Embryonic development of verongid demosponges supports the independent acquisition of spongin skeletons as an alternative to the siliceous skeleton of sponges

Approximately 85% of extant sponges (phylum Porifera) belong to the class Demospongiae, which contains 14 taxonomic orders. In the orders Verongida, Dictyoceratida, and Dendroceratida, jointly referred to as 'keratose demosponges', the skeleton does not contain siliceous spicules but only spongin fibres. This shared trait has encouraged placement of these orders together within Demospongiae, although their relationships remain uncertain. The present study documents for the first time embryo development in the order Verongida ( Aplysina aerophoba ), providing some clues for phylogenetic inference. Spawned eggs were enveloped by a follicle of maternal cells. Embryos and larvae were chimeric organisms, the blastocoel of which was filled with symbionts and maternal cells migrated from the follicle. The ultrastructure of epithelial larval cells revealed: (1) a basal apparatus characterized by a peculiar, angling accessory centriole; (2) a pear-shaped nucleus with a protruding beak connected to the rootlets of the basal body; and (3) a distinctive Golgi apparatus encircling the nuclear apex. Developmental and ultrastructural findings support the concept, in congruence with recent molecular studies, that Verongida are more closely related to Halisarcida (askeletal sponges) and Chondrosida (askeletal sponges + sponges with spongin + spiculate sponges) than to the remaining 'keratose' orders, making a monophyletic 'supra-ordinal unit' equivalent to a subclass (Myxospongia, new subclass). Hence, spongin skeletons have evolved at least twice in Demospongiae. Independent acquisition of 'corneous' materials as an alternative to silica could have been stimulated by the radiation of diatoms at the Cretaceous–Tertiary boundary (approximately 65 Mya), which depleted silicon in the photic zone of the world's ocean.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 427–447.     

Species: towards new, well-grounded practices

Velasco recently criticized our formal definition of the species concept, arguing for its inappropriateness both in fundamental and practical aspects [Velasco JD (2008) Biological Journal of the Linnean Society , 93 , 865–869]. Here, we clarify some misunderstandings that are at the basis of Velasco's paper. First, we show why and how the definition of the species concept can be grounded in the theory of evolution and what that implies. Then, we explain why Velasco's formal criticisms are unjustified. Finally, we point out the practical and methodological consequences of a rigorous conceptual framework for species study, and we show that today's development of species delimitation methods fully agrees with our proposal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 217–222.     

DNA C-values in seven families fill phylogenetic gaps in the basal angiosperms

The evolutionary significance of the c . 1000-fold range of DNA C-values in angiosperms (1C =  c . 0.1–127.4 pg) has often attracted interest. A recent analysis, which superimposed available C-value data onto the angiosperm phylogeny, that placed Ceratophyllaceae as the most basal angiosperm family led to the conclusion that ancestral angiosperms were characterized by small genomes (defined as 1C £ 3.5 pg). However, with the recent increase in DNA sequence data and large-scale phylogenetic analyses, strong support is now provided for Amborellaceae and/or Nymphaeaceae as the most basal angiosperm families, followed by Austrobaileyales (comprising Schisandraceae, Trimeniaceae and Austrobaileyaceae). Together these five families comprise the ANITA grade. The remaining basal angiosperm families (Ceratophyllaceae, Chloranthaceae and magnoliids), together with monocotyledons and eudicotyledons, form a strongly supported clade. A survey showed that C-value data were scarce in the basal angiosperm families, especially the ANITA grade. The present paper addresses these phylogenetic gaps by providing C-value estimates for each family in ANITA, together with C-values for species in Chloranthaceae, Ceratophyllaceae and a previously unrepresented family in the magnoliids, the Winteraceae.  © The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 175–179.     

