Morphological and ecological differences in two electrophoretically detected species of Cliona (Porifera, Demospongiae)

The taxonomie status of different morphotypes generally ascribed to the species of boring sponge Cliona viridis is examined. Morphometric, ecological and genetic differences between two morphotypes living sympatrically on rocky shores of the Ligurian Sea are considered. Morphotype 1 is characterized by small, well separated papillae (α stage), exclusively boring into the crust of coralline algae on the exposed rocky bottoms of the upper sublittoral zone. Morphotype 2 shows bigger papillae, often connected by portions of encrusting tissue ( β stage), and is predominantly found boring into organogenic concretions in slightly lighted habitats. Measurements of tylostyles indicate that morphotype 1 has spicules significantly smaller than morphotype 2. Electrophoretic analysis shows that the two morphotypes are fixed for different alleles at all four loci scored, indicating absence of gene flow between the two populations, i.e. they appear to be distinct biological species.     

Larval development in the Homoscleromorpha (Porifera, Demospongiae)

Abstract. Embryonic development from coeloblastula to fully developed larva was investigated in 8 Mediterranean homoscleromorph species: Oscarella lobularis, O. tuberculata, O. microlobata, O. imperialis, Plakina trilopha, P. jani, Corticium candelabrum , and Pseudocorticium jarrei. Morphogenesis of the larva is similar in all these species; however, cell proliferation is more active in species of Oscarella than in Plakina and C. candelabrum. The result of cell division is a wrinkled, flagellated larva, called a cinctoblastula. It is composed of a columnar epithelium of polarized, monoflagellated cells among which are scattered a few non-flagellated ovoid cells. The central cavity always contains symbiotic bacteria. Maternal cells are also present in O. lobularis, O. imperialis , and P. jarrei. In the fully developed larva, cell shape and dimensions are constant for each species. The cells of the anterior pole have large vacuoles with heterogeneous material; those of the postero-lateral zone have an intranuclear paracrystalline inclusion; and the flagellated cells of the posterior pole have large osmiophilic inclusions. Intercellular junctions join the apical parts of the cells, beneath which are other specialized cell junctions. A basement membrane underlying the flagellated cells lines the larval cavity. This is the first observation of a basement membrane in a poriferan larva. The basal apparatus of flagellated cells is characterized by an accessory centriole located exactly beneath the basal body. The single basal rootlet is cross striated. The presence of a basement membrane and a true epithelium in the larva of Homoscleromorpha—unique among poriferan clades and shared with Eumetazoa—suggests that Demospongiae could be paraphyletic.     

Differences between two sympatric species of Tethya (Porifera, Demospongiae) concerning the growth and final form of their megasters

The megasters of T. aurantium and T. citrina consist, in their mature stage, of spherasters and spheroxyasters respectively. A study of the ratio between the ray length and the diameter of the centre (R/C), which determines the megaster shape, has been carried out on spicules of different size.
While the small megasters of these two species have a similar shape, successive stages show in T. aurantium an isometric growth and in T. citrina an allometric growth with rays growing faster than the centre. The two species differ also in the mean size, size frequency distribution and number of rays of the asters. Differences in the mean total diameter and shape were also detected among spicules coming from different regions of the sponge body. The taxonomic and evolutionary meaning of these phenomena are discussed.     

Association of the sponge Tethya orphei (Porifera, Demospongiae) with filamentous cyanobacteria

Abstract. Specimens of the sponge Tethya orphei , collected in February 2005 on the underside of coral stones on Arì Athol (Maldives), have been processed for histological and ultrastructural investigations. The cortical layer of the sponge was found to be permeated by filamentous cyanobacteria, the trichomes of which measured 45–63 μm on average and were composed of 10–14 cells. The fine organization of the filaments was consistent with their taxonomic identification as Oscillatoria spongeliae. These filaments filled the cortical region of the sponge and penetrated inward into the upper choanosomal region, where they sometimes overlapped the siliceous spicule bundles. A budding specimen of T. orphei showed that the filaments were also present in the single bud protruding from the sponge surface, demonstrating that asexual reproduction can vertically transmit these symbionts from sponge to sponge. The occurrence of filaments in all the specimens studied is consistent with the assumption that filamentous cyanobacteria are not mere intruders but mutualistic symbionts with members of T. orphei.     

