Isopods on isopods: integrative taxonomy of Cabiropidae (Isopoda: Epicaridea: Cryptoniscoidea) parasitic on anthuroid isopods, with descriptions of a new genus and three new species from Japan.

Species in the parasitic isopod family Cabiropidae are known to utilise various isopods as hosts but there are currently no records of members parasitising anthuroid hosts. We describe Anthuroniscus gen. nov. for three new cabiropid species, Anthuroniscus shimomurai sp. nov. , Anthuroniscus dentatus sp. nov. and Anthuroniscus latus sp. nov. , all of which are parasitic on anthuroid isopods. Anthuroniscus gen. nov. differs from the other 14 cabiropid genera and 10 genera treated as family incertae sedis in females having an elongate, dorsally compressed, posteriorly tapering body with six pairs of lateral bulges; and cryptoniscus larvae in the following combination of characters: (1) eyes lacking, (2) antennular article 1 with eight teeth on the posterior margin, (3) uropodal exopod and endopod rectangular rather than tapering, and endopod longer than exopod, and (4) pleotelson trapezoidal, 2× as wide as long. Anthuroniscus shimomurai sp. nov. was parasitic on Mesanthura sp. from Kaichu Doro, Uruma, Okinawa, south-western Japan; A. dentatus sp. nov. on Accalathura sp. from Irabu Island, Miyako Islands, Okinawa; and A. latus sp. nov. on Colanthura nigra from Kanagawa, central Japan. In pairwise comparisons, the three new species showed p -distances of 0.6-1.3% for the 18S rRNA gene (1440 positions); and A. shimomurai sp. nov. and A. latus sp. nov. showed a p -distance of 36.2% for the 16S rRNA gene (412 positions). In an 18S -based maximum-likelihood tree, an Anthuroniscus gen. nov. clade was the sister group to Cryptoniscoidea sp., parasitic on an ostracod species. This is the first study reporting Cabiropidae from Japan and anthuroids as hosts for Cryptoniscoidea. ZooBank: urn:lsid:zoobank.org:pub:2EE042E2-AE48-4B87-B495-8436462146B9.

Description of Sandythoa gen. nov., a fish parasitic branchial cymothoid (Crustacea: Isopoda: Cymothoidae) from the Indian Ocean, with five species including one new species.

The comparative analysis of records of Elthusa samariscii (Shiino, 1951) from Japan and India, alongside corresponding illustrations, indicates that the records of E. samariscii from Samaris cristatus Gray in India represent a distinct and previously undescribed species. This study introduces Sandythoa tiranga gen. and sp. nov., providing comprehensive descriptions of various lifecycle stages, including the female, male, transitional, premanca, and manca larvae. The following combinations of characters identify the genus: cephalon anterior margin with acute rostrum; pleonite 1 is distinctly narrow, not extending laterally; presence of a narrow gap between pleonites; antenna with more than 10 articles; maxilliped with oostegital lobe. Sandythoa tiranga sp. nov. is specifically identified along the southwest coast of India. Furthermore, we propose transferring the following species from Elthusa: Sandythoa arnoglossi (Trilles and Justine 2006) comb. nov.; Sandythoa parabothi (Trilles and Justine, 2004) comb. nov.; Sandythoa samariscii (Shiino, 1951) comb. nov.; Sandythoa moritakii (Saito and Yamauchi, 2016) comb. nov. A revised key to the global marine branchial cymothoid genera is provided.

Finding Nemo! Description of a New Species of Branchial Fish Parasitic Cymothoid, Elthusa Schioedte and Meinert, 1884 (Crustacea: Isopoda), infesting Callionymus filamentosus Valenciennes, 1837 from the Northern Indian Ocean.