New carnivorous sponges (Porifera, Poecilosclerida) collected from manned submersibles in the deep Pacific

Nine new species of carnivorous sponges, namely Cladorhiza segonzaci , Chondrocladia koltuni , Chondrocladia lampadiglobus , Asbestopluma agglutinans , Asbestopluma ( Helophloeina ) formosa , Abyssocladia huitzilopochtli , Abyssocladia inflata , Abyssocladia dominalba and Abyssocladia naudur , are described. The subgenus Helophloeina Topsent, 1929 and the genus Abyssocladia Lévi, 1964 are revived and redefined, with an identification key for Abyssocladia which at present includes seven species. Eight of the sponges were collected from the French IFREMER manned submersible Nautile near active hydrothermal sites of the East Pacific Rise and of the North Fiji and Lau Basins, one from the Russian submersible Mir 2 in the Northwest Pacific near Bering Island, and one from the US submersible Alvin south of Easter Island. Their life conditions are described from direct observations from the submersibles. Some remarks are presented on the taxonomy of Cladorhizidae and more generally of carnivorous Poecilosclerida. The study suggests a very high degree of diversity in the deep Pacific carnivorous sponges.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 553–584.     

Chromosomal evolution of South American frugivorous butterflies in the Satyroid clade (Nymphalidae: Charaxinae, Morphinae and Satyrinae)

We describe the chromosome numbers of a monophyletic group of Satyroid subfamilies of primary fruit-attracted butterflies from South America: Charaxinae, Morphinae (including Brassolini) and Satyrinae. The charaxines do not have a distinct modal number. Their chromosome numbers are in the range n  = 6–50, with n  = 7–9, n  = 12, n  = 16, n  = 19–21, n  = 26, and n  = 28–31 being the most common numbers. Within the Morphinae, the Morphini have a modal n  = 28 and the Brassolini a modal n  = 29, with few exceptions. The Neotropical satyrines, in particular the basal species, have a weak modal n  = 29, which is a strong modal number in Palearctic satyrines. The African satyrines have an equally strong modal n  = 28. Most Neotropical satyrines have, like charaxines, chromosome numbers lower than the weak modal n  = 29, and often half this modal, but there are genera with stable numbers among the satyrines and charaxines. Evidently, the Neotropical satyroids descend from basal Nymphalidae with the typical lepidopteran modal number of n  = 31, which have also given rise to the Heliconiini with modal n  = 31 and 21 and Ithomiinae with modal numbers of n  = 14–15.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 467–481.     

A new species of Ferula (Apiaceae) from South Anatolia, Turkey

Ferula coskunii H. Duman & M. Sağıroğlu sp. nov. from the Amanos Mountains (C6: Hatay-Turkey) is described and illustrated. It is closely related to F. drudeana Korovin, from which it differs mainly in habit, basal leaves, stem and mericarp features.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 357–361.     

Diversity and specificity of Caribbean sponge–zoanthid symbioses: a foundation for understanding the adaptive significance of symbioses and generating hypotheses about higher-order systematics

The diversity and specificity of symbiotic associations may be useful in revealing the underlying ecology of symbioses and evolutionary relationships of symbiotic species. Symbioses between coral reef sponges and zoanthids are widespread and common in the greater Caribbean region, although the diversity and specificity of the species involved have only been explored at a few sites and the adaptive significance has only been examined for three combinations. We identified extensive diversity among sponges that associate with zoanthids by compiling sponge–zoanthid species associations from field surveys, the literature, and museum collections, and examined the patterns of specificity at multiple levels of sponge and zoanthid taxonomy. The results obtained indicate that facultative sponges are highly specific to the species of their partners whereas obligate zoanthids are not. The patterns of specificity among sponges and zoanthids suggest that many of these associations are not likely to be parasitic. Sponges harbouring photosynthetic endosymbionts associate at a disproportionately high frequency with zoanthids that harbour photosynthetic endosymbionts. Zoanthids embed in the surfaces of sponges to various degrees, resulting in a range of intimacy that negatively correlates with the number of hosts and polyp volume of zoanthids. Dendrograms based on the similarity among associations are largely consistent with current hypotheses of sponge higher-order systematics, but inconsistent with the current hypotheses of zoanthid systematics, and they highlight the potential utility of ecological characters in systematic analyses.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 695–711.     

Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia

We report on and name two new taxa of basal crocodylomorph archosaurs from the Lower Jurassic, Litargosuchus leptorhynchus gen. et sp. nov. , from the upper Elliot Formation (Stormberg Group) of South Africa, and Kayentasuchus walkeri gen. et sp. nov. , from the Kayenta Formation (Glen Canyon Group) of Arizona, USA. Examination of this material led to a reconsideration of basal crocodylomorph interrelationships. A phylogenetic analysis found no support for the monophyly of Sphenosuchia.  © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 136 , 77–95.     

