circumstances, if at all, such relationships could be hump shaped. We review the empirical evidence about the shape of such relationships and show that species richness on islands usually increases with habitat diversity and that it never decreases. We also critically examine the assumptions of the theoretical model and modify them to incorporate a less restrictive definition of niche width. The modified assumptions lead to simulations that better capture real patterns, using either simple parameters or observed distributions of niche breadth. Further work is needed to incorporate ecological interactions and metacommunity dynamics if the aim is to merge niche and island biogeography theories in a realistic modeling framework.

Genetic evidence against monophyly of Oniscidea implies a need to revise scenarios for the origin of terrestrial isopods.

Among the few crustacean taxa that managed to inhabit terrestrial environments, Oniscidea includes the most successful colonizers in terms of species richness and abundance. However, neither morphological traits nor molecular markers have definitively resolved phylogenetic relationships among major Oniscidea clades or established the monophyly of the taxon. Herein, we employed the highly conserved, nuclear protein-coding genes Sodium-Potassium Pump (NAK) and Phosphoenolpyruvate Carboxykinase (PEPCK), along with the traditionally used 18 s and 28 s ribosomal RNA genes, in an attempt to clarify these questions. Our dataset included sequences representing all major Oniscidea clades and closely related aquatic taxa, as suggested by previous studies. We applied Bayesian Inference and Maximum Likelihood methods and produced a robust and fully resolved phylogenetic tree that offers strong evidence against the monophyly of Oniscidea. The amphibious genus Ligia appears to be more closely related to representatives of marine suborders, while the phylogenetic pattern of the remaining Oniscidea implies a complex history of the transition from the marine environment to land. With the exception of the basal clade, all other established major clades have been recovered as monophyletic, even though relationships within these clades call for a revised interpretation of morphological characters used in terrestrial isopod taxonomy.

Promoting evolution: the brand new Hellenic Evolutionary Society (HEVOS).

Herein we present the recently founded Hellenic Evolutionary Society (HEVOS) that has been recently instituted to promote evolution and scientific thinking among the Greek-speaking public. HEVOS is a timely initiative, given the low levels of acceptance of evolution by Greek society and the almost complete lack of evolution teaching in primary and secondary education in Greece. Herein, the main aims of the Society are presented.

Response of the temperate scleractinian coral Cladocora caespitosa to high temperature and long-term nutrient enrichment.

Anthropogenic nutrient enrichment and increased seawater temperatures are responsible for coral reef decline. In particular, they disrupt the relationship between corals and their dinoflagellate symbionts (bleaching). However, some coral species can afford either high temperatures or nutrient enrichment and their study can bring new insights into how corals acclimate or adapt to stressors. Here, we focused on the role of the nutrient history in influencing the response of the Mediterranean scleractinian coral Cladocora caespitosa to thermal stress. Colonies living naturally in nutrient-poor (<0.5 µM nitrogen, <0.2 µM phosphorus, LN) and nutrient-rich (ca. 10-20 µM nitrogen, 0.4 µM phosphorus, HN) locations were sampled, maintained under the right nutrient conditions, and exposed to a temperature increase from 17 °C to 24 °C and 29 °C. While both HN and LN colonies decreased their concentrations of symbionts and/or photosynthetic pigments, HN colonies were able to maintain significant higher rates of net and gross photosynthesis at 24 °C compared to LN colonies. In addition, while there was no change in protein concentration in HN corals during the experiment, proteins continuously decreased in LN corals with increased temperature. These results are important in that they show that nutrient history can influence the response of scleractinian corals to thermal stress. Further investigations of under-studied coral groups are thus required in the future to understand the processes leading to coral resistance to environmental perturbations.

Isopod distribution and climate change.