PURPOSE: Specimens of Elthusa from trashfish of the Indian SW coast were described as a new parasitic cymothoid, Elthusa nemo sp. nov., (Crustacea: Isopoda). Later, the branchial cavity of the Blotchfin dragonet Callionymus filamentosus Valenciennes, 1837 were documented as its micro- and macro-habitat (host). METHODS: Fresh/live specimens of unidentified cymothoid samples were collected from the trash fishes obtained from Neendakara (08°30.0' N 76°53.30' E) fish landing centre, Kollam district, Kerala state, southwest coast of India. After a long search for the host species, we recovered the isopod from the branchial cavity of the deep-sea fish Callionymus filamentosus Valenciennes, 1837 (Callionymiformes: Callionymidae). The new species is described and illustrated based on ovigerous females. RESULTS: Elthusa nemo sp. nov., has the following sets of combinations of characters: body slightly twisted, elongated, dorsal surfaces smooth, nearly twice as long as greatest width; pleon short, ~ 14% body length, 0.8 times as wide as pereon maximum width; widest at pleonite 2 and narrowest at pleonite 1; only pleonite 1, laterally overlapped by pereonite 7 posterolateral expansion and coxa 7; presence of appendix masculina on pleopod 2; uropods 0.8 times as the length of pleotelson; antenna with two plumose setae on article 4. CONCLUSION: Elthusa nemo sp. nov., is the sixth species of the genus known from Indian waters.

Redescription and molecular characterization of Mothocya parvostis Bruce, 1986 (Crustacea: Isopoda: Cymothoidae) parasitic on Japanese halfbeak, Hyporhamphus sajori (Temminck & Schlegel, 1846) (Hemiramphidae) with Mothocya sajori Bruce, 1986 placed into synonymy.

Two species of Mothocya have previously been recorded from Hyporhamphus sajori: M. parvostis Bruce, 1986 and M. sajori Bruce, 1986. Mothocya parvostis is re-described based on the ovigerous female type and additional materials collected from the host from in and around the type locality. Morphological re-examination of fresh specimens and the type materials together with genetic data show that the M. sajori and M. parvostis are the same species, differing primarily in size, therefore we have placed Mothocya sajori Bruce, 1986 into a junior synonym of Mothocya parvostis Bruce, 1986. Mothocya parvostis is characterized by the following combinations of characters: 1) body slightly to moderately twisted to one side; 2) pereonite 7 posterior margin moderately to deeply recessed; 3) uropodal rami extending to pleotelson posterior margin; and 4) uropod rami bluntly rounded, exopod 1.5 times as long as peduncle. The differences of four morphological features for M. parvostis and M. sajori was quantified. Furthermore, a total of 635 isopods infesting H. sajori were collected from all over Japan to conduct quantitative morphological and molecular sequence analyses (mitochondrial cytochrome c oxidase subunit I and 16S rRNA). Although the four quantitative features did not overlap between the two species in type specimens, all quantitative morphological values of newly collected specimens in this study did not display a bimodal distribution. In addition, our molecular analyses found only a single clade for our newly collected specimens in neighbor-joining tree.

Mothocya kaorui n. sp. (Crustacea: Isopoda: Cymothoidae), a fish-parasitic isopod with unique antennules from the Izu Islands, Japan.

A new species of fish-parasitic isopod in the family Cymothoidae is described from the Izu Islands, Japan. Mothocya kaorui n. sp. is reported from the gill cavities of the keeled needlefish, Platybelone argalus platyura (Bennett). Despite its unique morphological characters, such as completely article-fused antennules, mitochondrial DNA analysis indicated that it belongs to Mothocya Costa. The new species is clearly distinguished from all other species of Mothocya by having completely fused, stout antennules and partially fused, slender antennae; maxilla mesial lobe with 3 or 4 recurved robust setae, lateral lobe with 4-6 recurved robust setae; maxilliped with 5-8 robust setae on article 3; coxae 2 and 3 wide; black subtriangular pleotelson; and black uropods.

New species of Sargassum-boring Limnoria Leach, 1814 (Crustacea, Isopoda, Limnoriidae) from Japan.