Epibiontic and endobiontic polychaetes of Geodia cydonium (Porifera,Demospongiae) from the Mediterranean Sea

Polychaete assemblages associated to the sponge Geodia cydonium were investigated at two sampling sites in the Mediterranean Sea: Porto Cesareo Basin (Apulia) and Marsala Lagoon (Sicily), both characterized by sheltered hydrodynamic conditions. Samples were seasonally performed during 1997, in order to compare the assemblages coming from the two localities studied, considering separately the internal and external tissues of the sponge, and with the aim of evaluating the influence of sponge size on polychaete colonization. The examined sponge is characterized by a peculiar stratification of its tissues: an external thick and hard layer, the cortex, and an internal softer one, the choanosome. Statistical analysis showed that this was the main factor controlling polychaete assemblage, with the internal tissue, less rich and diversified, appearing impoverished with respect to the external layer. A similarity in species composition was observed between sites, even though some differences were evidenced in the abundance of some species, mainly reflecting differences in local environmental conditions. Species richness and density increased with the increasing sponge size. Such a situation is particularly evident at Porto Cesareo, where sponges are covered by an algal layer which is particularly rich on the largest specimens, thus suggesting that most of the species of polychaetes were linked more to the neighbouring environment than to the sponge itself.     

Taxonomic revision of the Mediterranean Plakina Schulze (Porifera, Demospongiae, Homoscleromorpha)

Only three species of the sponge genus Plakina Schulze have been described from the Mediterranean since 1880, in spite of a large amount of allegedly intraspecific variability in morphological characters. However, recent genetic studies based on electrophoretic techniques have revealed extensive cryptic speciation in north-western Mediterranean Plakina , demonstrating that most of this variation was interspecific rather than intraspecific. We describe in detail the morphology and anatomy of four new species of Plakina from the Mediterranean– P. crypta, P. weinbergi, P. endoumensis and P. jani –of which the latter two were discovered through allozyme electrophoresis. Plakina monolopha Schulze and P. trilopha Schulze are redescribed, and their morphological and geographical limits are discussed along with those of P. dilopha Schulze. Accurate analysis of the internal anatomy and of the shape and ramification pattern of lophose spicules in scanning electron microscopy provides new, powerful morphological criteria for species discrimination in Plakina. More widespread use of such new taxonomic characters should provide evidence against the alleged cosmopolitanism of some Plakina species, thus generating an increase in estimates of the biodiversity of plakinids.     

An unexpectedly sophisticated, V-shaped spermatozoon in Demospongiae (Porifera): reproductive and evolutionary implications

The demosponge Crambe crambe shows a peculiar spermatogenesis, hard to be reconciled with the basal position of sponges in the animal phylogeny. Early spermatogenesis stages showed most of the simple features expected in sponges. However, spermiogenesis departed from the anticipated process. Spermatids lengthened remarkably, forming a deep cytoplasmic pit around the cilium insertion, with the proximal axoneme bending to produce a V-shaped spermatozoon surprisingly similar to that known in the phylum Phoronida. The cytology was unexpectedly complex, with a needle-like nucleus of helically condensed chromatin, a conical acrosome with a subacrosomal rod, and a mitochondrion connected to the basal body by striated rootlets. These findings establish that the spermatozoon of broad-casting demosponges occurs in two structural categories ('primitive' and 'modified' type). This dualistic condition must necessarily have pre-dated the evolutionary apparition of higher metazoans, if we are to keep regarding sponges as the most primitive animals. We hypothesize that internal fertilization in C. crambe – and incidentally other demosponges – may depart from the general model assumed for spermcasting sponges. The V-shape of this spermatozoon suggests a design to favour autonomous penetration through the dense mesohyl to reach the oocytes, rather than engulfment and transportation by carrier cells towards the oocyte.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 413–426.     