The unique properties of terrestrial isopods regarding responses to limiting factors such as drought and temperature have led to interesting distributional patterns along climatic and other environmental gradients at both species and community level. This paper will focus on the exploration of isopod distributions in evaluating climate change effects on biodiversity at different scales, geographical regions, and environments, in view of isopods' tolerances to environmental factors, mostly humidity and temperature. Isopod distribution is tightly connected to available habitats and habitat features at a fine spatial scale, even though different species may exhibit a variety of responses to environmental heterogeneity, reflecting the large interspecific variation within the group. Furthermore, isopod distributions show some notable deviations from common global patterns, mainly as a result of their ecological features and evolutionary origins. Responses to human disturbance are not always traceable, but a trend towards community homogenisation is often found under strong global urbanisation processes. In general, even though it is still not clear how predicted climate change will affect isopod distribution, there is evidence that mixed effects are to be expected, depending on the region under study. We still lack robust and extensive analyses of isopod distributions at different scales and at different biomes, as well as applications of distribution models that might help evaluate future trends.

Functional response and multiple predator effects of two generalist predators preying on Tuta absoluta eggs.

BACKROUND: Interactions among invertebrate predators could affect a pest suppression. The hemipteran species Macrolophus pygmaeus (Rambur) and Nesidiocoris tenuis (Reuter) (Hemiptera: Miridae) are natural enemies of several pests in agroecosystems and coexist in tomato crops in Mediterranean countries. By using the multiplicative risk model (MRM) and the substitutive model, the multiple predator effects (MPEs) on prey suppression were calculated when two individuals of the predators foraged at the same densities on South American tomato pinworm, Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae), eggs. RESULTS: Egg consumption increased with increasing egg density and the two predators exhibited a type III functional response. Predation rates were strongly affected by prey density. Using the MRM, we found risk reduction at intraspecific treatments at high prey density. Applying the substitutive model, we detect risk enhancement at interspecific treatments at high egg density. CONCLUSIONS: At low prey densities, most of the interactions were independent, whereas at high densities most interactions were not independent and resulted in prey risk reduction, indicating antagonism between the individuals involved. We also showed that N. tenuis is a more competitive predator species for T. absoluta eggs than M. pygmaeus; however, combination of the two predator species will lead to better pest suppression at high T. absoluta population densities. © 2017 Society of Chemical Industry.

A molecular phylogeny of Porcellionidae (Isopoda, Oniscidea) reveals inconsistencies with present taxonomy.

Porcellionidae is one of the richest families of Oniscidea globally distributed but we still lack a comprehensive and robust phylogeny of the taxa that are assigned to it. Employing five genetic markers (two mitochondrial and three nuclear) we inferred phylogenetic relationships among the majority of Porcellionidae genera. Phylogenetic analyses conducted via Maximum Likelihood and Bayesian Inference resulted in similar tree topologies. The mtDNA genes cytochrome oxidase I (COI) and 16s rRNA (16s) were used for clade dating using previously published mutation rates. Our results provide evidence against the monophyly of both Porcellionidae and the largest genus of the family Porcellio. These results are compared to previous published work based on morphological evidence. The genera Leptotrichus and Brevurus are not grouped with the rest of Porcellionidae whereas Agnaridae are grouped with part of Porcellionidae. Armadillidium and Schizidium (Armadillidiidae) occupy a basal position on the phylogenetic tree. Even though the African genera Tura and Uramba (distributed in East Africa) are grouped together there is no general geographical pattern in other sub-clades. Additional taxonomic issues that arise in this work such as the assignment of the recently described genus Levantoniscus, arealso discussed. The status of Porcellionidae should be further revised and morphological characters traditionally used in Oniscidea taxonomy should be reconsidered in view of molecular evidence. The origin of the monophyletic clade within Porcellionidae as indicated in the present work is dated back to the Oligocene (~32 mya).