A new marine Sargassum-boring species of Limnoria (Limnoriidae) is described on the basis of specimens obtained at Kamogawa-shi, Chiba Prefecture, Japan. Limnoria aspera sp. nov. shares a reduced mandibular palp to a seta, algal-feeding, and the clavate shaped epipod of the maxilliped with the other species of non-mandibular-palp group. L. aspera sp. nov. differs morphologically from the congeneric species by secondary unguis of pereopods and unique carinae of pleonite 5 and pleotelson. We describe the sequences of the mitochondrial COI gene and the nuclear 28S rDNA gene. L. aspera sp. nov. differs by 14.218.0% in p-distance based on COI sequences from other Japanese species, L. furca and L. nagatai.

A new genus and species of groundwater isopod of the family Protojaniridae (Isopoda: Asellota: Gnathostenetroidoidea) from southern Chile.

A new asellotan isopod of the family Protojaniridae Fresi, Idato Scipione, 1980 is described from freshwater springs in the Osorno province, Los Lagos region, southern Chile. Wiyufiloides osornoensis gen. sp. n. is the third South American protojanirid species and the first known groundwater isopod in Chile. The new genus and species is principally characterized by the presence of a vestigial antennal scale, a strongly subchelate pereiopod I and the absence of an apical lobe on the protopod of pleopod II. The new taxon is described in detail and figures are given.

Six new epigean species of Caecidotea/ (Isopoda: Asellidae) distributed along the Trans-Mexican Volcanic Belt in Central Mexico.

Six new epigean freshwater species of the genus Caecidotea in Mexico are described. These species were collected in waterbodies located along the Trans-Mexican Volcanic Belt (TMVB) and the eastern slope of the country: Caecidotea buzwilsoni sp. nov., Caecidotea chicoensis sp. nov., Caecidotea alvarezi sp. nov., Caecidotea mintzita sp. nov., Caecidotea zacapuensis sp. nov., and one additional species Caecidotea villalobosi sp. nov. is described from the Papaloapan basin at the Atlantic Slope of Mexico. The taxonomic treatment for each species includes a detailed description, figures of characters of diagnostic importance and comments. We also provide an updated map of Caecidotea records in Mexico. This work substantially increases the number of epigean Caecidotea known to occur in Mexico to 11 species, a new total of 99 species described to North America.

Two new species and a new record of Bopyrinae (Isopoda: Bopyridae) infesting Alpheidae and Hippolytidae, with comments on the genus Bopyrina Kossmann, 1881.

Two new bopyrids, Bopyrinina articulata n. sp. and Bopyrinella asymmetrica n. sp. are described from French Polynesia, the Red Sea, and the Philippines. Bopyrinina articulata n.sp. infests Salmoneus cf. gracilipes Miya, and is distinguishable from other species of this genus by the structure of the maxilliped, pleopods and oostegite 1. Bopyrinella asymmetrica n. sp. is most similar to Bopyrinella albida Shiino, 1958, but females differ from that species because all pereomeres on the short side have round dorsolateral bosses and its pleotelson is greatly distorted. Bopyrina ocellata (Czerniavsky, 1868) is newly recorded from the Western Atlantic, from Florida, USA. Review of the species of Bopyrina revealed that B. choprai Nierstrasz & Brender à Brandis, 1929 and B. sewelli Chopra, 1930 are intermediate in morphology between Bopyrina and Schizobopyrina, the taxonomic assignment of these two species needs further evaluation. Keys to Bopyrinella Nierstrasz & Brender à Brandis, 1925 and Bopyrina Kossmann, 1881 are presented. Hosts and distributions of the seven species of Bopyrinella are summarized.

More than one way to smell ashore - Evolution of the olfactory pathway in terrestrial malacostracan crustaceans.