Morphology and ultrastructure of the swimming larvae of Crambe crambe (Demospongiae, Poecilosclerida)

Abstract. We describe the morphology and ultrastructure of the free-swimming larvae of the sponge Crambe crambe , one of the most abundant encrusting sponges on shallow rocky bottoms of the western Mediterranean. Larvae of C. crambe are released in July and August. The larva is uniformly flagellated except at the posterior zone. Flagellated cells are extraordinarily slender, elongate, and sinuous and form a pseudo-stratified layer. Their distal zone contains abundant mitochondria, some small vesicles, a Golgi complex, and the basal apparatus of the flagellum. Abundant lipid droplets are present throughout the cell. The nucleus is most often in a basal position. The flagellum projects from the bottom of an asymmetrical socket formed by cytoplasmic expansions. The basal body extends in a conical tuft and a laminar rootlet in close association with the Golgi system. The cells at the posterior pole are flat and polygonal on the surface, with long overlapping pseudopodia in the typical shape of a pinacoderm. Sparse collagen is present throughout the whole larva including the flagellated layer. Archeocytes and sclerocytes are abundant in the posterior region. Typical collencytes and spherulous cells seem to be absent. Intracellular and extracellular rod-like bacteria with conspicuous fimbria occur exclusively in the posterior region of the larva. The asymmetrical cytoplasmic prolongations, which surround the flagellum, and the basal apparatus of the flagellum are suggested as the sites of stimulus reception and triggering of locomotor responses, respectively. This ultrastructural study of the larva of C. crambe has shown features directly linked to its behavior and ecology.     

Fauna of Sponges (Porifera) of Peter the Great Bay,Sea of Japan

A complete list of sponges of Peter the Great Bay (northwestern Sea of Japan) is presented comprising 40 species belonging to 27 genera, 19 families, 9 orders, and 2 classes. Information is presented about the biogeographical composition of sponge fauna of the bay, the depths of their habitats, and their substrates.Original Russian Text Copyright ¢ 2005 by Biologiya Morya, Khodakovskaya.     

Symbiosis of Mycale (Mycale) vansoesti sp. nov. (Porifera, Demospongiae) with a coralline alga from North Sulawesi (Indonesia)

Abstract. The symbiotic association between the new sponge species Mycale vansoesti and the coralline alga Amphiroa sp. from the Bunaken Marine Park (North Sulawesi, Indonesia) is described. The alga completely pervades the sponge. The color of the sponge ectosome is white, both on the external surface and on the atrial wall, but where the alga is present the sponge takes on the light pink color of the alga. The sponge spicular complement is characterized by mycalostyles, anisochelae of two types, sigmas (often "C" shaped), and extremely abundant toxas organized in bundles forming toxadragma. In the association, the sponge shows very low silicate value, and consequently the alga represents the main skeleton of the sponge. On the other hand, the sponge affects the morphology of the alga, leading to a cylindrical shape, with thalli running parallel to the sponge surface. This association seems to be obligate for the sponge, as we found no sponges of this species living in isolation.     

The architecture of the canal systems of Petrosia ficiformis and Chondrosia reniformis studied by corrosion casts (Porifera,Demospongiae)

Summary The three-dimensional organization of the canal system in two sponge species, Petrosia ficiformis and Chondrosia reniformis, was studied using corrosion casts. Casts were made of live animals, in situ, and canal replicas were analzyed by scanning electron microscopy (SEM). In P. ficiformis the incurrent system consists of a superficial canal network giving rise to large radial canals, which ramify and anastomosize forming an internal web. Excurrent canals are arranged into modular ramified systems radiating from atrial cavities opening to the exterior. Main excurrent canals run at various depths within the sponge, even through the superficial incurrent network. Both incurrent and excurrent canal replicas show smooth, blind-ending capillaries. Some large incurrent canals merge with excurrent ones, thus bypassing choanocyte chambers. In C. reniformis there is a cortical collagen layer crossed by three-like incurrent canals, the twigs of which communicate with groups of inhalant pores. The stems of tree-like canals penetrate into the sponge medulla where they ramify and anastomosize to form a web. Main excurrent canals arise from large cloacal ducts leading to the oscular openings. They give rise to a sequence of branches intersecting the incurrent web. Both incurrent and excurrent canals have sharp, blind-ending capillaries. Morphometric data functions show that diameter scaling in canal branches is exponential in Petrosia and linear in Chondrosia. Structural differences and homologies between the two species are discussed.     

