The effects of the excretory-secretory products of fouling organisms on settlement of larvae of the sponge Halichondria panicea (Pallas, 1766) (Porifera: Demospongiae)

The effects of the excretory-secretory products (ESPs) of several fouling organisms on the larvae of the sponge Halichondria panicea were assessed in laboratory experiments. The ESPs of the brown alga Laminaria saccharina significantly stimulated larval settlement and metamorphosis, while the metabolites excreted by conspecific adult colonies were harmful to H. panicea larvae. The ESPs of the ascidians Styela rustica and Molgula citrina and the blue mussel Mytilus edulis impeded both the settlement and metamorphosis of the sponge larvae to varying degrees. The chemical cues of the bivalve Hiatella arctica had no significant effect on the number of settled larvae of H. panicea but retarded their metamorphosis.     

Structure and swimming behavior of the larva of Halichondria melanadocia (Porifera: Demospongiae)

Larvae of the sponge Halichondria melanadocia are of the parenchymella type and, during swimming, can change shape rapidly from cigar-like to ovoid. Larvae collected in Hawaii displayed neither qualitative nor quantitative differences in behavior or structure from those collected in Florida. Floridian larvae were examined at 2, 28, 48, and 72 hr after release to assess anatomical changes correlated with duration of the free-swimming period. Although 2 hr larvae were significantly longer than 48 or 72 hr larvae, other differences were not observed. Positively phototactic throughout the free-swimming period, the larvae eventually begin to swim on or near the bottom of dishes, settle temporarily, but can resume swimming before permanent settlement is achieved. The larva is extensively flagellated and a band of long flagella separates the lateral and posterior regions. The epidermis is a tall, simple columnar epithelium composed of highly polarized, monoflagellated cells. The interior contains at least two distinct amoeboid cell types that intermesh with the basal ends of epidermal cells within a loosely defined cavity. No spicules are present. Choanocyte chambers, found within 3 of the 50 larvae that were serially sectioned, varied in size and complexity, but were not associated with canals. This report is the-first of such chambers in Halichondria larvae. Spicules and choanocyte chambers are somatic structures associated with adults, and their appearance in larvae is presumably a consequence of a heterochronic event, most likely acceleration. The evolutionary significance of the occurrence of these traits in Halichondria larvae awaits further developmental analysis and greater phylogenetic resolution.     

The spermatogenesis ofHalichondria panicea (Porifera,Demospongiae)

Summary Spermatogenesis of the marine spongeHalichondria panicea begins with the break up of choanocyte chambers, choanocytes constituting the origin of spermatogonia. The transition from choanocytes to spermatogonia is direct, without cell division. Already the spermatogonia are flagellated. The ensuing large aggregates of spermatogonia are enclosed by spermatocyst-building cells. Further development takes place within the spermatocysts, mostly arranged in fields which, however, lack any developmental gradient. Within a single spermatocyst development is mostly synchronous. Spermatogonia transform into first order spermatocytes directly. The transition from spermatid to spermatozoon is characterized by an unusual prolongation of the chromatin, often resulting in a helical form of the chromosome material and a strong enlargement of the mitochondria which align with the nucleus, leading to an irregular shape of the spermatozoon. Another exceptional feature is the virtual absence of a Golgi apparatus during all stages of spermatogenesis. TheH. panicea investigated here contained only male reproductive elements, thus appear to be gonochorists. Some features of the spermatogenesis ofH. panicea, such as dissolving choanocyte chambers, the enclosure of spermatogonia by spermatocyst-building cells and the formation of a synaptonemal complex in first order spermatocytes occur in other sponge species as well; however, the early presence of flagella in spermatogonia, the absence of the Golgi apparatus and the later irregular development of nuclei, mitochondria and the spermatozoa themselves represent features hitherto not observed in sponges.     