Crustaceans provide a fascinating opportunity for studying adaptations to a terrestrial lifestyle because within this group, the conquest of land has occurred at least ten times convergently. The evolutionary transition from water to land demands various morphological and physiological adaptations of tissues and organs including the sensory and nervous system. In this review, we aim to compare the brain architecture between selected terrestrial and closely related marine representatives of the crustacean taxa Amphipoda, Isopoda, Brachyura, and Anomala with an emphasis on the elements of the olfactory pathway including receptor molecules. Our comparison of neuroanatomical structures between terrestrial members and their close aquatic relatives suggests that during the convergent evolution of terrestrial life-styles, the elements of the olfactory pathway were subject to different morphological transformations. In terrestrial anomalans (Coenobitidae), the elements of the primary olfactory pathway (antennules and olfactory lobes) are in general considerably enlarged whereas they are smaller in terrestrial brachyurans compared to their aquatic relatives. Studies on the repertoire of receptor molecules in Coenobitidae do not point to specific terrestrial adaptations but suggest that perireceptor events - processes in the receptor environment before the stimuli bind - may play an important role for aerial olfaction in this group. In terrestrial members of amphipods (Amphipoda: Talitridae) as well as of isopods (Isopoda: Oniscidea), however, the antennules and olfactory sensilla (aesthetascs) are largely reduced and miniaturized. Consequently, their primary olfactory processing centers are suggested to have been lost during the evolution of a life on land. Nevertheless, in terrestrial Peracarida, the (second) antennae as well as their associated tritocerebral processing structures are presumed to compensate for this loss or rather considerable reduction of the (deutocerebral) primary olfactory pathway. We conclude that after the evolutionary transition from water to land, it is not trivial for arthropods to establish aerial olfaction. If we consider insects as an ingroup of Crustacea, then the Coenobitidae and Insecta may be seen as the most successful crustacean representatives in this respect.

First report of Livoneca guianensis (Isopoda: Cymothoidae) in Leporinus fasciatus (Pisces: Anostomidae) in Brazil.

This study aimed to report the first record of the occurrence of the parasite Livoneca guianensis in Leporinus fasciatus in Brazil. In September 2019, during surveys of the parasites in 12 specimens of L. fasciatus from the lower Jari River basin, in Amapá State, Brazil, one specimen of L. guianensis was found in the tongue of one host. The prevalence of L. guianensis was 8.3%, and it was observed that this parasite has a geographic distribution restricted to South America, parasitizing fish species from Guyana and Brazil. This study contributed to the first report on the occurrence of L. guianensis for L. fasciatus in Brazil. In addition, increased the knowledge on the geographic distribution of this parasite with morphometric data, which are scarce for this Cymothoidae.

Infestation of the Parasitic Isopod Mothocya parvostis on Juveniles of the Black Sea Bream Acanthopagrus schlegelii as an Optional Intermediate Host in Hiroshima Bay.

In Hiroshima Bay, parasitic isopods of the genus Mothocya infest the black sea bream Acanthopagrus schlegelii (Bleeker, 1854) and the Japanese halfbeak Hyporhamphus sajori (Temminck and Schlegel, 1846), two fish species that are abundant and commercially important in the Seto Inland Sea of Japan. Immature and mature Mothocya individuals can infect both juveniles and adults of H. sajori, while immature Mothocya are known to parasitize juveniles of A. schlegelii; i.e., no Mothocya parasites are found in adult A. schlegelii. The identification of the immature Mothocya parasitizing juveniles of A. schlegelii remains uncertain, because Mothocya species are morphologically identifiable only based on adult females. Also, the biological/ecological relationship between the hosts and parasites has not been studied. Here, we identified the parasites on A. schlegelii as Mothocya parvostis Bruce, 1986 by molecular sequence analyses along with other parasites obtained from H. sajori, the latter being morphologically confirmed by comparison with paratype materials of M. parvostis as well as the similar congener Mothocya sajori Bruce, 1986. The growth rates of the infected A. schlegelii juveniles from June to September in the years 2013-2015 and 2018 were significantly lower than those of the uninfected ones, suggesting a negative effect of the infection on the hosts. Our data on the prevalence and duration of the infection, as well as the body size gain of the hosts and parasites, corroborate a hypothesis that M. parvostis would utilize A. schlegelii as an optional intermediate host before it reaches the final host, H. sajori.