Branching out: a remarkable new branching syllid (Annelida) living in a Petrosia sponge (Porifera: Demospongiae)

We describe the morphology and biology of a previously unknown form of branching annelid, Ramisyllis multicaudata gen. et sp. nov. , an endosymbiont of shallow‐water marine sponges (Petrosia sp., Demospongiae) in northern Australia. It belongs to the polychaete family Syllidae, as does Syllis ramosa McIntosh, 1879, the only other named branching annelid, which was collected from deep‐water hexactinellid sponges during the 1875 Challenger expedition. It differs from S. ramosa in parapodial and chaetal morphology. Ramisyllis multicaudata gen. et sp. nov. has segments of several types, including specialized posterior segments on the emergent portions of the worm, and simplified elongate segments that bridge larger cavities in the sponge interior. Aside from the obvious branching form, the new annelid is similar to Parahaplosyllis, differing from it in lacking pharyngeal armature and in the details of the parapodial chaetae and dorsal cirri. Molecular evidence from 16S and 18S rDNA supports a sister‐group relationship with Parahaplosyllis, with both being sister to Trypanosyllis and Eurysyllis. The phylogenetic position of R. multicaudata gen. et sp. nov. indicates that branching has evolved independently in Ramisyllis gen. nov. and Syllis. This is supported by differences in the branching process between the two taxa: in S. ramosa branching is initiated by segment addition at the parapodium, whereas in R. multicaudata gen. et sp. nov. segments are added from a region between parapodia. A model for branching in R. multicaudata gen. et sp. nov. is proposed and possible developmental processes underlying branching in Annelida, and body symmetry comparisons with other invertebrates, are also discussed. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 481–497.     

Choanocyte ultrastructure in Halisarca dujardini (Demospongiae,Halisarcida)

Understanding poriferan choanocyte ultrastructure is crucial if we are to unravel the steps of a putative evolutionary transition between choanoflagellate protists and early metazoans. Surprisingly, some aspects of choanocyte cytology still remain little investigated. This study of choanocyte ultrastructure in the halisarcid demosponge Halisarca dujardini revealed a combination of minor and major distinctive traits, some of them unknown in Porifera so far. Most significant features were 1) an asymmetrical periflagellar sleeve, 2) a battery of specialized intercellular junctions at the lateral cell surface complemented with an array of lateral interdigitations between adjacent choanocytes that provides a particular sealing system of the choanoderm, and 3) a unique, unexpectedly complex, basal apparatus. The basal apparatus consists of a basal body provided with a small basal foot and an intricate transverse skeleton of microtubules. An accessory centriole, which is not perpendicular to the basal body, is about 45°. In addition, a system of short striated rootlets (periodicity = 50–60 nm) arises from the proximal edge of the basal body and runs longitudinally to contact the nuclear apex. This is the first flagellar rootlet system ever found in a choanocyte. The accessory centriole, the rootlet system, and the nuclear apex are all encircled by a large Golgi apparatus, adding another distinctive feature to the choanocyte cytology. The set of distinct features discovered in the choanocyte of H. dujardini indicates that the ultrastructure of the poriferan choanocyte may vary substantially between sponge groups. It is necessary to improve understanding of such variation, as the cytological features of choanocytes are often coded as characters both for formulation of hypotheses on the origin of animals and inference of phylogenetic relationships at the base of the metazoan tree. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Ultrastructural study of spermatogenesis in Oscarella lobularis (Porifera,Demospongiae)