Can a sponge feeder be a herbivore? Tylodina perversa (Gastropoda) feeding on Aplysina aerophoba (Demospongiae)

Feeding biology in mollusks has important biological, ecological and evolutionary implications because many of the characteristics we observe in mollusks arise from their co-evolution with diet organisms. We investigated the relationship between the opisthobranch Tylodina perversa and the sponges Aplysina aerophoba and Aplysina cavernicola in order to ascertain the trophic interactions between them. The opisthobranch preferred specimens of Aplysina aerophoba inhabiting shallow overdeep waters, ectosome of Aplysina aerophoba over choanosome, and showed no preference for Aplysina cavernicola . The sponge Aplysina cavernicola lacks the cyanobacteria abundant in the ectosome of Aplysina aerophoba . Our study shows that the opisthobranch Tylodina perversa actively selects for sponges or sponge zones with high concentration of cyanobacteria, i.e. only a fraction of the ingested material is of animal origin. Addition of cyanobacteria to symbiont-free sponge material induced a shift in mollusk preference. Our results cast doubt over the widely recognized qualification of Tylodina perversa as a carnivorous sponge feeder and show evidence that cyanobacteria determine the opisthobranch food selection. Whether this is an isolated example of how symbionts may determine trophic interactions between hosts and predators or it is widespread in benthic organisms remains an open question that requires further investigation.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 429–438.     

Species' diversity in the New Caledonian endemic genera Cephalidiosus and Nobarnus (Insecta: Heteroptera: Tingidae), an approach using phylogeny and species' distribution modelling

The patterns of local endemism in New Caledonia were analysed in two endemic genera of Tingidae (Insecta, Heteroptera), Cephalidiosus and Nobarnus , through a phylogenetic analysis and species' distribution modelling. The aim was to determine the possible causes of diversification and endemism in New Caledonia. Our results show that environmental conditions are probably important for the distribution of the genus Cephalidiosus , in conjunction with other factors such as resource (host plant) distribution, but suggest that the same environmental conditions have not influenced the speciation processes and diversification in the genus.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 177–184.     

Molecular characterization of an ancient Hepatozoon species parasitizing the 'living fossil' marsupial 'Monito del Monte' Dromiciops gliroides from Chile

The Microbiotheriid Dromiciops gliroides , also known as 'Monito del Monte', is considered to be a threatened species and the only living representative of this group of South American marsupials. During the last few years, several blood samples from specimens of 'Monito del Monte' captured at Chiloé island in Chile have been investigated for blood parasites. Inspection of blood smears detected a Hepatozoon species infecting red blood cells. The sequences of DNA fragments corresponding to small subunit ribosomal RNA gene revealed two parasitic lineages belonging to Hepatozoon genus. These parasite lineages showed a basal position with respect to Hepatozoon species infecting rodents, reptiles, and amphibians but are phylogenetically distinct from Hepatozoon species infecting the order Carnivora. In addition, the Hepatozoon lineages infecting D. gliroides are also different from those infecting other micro-mammals living in sympatry, as well as from some that have been described to infect an Australian species of bandicoot. The potential vector of this parasite appears to be the host-specific tick Ixodes neuquenensis because the sequencing of a long amplicon determined the presence of one of the two lineages found in the marsupial.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 568–576.     

Polymorphism hides cryptic species in Oerstedia dorsalis (Nemertea, Hoplonemertea)

The marine hoplonemertean Oerstedia dorsalis is considered to be a highly polymorphic species with extensive geographic distribution. We show, based on mitochondrial cytochrome c oxidase subunit I, 16S and nuclear internal transcribed spacer sequences, that there are genetic subsets withing this species. Seventy-one specimens of various colours from different geographic localities (in Europe) were sequenced and analysed using statistical parsimony and Bayesian analysis. Both analyses supported nine major clades. We conclude that O. dorsalis hides different species with geographic resolution. These species, however, appear to be polymorphic as well, and we find no diagnostic features in pigmentation or external characters to separate species within this complex.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 556–567.     