A new species of Crinoniscus Pérez, 1900 (Crustacea: Isopoda: Crinoniscidae) parasitising the pedunculate barnacle Heteralepas newmani Buhl-Mortensen & Mifsud (Cirripedia: Heteralepadidae) with notes on its ecology and a review of the genus.

The new species Crinoniscus stroembergi n. sp. belonging to the parasitic isopod family Crinoniscidae Bonnier, 1900, is described from a pedunculate barnacle host collected in the Mediterranean Sea. This is the first species of Crinoniscus Pérez, 1900 described from a host in the genus Heteralepas Pilsbry. The male cryptoniscus larva is distinguished from congeneric species by having a dorsoventrally flattened body with the posterolateral margins of the cephalon scarcely extending beyond the anterior margin of pereomere 1 (in contrast to C. cephalatus Hosie, 2008 with extended margins); articles 1 and 2 of the antennule being subequal in width and the anterodistal angle of antennule article being low and rounded; propodus of pereopods 6 and 7 having a sinuous dorsal margin that is distally narrowing and the posterior margin of the pleotelson being short and rounded. The mature females of species in Crinoniscus are of two basic forms: those with ventrolateral lobes on the pereomeres (C. alepadis (Gruvel, 1901) n. comb., and C. politosummus Hosie, 2008) and those lacking such lobes (C. cephalatus and C. stroembergi n. sp.). The mature females of C. stroembergi n. sp. can be distinguished from other species of Crininiscus based on their lacking lobes on the anterior end. The material examined includes the male and three female developmental stages of the parasite in the host, Heteralepas newmani Buhl-Mortensen & Mifsud. The mouthparts of the immature female are described and the feeding biology and effects on the host are discussed along with a review of feeding modes in species of the Cryptoniscoidea. Leponiscus alepadis is transferred to Crinoniscus; the latter genus now contains five species. A modified diagnosis of Crinoniscus is provided.

Morphostructural and immunohistochemical study for evaluation of nano-TiO2 toxicity in Armadillo officinalis Duméril, 1816 (Crustacea, Isopoda, Oniscidea).

Invertebrates are precious organisms in order to study environmental pollution. In particular, they appear to be suitable as a bioindicator species for pioneer ecotoxicity studies on new xenobiotics such as nanoparticles. In fact, they are able to absorb nanomaterials scattered in the environment in different ways and it's known the compartmentalization of nano-sized contaminants in selected tissues and intracellular organelles. Titanium dioxide represents the most used nanoparticulate, destined to become probably ubiquitous in the environment. Recently, some research has been published on the toxic potential of nano-TiO2 in several animal species. Among all invertebrates, Oniscidean Isopods are the only taxon of Crustaceans that has become completely terrestrial, known as excellent bioindicators and bioaccumulators. They have a digestive gland, the hepatopancreas, which is the location of election for the accumulation of pollutants. For this reason, they are considered efficient animal models to ecological studies. For this study, we collected Armadillo officinalis from Natural Oriented Reserve of "Vendicari" (Sicily, Italy), to evaluate the toxicity of titanium dioxide (TiO2 ) on their hepatopancreas, after a short period of exposure. We conducted morphostructural and immunohistochemistry assays. The results suggested a great capacity of the species of bioaccumulation of nanoparticles in the hepatopancreas, where a strong positivity to the metallothioneins was highlighted. Our study confirms that Oniscidean Isopods, in particular Armadillo officinalis, proved to be an appropriate indicator of pollution in terrestrial ecosystems from nanoparticles.