Zoobenthos-inhabiting Sarcotragus muscarum (Porifera: Demospongiae) from the Aegean Sea

The cavities and outer surfaces of 20 individuals of the sponge Sarcotragus muscarum collected from the Turkish Aegean coast during July–September 1993 contained a total of 148 zoobenthic species with 5299 individuals belonging to seven taxonomic groups. Among these, Polychaeta accounted for 60% of the total number of species, while Crustacea comprised 71% of the total number of individuals and 40% of the wet weight of biomass. The amphipod and decapod crustaceans Tritaeta gibbosa and Synalpheus gambarelloides, and the bivalve Hiatella arctica, were the most abundant species on the sponge samples both in terms of number of individuals and biomass values. The community structure inside the sponges was greatly influenced by the nature of the substratum around the sponge, and by the overwhelming population of the Lessepsian echinoderm Ophiactis savignyi. Number of species, biomass and diversity index values were found to be positively related to the sponge volume. Mean density and biomass of the associated taxa are 0.22 ind cm^–3 and 11.63 mg cm^–3, respectively.     

Two new species in the family Axinellidae (Porifera,Demospongiae) from British Columbia and adjacent waters

Two new species of Demospongiae are described for British Columbia and adjacent waters in the family Axinellidae, Auletta krautteri sp. n. and Dragmacidon kishinensis sp. n. They represent range extensions for both of these genera. Both are fairly commonly encountered, Auletta krautteri below diving depths (87 to at least 300 m) and Dragmacidon kishinensis in shallow water (intertidal to 30 m). We propose an amended genus diagnosis for Auletta to account for the variability among species in principal spicules that form the ascending tracts to be either oxeas, styles or strongyles rather than just oxeas.     

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.     

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.     

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.     

Reproductive biology of three sponge species of the genus Xestospongia (Porifera: Demospongiae: Petrosida) from the Great Barrier Reef

The reproductive development of three species of the Petrosida, Xestospongia bergquistia, X. exigua, and X. testudinaria, was monitored for four years on a fringing reef at Orpheus Island, Great Barrier Reef, Australia. All three species were oviparous and female reproductive activity began prior to males becoming active. X. bergquistia and X. testudinaria were gonochoric and broadcast eggs in spawning events that were synchronous within species. Egg development occurred over more than five months in X. bergquistia and X. testudinaria and two months in X. exigua. Spawning was during periods of warm temperature and occurred in October or November for X. bergquistia and X. testudinaria, and January or February for X. exigua. Lunar phase was implicated in timing of spawning of X. testudinaria. Diel timing of spawning in X. testudinaria and X. bergquistia was consistently a morning event.     

Biological and chemical characterizations of three new species of Dysidea (Porifera: Demospongiae) from the Pacific Mexican coast

The integration of biological and chemical data has provided patterns to identify three new Dysidea species from the Gulf of California (Pacific Ocean, Mexico). These three species are differentiated by distinctive morphological characters, such as the prominent subdermal fibers running along the surface in Dysidea reformensis n. sp., the granular appearance of the surface in conjunction with the consistent habit in Dysidea cachui n. sp., and the skeletal details such as tertiary network between the secondary fibers in Dysidea uriae n. sp. Differences are also observed in the natural products content of these species. Sesquiterpene-hydroquinone derivatives are the main secondary metabolites of D. reformensis and D. cachui, with each species displaying a different chemical profile, while the furanosesquiterpene dendrolasin has been isolated from D. uriae. The distinctive morphological characters and the natural products contained by each species are also compared with those of related Dysidea species.     

Molecular phylogenies challenge the classification of Polymastiidae (Porifera,Demospongiae) based on morphology

Polymastiidae Gray, 1867 is a worldwide distributed sponge family, which has a great significance for understanding of the demosponge deep phylogeny since the former order Hadromerida Topsent, 1894 has been recently split based on the molecular evidence and a new separate order has been established for the polymastiids. However, molecular data obtained from Polymastiidae so far are scarce, while the phylogenetic reconstruction based on morphology has faced a deficit of characters along with the vagueness of their states. The present study is a phylogenetic reconstruction of Polymastiidae based on novel data on two molecular markers, cytochrome oxidase subunit I and large subunit ribosomal DNA, obtained from a broad set of species. Monophyly of the family and nonmonophyly of four polymastiid genera are revealed, suggesting a high level of homoplasy of morphological characters, which are therefore not an appropriate base for the natural classification of Polymastiidae. Although the presented phylogenies cannot yet provide an alternative classification scheme, several strongly supported clades, which may be used as reference points in future classification, are recovered and three taxonomic actions are proposed: transfer of one species from Radiella to Polymastia Bowerbank, 1862; transfer of three species from Radiella Schmidt, 1870 to Spinularia Gray, 1867; and the consequent abandonment of Radiella.     