Summary

The various phases of spermatogenesis in the demosponge Oscarella lobularis were studied by electron microscopy. Spermatogenesis occurs within spermatic cysts, which are presumed to derive from choanocyte chambers by transformation of choanocytes into spermatogonia. Germ cells develop asynchronously within spermatocysts, and cytoplasmic bridges, indicating incomplete cells division, connect several germ cells. Attached spermatogonia suggest gonial generations. Spermatocytes I typically show the presence of synaptonemal complexes indicating meiotic divisions. Spermatocytes II have a small size probably because of the meiotic divisions of spermatocytes I. Spermatids are characterized by an acrosome, a big mitochondrion and a peripheral sheath of condensed chromatin surrounding a clearer central area in the nucleus. The mature spermatozoon shows a lateral flagellum and a flattened acrosome capping the nucleus. The phylogenetic implications of some features of the spermatozoon are suggested.     

Interspecific variation in arrangement and morphology of micrasters of Tethya species (Porifera,Demospongiae)

Summary A new distinctive feature between the two Mediterranean species of Tethya, T. aurantium and T. citrina has been found in the body arrangement of different types of micrasters. Contrary to the previous assumptions, T. aurantium has two clearly distinct categories of micrasters: the chiaster-tylaster in the cortex and the larger, slender oxyaster in the choanosome. T. citrina has only slightly differentiated micraster sets in the cortex and choanosome; in the latter the shape of micrasters is close to that of oxyasters. SEM analysis shows that differences in micraster shape depend on the cylindrical or conical form of rays and on the distribution, density and strength of the microspines along their axis. The relationship between the degree of micraster differentiation and the development of the cortex in the two species is discussed.     

Dynamic structure of the mesohyl in the sponge Chondrosia reniformis (Porifera, Demospongiae)

The common demosponge Chondrosia reniformis possesses the capacity to undergo an unusual creep process which results in the formation of long outgrowths from the parent body. These shape changes, which have been interpreted as adaptive strategies related to environmental factors, asexual reproduction or localised locomotor phenomena, are due mainly to the structural and mechanical adaptability of the collagenous mesohyl. This contribution describes the morphological correlates of mesohyl plasticisation in C. reniformis. The microscopic anatomy of the mesohyl was examined when it was in different physiological conditions: (1) standard ”resting” condition, (2) ”stiffened” condition and (3) dynamic ”creep” condition. In this last case four representative regions of the sponge body were analysed: the parent region, the elongation region, the transition region and the propagule region. The results show that the histological modification of the sponge mesohyl during plasticisation is limited and localised. The most significant structural changes involve mainly cytological features of specific cellular components characterised by granule inclusions (i.e. the spherulous cells) and the arrangement and density of the collagenous extracellular framework, though the integrity of the collagen fibrils themselves is not affected. Morphological and functional aspects of mesohyl plasticisation invite comparison with the mutable collagenous tissue of echinoderms. Possible functional analogies between these two tissues are hypothesised. Accepted: 29 June 2001     

Investigation of the budding process in Tethya citrina and Tethya aurantium (Porifera, Demospongiae)

The budding process has been studied in two congeneric Mediterranean species belonging to Tethya from different sampling sites: Marsala and Venice Lagoons (Tethya citrina); Marsala Lagoon and Porto Cesareo Basin (Tethya aurantium). Buds, connected to the adult by a spiculated stalk, differ between the two species in morphology and size, since those of T. citrina are small with elongated bodies, showing only a few spicules protruding from the apical region, whereas those of T. aurantium are round, larger, and show spicules radiating from the peripheral border. In T. citrina, cells with inclusions, varying in electron density and size, represent the main cell types of the buds. In T. aurantium, the cell component shows a major diversification, resulting from spherulous cells, grey cells, vacuolar cells and peculiar micro-vesicle cells. Neither canals nor choanocyte chambers were observed in the buds of the two species. In T. citrina, bud production is similar in both sampling sites. In T. aurantium, budding occurs more rarely in Porto Cesareo Basin, probably in relation with environmental factors, such as the covering of the cortex by sediment and micro-algae. Finally, in the buds of both species, the spicule size does not differ from that of the cortex of the adult sponges, further supporting the main involvement of the cortex in organizing the skeletal architecture of the buds.