First report of Livoneca guianensis (Isopoda: Cymothoidae) in Leporinus fasciatus (Pisces: Anostomidae) in Brazil / Primeiro registro de Livoneca guianensis (Isopoda: Cymothoidae) em Leporinus fasciatus (Pisces: Anostomidae) no Brasil

Abstract This study aimed to report the first record of the occurrence of the parasite Livoneca guianensis in Leporinus fasciatus in Brazil. In September 2019, during surveys of the parasites in 12 specimens of L. fasciatus from the lower Jari River basin, in Amapá State, Brazil, one specimen of L. guianensis was found in the tongue of one host. The prevalence of L. guianensis was 8.3%, and it was observed that this parasite has a geographic distribution restricted to South America, parasitizing fish species from Guyana and Brazil. This study contributed to the first report on the occurrence of L. guianensis for L. fasciatus in Brazil. In addition, increased the knowledge on the geographic distribution of this parasite with morphometric data, which are scarce for this Cymothoidae.

Resumo Este estudo objetivou registrar a primeira ocorrência de Livoneca guianensis em Leporinus fasciatus no Brasil. Em setembro de 2019, durante uma pesquisa sobre parasitos em 12 espécimes de L. fasciatus do baixo Rio Jari, Estado do Amapá, norte do Brasil, um espécime de L. guianensis foi encontrado na língua de um hospedeiro. A prevalência de L. guianensis foi de 8,3% e foi observado que esse parasito tem distribuição geográfica restrita à América do Sul, parasitando espécies de peixes da Guiana e do Brasil. Este estudo contribuiu com o primeiro relato da ocorrência de L. guianensis para L. fasciatus no Brasil. Além disso, aumentou o conhecimento sobre a distribuição geográfica desse parasito, contribuindo com dados morfométricos que são escassos para esse Cymothoidae.

CO2 leakage simulation: Effects of the decreasing pH to the survival and reproduction of two crustacean species.

The effects of CO2-related acidification on two crustacean populations, the isopod Cyathura carinata and the amphipod Elasmopus rapax, were studied. Three pH levels were tested: artificial seawater without CO2 injection and two levels of reduced pH. Even though RNA:DNA ratio was reduced for both species, no statistical significant differences were found between the control and the treatments. Both species experienced a reduction in survivorship, longevity and the body length of surviving animals; although the impairment observed in E. rapax was more severe than in C. carinata. The long life span isopod and the short life span amphipod experienced a high degree of impairment in the reproduction, likely due to the reallocation of resources from reproduction to body maintenance and increasing survival by postponing the brood production. Regardless of the underlying processes and the energetic pathways, both experienced failure to reproduce, which could lead to the local extinction of these species.

Hooked on you: shape of attachment structures in cymothoid isopods reflects parasitic strategy.

BACKGROUND: Parasite attachment structures are critical traits that influence effective host exploitation and survival. Morphology of attachment structures can reinforce host specificity and niche specialisation, or even enable host switching. Therefore, it is important to understand the determinants of variation in attachment structures. Cymothoid isopods are striking ectoparasites of fishes that include the infamous 'tongue-biters.' They are known to parasitise hosts in one of four qualitatively distinct anatomical regions. Here, we quantify variation in cymothoid attachment structures - hook-like appendages called dactyli - and test whether differences in dactylus shape are correlated with parasite mode (where they attach), allometry, or both, using multivariate ordinary least squares regression. We also assess the influence of shared ancestry on shape using a molecular phylogeny to weight our models using phylogenetic generalised least squares regression. RESULTS: We find clear differences in shape between externally-attaching and internally-attaching cymothoids but also between anterior and posterior dactyli across various species with the same attachment mode. Allometric effects are significant for anterior but not posterior dactyli. Mouth-attaching species show greater shape variability than gill- and mouth-attaching species. We find no evidence that there are clade-specific patterns of association between parasite mode and dactylus shape. CONCLUSIONS: Parasite mode appears to be the main driver of attachment morphology. This likely reflects several components of parasite ecology including feeding and functional demands of attachment in different microhabitats. Geometric morphometric approaches to the quantification of shape variation of simple structures is an effective tool that provides new insights into the evolvability of parasite attachment.

Redescription and neotype designation for the poorly known fish parasitic cymothoid Joryma brachysoma (Pillai, 1964) (Crustacea: Isopoda) from India.