The vacuolar cells of Oscarella lobularis (Porifera,Demospongiae): Ulatrastructural organization,origin, and function

This work is concerned with the ultrastructural organization and some histochemical features of the vacuolar cells of the demosponge Oscarella lobularis. Vacuolar cells are characterized by large clear vacuoles containing an aqueous fluid. They are commonly found in the mesohyl of this sponge and tend to constitute a sort of parenchyma in the choanosome. Mobile cells of the mesohyl appear to differetiate into vacuolar cells through the progressive formation of wide cytoplasmic lacunae. We have identified four types of cells showing progressive transformation toward the vacuolar cell type. Precursors (types 1–4) of the vacuolar cells probably derive from endopinacocytes, since they share several histochemical and ultrastructural characteristics with them. Our data support the notion that vacuolar cells are involved in the synthesis of collagen, act as a mechanical support of the sponge body, and are eventually extruded from the sponge through the canals of the aquiferous system.     

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.     

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     

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.     

Silica spicules and axial filaments of the marine sponge Stelletta grubii (Porifera,Demospongiae)

Summary In all cases an organic axial filament within the silica spicules of Stelletta grubii forms the core of the major axes of the glass. In the small, star-shaped silica spicules (asters) the filament is shown for the first time to be radial with an enlarged center; in the large four-rayed spicules (triaenes) it is four-rayed; and in the large single-rayed spicules (oxeas) the filament is single-rayed. In situ, the filament is not dissolved by boiling nitric acid and thus is apparently protected by encasement within the glass which can also be stratified. The small silica asters are formed by single cells which resemble the so-called spherulous cells of other sponges. The very large size of triaenes and oxeas suggests that they may possibly be formed by more than one cell. The diameter of the filament in the much smaller asters is much narrower than the filament in the larger spicules, indicating a possible relationship between filament diameter and spicule diameter. While the axial filament in larger spicules frequently has a triangular cross-section it can also be hexaognal. Some aster filaments also retain a close to hexagonal cross-section. Filaments freed from large spicules by hydrofluoric acid display a complex morphology; possibly there is an internal silicified core. Some reported aspects of filament morphology are, however, probably artefacts of desilicification with hydrofluoric acid. Offprint requests to: T.L. Simpson, Department of Biology, University of Hartford, West Harford, Connecticut 06117, USA (Permanent affiliation)     

Post-larval development of the commercial sponge Spongia officinalis L. (Porifera, Demospongiae)

This study investigated the development of the larvae of Spongia officinalis in experimental conditions, after settlement on plastic substrates, using electron and light microscopy. The released larvae show a dark pigmented ring distinguishes the posterior larval pole. The youngest larvae, covered with a flagellate epithelium, move onwards by rotating on their longitudinal axis. Over time a creeping-like motion prevails, probably linked to the need for settlement. After a free-swimming period of 24-48 h, larvae settle on the artificial substrate by the anterior pole. At settlement, the flagellate epithelium is substituted by flattened cells, which delimit the outermost surface. Post-larvae were reared to about three months. The early phase of post-larval differentiation shows a solid interior mainly consisting of granular cells varying in shape and size. They are included in a dense collagen matrix that contains a conspicuous amount of bacteria. Lacunae are already evident in the initial phase of metamorphosis. In several of them, cell debris and nucleate cells are visible. This feature is consistent with a progressive reduction of the cell mass (autolysis). Neither choanocyte chambers nor canals differentiate. The morphogenetic process leads to a metamorph only consisting of vacuolated cells and collagen fibrils included in a thin fibrous coat.     

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.