Changes in ventral head width, a discriminating shape factor among African cichlids, can be induced by chronic hypoxia

A massive enlargement of the gill surface proved to be an important factor in the hypoxia survival of young cichlids. Because the heads of cichlids are densely packed with structures related to both feeding and breathing, we hypothesized that the extra space needed for gill enlargement requires such large structural reorganizations that outer head shape is affected. We used a three-dimensional model to describe changes in the outer head shape of cichlids. Broods of cichlids of different phylogenetic lineages, habitats, and trophic specialization were split and raised at either 10% or 80–90% air saturation. Despite the above-mentioned differences between the species that were used, all hypoxia raised groups showed similar volume enlargements. Volume increases were most prominent in the ventral suspensorial and ventral opercular subcompartments. A relation with the enlarged gills of hypoxia raised fish is likely because the gills are mainly located in these compartments. The differences in ventral width correspond to those found in other studies comprising a wide variety of genotypic and phenotypic variations. The present study shows that such variation in the ventral width is conceivable by phenotypic plasticity alone.   © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 608–619.     

Phylogenetic patterns of diversification in a clade of Neotropical frogs (Anura: Aromobatidae: Mannophryne)

We used partial sequences of mitochondrial 16S and cytochrome oxidase I genes to perform a phylogenetic study of collared frogs (Anura: Aromobatidae: Mannophryne ), a genus endemic to Venezuela and the islands of Trinidad and Tobago. We analysed 1.2 kb from 13 of the 15 described species of Mannophryne . Maximum parsimony, maximum likelihood and Bayesian analyses support the monophyly of Mannophryne . Mannophryne consists of three deeply differentiated clades that split from each other in a relatively short period of time. The diversification of Mannophryne occurred well before the glacial-interglacial periods of the Quaternary. Our data support the taxonomic validity of M. olmonae , a species endemic to Tobago Island. Mannophryne olmonae is more closely related to the continental species Mannophryne riveroi than to the Trinidad island endemic Mannophryne trinitatis . As in most tropical clades of frogs, molecular evidence indicates that species richness in Mannophryne is largely underestimated and, consequently, current priorities for conservation are inadequate.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 185–199.     

Amana kuocangshanica (Liliaceae), a new species from south-east China

Amana kuocangshanica D. Y. Tan & D. Y. Hong is described as a new species from Mt. Kuocang, Zhejiang Province, south-east China. Morphological observations and a statistical analysis of the new species and its relatives show that it is most closely related to A. anhuiensis and A. erythronioides , but differs in having the lower leaf oblanceolate, the widest at two-thirds length from the base, tunics glabrous inside, and fruit beaks 5–7.5 mm long. The morphologies of the pollen, seeds, and epidermal cells of the leaves of this new species are described.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 435–442.     

A new species of Arabis (Brassicaceae) from inner Anatolia

A new species of Arabis L., A. erikii Mutlu sp. nov. , is described and illustrated. The species is restricted to inner Anatolia, south-west of Sivas. It grows on steppes. Diagnostic morphological characters are given for discrimination from the most similar taxa.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 251–256.     

Sex ratio, mating behaviour and Wolbachia infections in a sheetweb spider

The solitary sheetweb spider Pityohyphantes phrygianus has a female-biased primary sex ratio. Earlier studies have indicated that female post-copulatory position may influence the degree of bias, but the adaptive significance of the departure from a Fisherian 1 : 1 sex ratio is not known. In this study we show that there is a significant relationship between abdominal position and offspring sex ratio in undisturbed females. Moreover, female mating behaviour (and thus sex ratio) is correlated with female size, but this relationship may shift from negative to positive depending on the presence of the maternally acquired endosymbiotic bacterium Wolbachia in both sexes. A survey of wild-caught individuals shows that the incidence of Wolbachia infection in natural populations is high. This indicates that Wolbachia may be a significant factor influencing female post-copulatory behaviour and sex ratio in natural populations. We discuss various adaptive reasons for sex-ratio bias and behavioural plasticity in females.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 181–186.