A neotype is designated for Joryma brachysoma (Pillai, 1964). The general morphology and appendages of the female and male stages are illustrated and re-described based on the neotype and several additional fresh specimens collected from the type host Pellona brachysoma Bleeker (=Sardinella brachysoma Bleeker) from nearby the type locality. The present redescription and neotype designation further resolves the taxonomic ambiguity regarding the species identification and conserves the name of J. brachysoma.

Genetic and morphological divergence at a biogeographic break in the beach-dwelling brooder Excirolana hirsuticauda Menzies (Crustacea, Peracarida).

BACKGROUND: There is a biogeographic break located at 30°S in the southeast Pacific, in a coastal area of strong environmental discontinuities. Several marine benthic taxa with restricted dispersal have a coincident phylogeographic break at 30°S, indicating that genetic structure is moulded by life history traits that limit gene flow and thereby promote divergence and speciation. In order to evaluate intraspecific divergence at this biogeographic break, we investigated the genetic and morphological variation of the directly developing beach isopod Excirolana hirsuticauda along 1900 km of the southeast Pacific coast, across 30°S. RESULTS: The COI sequences and microsatellite data both identified a strong discontinuity between populations of E. hirsuticauda to the north and south of 30°S, and a second weaker phylogeographic break at approximately 35°S. The three genetic groups were evidenced by different past demographic and genetic diversity signatures, and were also clearly distinguished with microsatellite data clustering. The COI sequences established that the genetic divergence of E. hirsuticauda at 30°S started earlier than divergence at 35°. Additionally, the three groups have different past demographic signatures, with probable demographic expansion occurring earlier in the southern group (south of 35°S), associated with Pleistocene interglacial periods. Interestingly, body length, multivariate morphometric analyses, and the morphology of a fertilization-related morphological character in males, the appendix masculina, reinforced the three genetic groups detected with genetic data. CONCLUSIONS: The degree of divergence of COI sequences, microsatellite data, and morphology was concordant and showed two geographic areas in which divergence was promoted at differing historical periods. Variation in the appendix masculina of males has probably promoted reproductive isolation. This variation together with gene flow restrictions promoted by life history traits, small body size, oceanographic discontinuities and sandy-beach habitat continuity, likely influenced species divergence at 30°S in the southeast Pacific coast. The degree of genetic and morphological differentiation of populations to the north and south of 30°S suggests that E. hirsuticauda harbours intraspecific divergence consistent with reproductive isolation and an advanced stage of speciation. The speciation process within E. hirsuticauda has been shaped by both restrictions to gene flow and a prezygotic reproductive barrier.

Tailored disorder in calcite organization in tergite cuticle of the supralittoral isopod Tylos europaeus Arcangeli, 1938.

The crustacean cuticle forms skeletal elements consisting of chitin-protein fibrils reinforced by amorphous and crystalline calcium carbonate and phosphate minerals. The edges of skeletal elements are of particular interest. They are subject to repeated strain and stress because they form transitions to the arthrodial membranes connecting them. These allow for relative movements of skeletal elements. In this study, we investigate structure, chemical composition, mineral organization and local mechanical properties of the anterior and posterior edges of the tergite cuticle in the conglobating beach isopod Tylos europaeus and compare these with the protective dorsal region of the tergites. The distribution of mineral phases at the edges resembles that of dorsal regions of the tergites. At the transition with the unmineralized arthrodial membrane the calcite containing distal exocuticle is replaced by epicuticular material and the subjacent cuticular layers containing amorphous calcium carbonate become enriched with amorphous calcium phosphate. At the edges, the local elastic modulus and hardness values are significantly lower compared to dorsal regions of the tergite cuticle, for both, the calcite and the amorphous mineral containing layers. The calcite within the tergite cuticle is assembled in different texture patterns: (i) almost random co-orientation, (ii) almost single crystalline calcite, and (iii) a graded organization. Calcite organization and co-orientation strength is highly variable, not only on very few tens of micrometres, but also between regions with different skeletal functionality. Our results show that besides structure and composition, patterns of calcite organization contribute to the hierarchical architecture and functionality of biological composites.