Everything is similar, everything is different! Trichorhina (Oniscidea, Platyarthridae) from Brazilian caves, with descriptions of 11 new species / Tudo é parecido, tudo é diferente! Trichorhina (Oniscidea, Platyarthridae) de cavernas brasileiras, com descrições de 11 novas espécies

Abstract The present work describes 11 new species of the genus Trichorhina from Brazilian caves in the states of Bahia, Mato Grosso do Sul, Minas Gerais, Pará, and São Paulo. Moreover, the distribution of T. tomentosa has been expanded to include the states of Mato Grosso and Minas Gerais. Additionally, a distribution map and a key for all Brazilian species of Trichorhina, including epigean ones, are provided.

Resumo O presente trabalho descreve 11 novas espécies do gênero Trichorhina de cavernas brasileiras nos estados da Bahia, Mato Grosso do Sul, Minas Gerais, Pará e São Paulo. Além disso, a distribuição de T. tomentosa foi ampliada para incluir os estados de Mato Grosso e Minas Gerais. Adicionalmente, são fornecidos um mapa de distribuição e uma chave para todas as espécies brasileiras de Trichorhina, incluindo as epígeas.

The Aegean archipelago: a natural laboratory of evolution, ecology and civilisations.

The Aegean archipelago, comprising numerous islands and islets with great heterogeneity in topographic, geological, historical and environmental properties, offers an ideal natural laboratory for ecological and evolutionary research, and has been the stage for a very long interaction between human civilizations and local ecosystems. This work presents insights that have been gained from past and current relevant research in the area, highlighting also the importance of the Aegean archipelago as a useful model to address many major questions in biogeography, ecology and evolutionary processes. Among the most interesting findings from such studies concern the role of habitat heterogeneity as the most important determinant of species richness, the development of a new model (Choros) for the species-area-habitats relationship, the mechanistic aspects of the Small Island Effect, the very high rates of species turnover, the lack of a role for interspecific competition in shaping species co-occurrence patterns in most cases, the importance of non adaptive radiation in diversification of several taxa, the insights into the relative roles of vicariance and dispersal in speciation, the understanding of the interplay between human presence and the establishment of exotic species and extinction of indigenous biotas. Concluding, the Aegean archipelago is an ideal stage for research in evolution, ecology and biogeography, and has the potential to become a model study area at a global level, especially for land-bridge, continental islands.

Patterns of taxonomic diversity among terrestrial isopods.

The publication of the world catalog of terrestrial isopods some ten years ago by Schmalfuss has facilitated research on isopod diversity patterns at a global scale. Furthermore, even though we still lack a comprehensive and robust phylogeny of Oniscidea, we do have some useful approaches to phylogenetic relationships among major clades which can offer additional insights into isopod evolutionary dynamics. Taxonomic diversity is one of many approaches to biodiversity and, despite its sensitiveness to biases in taxonomic practice, has proved useful in exploring diversification dynamics of various taxa. In the present work, we attempt an analysis of taxonomic diversity patterns among Oniscidea based on an updated world list of species containing 3,710 species belonging to 527 genera and 37 families (data till April 2014). The analysis explores species diversity at the genus and family level, as well as the relationships between species per genera, species per families, and genera per families. In addition, we consider the structure of isopod taxonomic system under the fractal perspective that has been proposed as a measure of a taxon's diversification. Finally, we check whether there is any phylogenetic signal behind taxonomic diversity patterns. The results can be useful in a more detailed elaboration of Oniscidea systematics.

A new genus of Trachelipodidae Strouhal, 1953 (Crustacea, Isopoda, Oniscidea) from the eastern Mediterranean.

Levantoniscus n. gen. is erected for two new species from Cyprus: Levantoniscus bicostulatus n. sp. and Levantoniscus makrisi n. sp. Levantoniscus wahrmani (Strouhal, 1968) n. comb. from Israel and southern Turkey is transferred from the genus Bathytropa Budde-Lund, 1885 and family Bathytropidae. The new genus is included in the family Trachelipodidae and is characterized by distinct dorsal ornamentation, interlocking pleopods and uncovered pleopodal lungs which are located in invaginations on pleopod 3-5 exopodites.