Overgrowth competition between clades: implications for interpretation of the fossil record and overgrowth indices

Overgrowth interactions (2693 in total) were observed among three major groups (arguably clades) of bryozoans--cheilostomatids (57 species), ctenostomatids (3 species), and cyclostomatids (14 species). The bryozoans studied here occur in shallow water at high-temperate polar latitudes where they encrust hard substrata such as rock piles. The main study site was the intertidal and infralittoral zones of Kodiak Island, Alaska, but observations were also made in similar zones of South Georgia Island and the Falkland Islands in the South Atlantic Ocean. Cheilostomatids dominated the number of species, individuals, and interactions at all depths. Intraclade interactions formed 73.7% of the encounters for cheilostomatids, 1.6% for ctenostomatids, and 5.7% for cyclostomatids. The competitive ranking of the three clades was broadly ctenostomatids > cyclostomatids > cheilostomatids. Significantly, these results contradict all previous quantitative studies of bryozoan overgrowth, in which cheilostomatids are reported to overgrow cyclostomatids at a higher rate. From these studies and the literature, we calculated win indices to vary from 0 to 0.42 for living cyclostomatids, from 0.08 to 0.9 for living cheilostomatids, and from 0.25 to 0.75 for living ctenostomatids. The win indices of cyclostomatid and cheilostomatid clades show significantly more variation in living assemblages than in fossil assemblages. This disparity may be due to differential preservation (polar and subpolar assemblages last less than 4 years). The diversity was very high in terms of both species richness and interaction types (outcomes between competitor pairs). Comparison with the literature suggests the possibility that nearshore diversity of bryozoans may be bimodal (have two peaks) between high arctic and antarctic latitudes. Indices of success in overgrowth competition have been constructed in various ways. For cheilostomatids, the method of calculation had little influence on the ranking of representatives. In contrast, the apparent success of ctenostomatids and cyclostomatids varied hugely with how the index was calculated. This inconsistency is due to the use of very different strategies in overgrowth competition; among the two latter groups, many interactions involve tied outcomes.     

Search mechanism of a stream grazer in patchy environments: the role of food abundance

Summary The search behavior of the grazing stream insect Baetis tricaudatus (Ephemeroptera: Baetidae) was examined in field and laboratory experiments. Regardless of food abundance in experimental habitats, nymphs spent significantly more time in food patches than predicted if they had moved randomly with respect to patches. A significant reduction in movement rate within patches relative to movement rate between patches largely accounted for these results. The movement pattern within patches was highly systematic and in agreement with predictions of optimal foraging theory since food was uniformly distributed within patches. Between-patch search movements were affected by food abundance in the most recently grazed patch. Search intensity after departure from a patch was positively related to food abundance in the patch while movement rate after patch departure was inversely related to patch food level. These effects produced between-patch movement patterns that were suboptimal in the experimental habitats because they resulted in revisitation of previously depleted patches. However, differences between experimental and natural habitats in the spatial occurrence of patch types suggest that Baetis between-patch search behavior may be adaptive in natural habitats.     

Structural and taphonomic analysis of a columnar coralline algal build-up from SE Sicily

Unusual “coralligenous” build-ups were found at the infra–circalittoral boundary from SE Sicily. They are columnar, rising up perpendicularly from the bottom, and consist of an inter-growth of encrusting algae and invertebrates. A selected build-up and its longitudinal section, together with some thin sections of selected parts, were studied to detail the growth structures, composition and fabric of the trapped sediments. The coralline algae show a succession of “concentric”, “fruticulose” and “foliaceous” structures. The framework delineates cavities bounded by algal thalli, sometimes filled by sediments. The organisms constituting the framework were grouped into functional guilds (Fagerström, 1991): “primary frame builders” (mainly coralline algae); “secondary frame builders” (vermetids, serpulids and some bryozoans); “bafflers” (erect, flexible bryozoans, some sponges and soft algae); “binders” (encrusting bryozoans, some crustose coralline algae and serpulids); “destroyers” (a few endolithic algae, rare sponges and bivalves); “dwellers” (brachiopods, foraminifers, bivalves, small bryozoans and some motile organisms). The distribution pattern of taphonomic features (composition, grain size and distribution of sediments filling cavities; presence of cement and its location; mineralogical composition and crystal size), together with interactions between growth structures and functional guilds, allowed one to distinguish three taphofacies, each recording different environmental conditions. They have been tentatively related with cyclic sequences testifying to local hydrodynamic regimes, with subsequent slighter and stronger intensity phases, during the last 2200 years.     

Sexual dimorphism in between and within patch movements of a monophagous insect: Tetraopes (Coleoptera: Cerambycidae)

Summary Movements of milkweed beetles (Tetraopes tetraophthalmus and Tetraopes femoratus) were monitored using mark-release-recapture techniques to compare the within- and between-patch movement patterns of individuals in a naturally occurring population. Within-patch and between-patch movement patterns differ in both frequency and distance traveled for males and females of both study species. Males move farther than females and are more likely to move between patches. Individuals recognize and alter movement upon encountering patch edges. Results suggest that laboratory estimates of Tetraopes vagility give a misleading indication of actual dispersal.     

Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea

The shell of a living specimen of the Indo-Pacific gryphaeid giant oyster Hyotissa hyotis was colonized by numerous encrusting, boring, nestling and baffling taxa which show characteristic distribution patterns. On the upper valve, sponge-induced bioerosion predominates. On the lower valve intergrowth of chamid bivalves and thick encrusting associations—consisting mostly of squamariacean and corallinacean red algae, acervulinid foraminifera, and scleractinian corals—provides numerous microhabitats for nestling arcid and mytilid bivalves as well as for encrusting bryozoans and serpulids. Such differences between exposed and cryptic surfaces are typical for many marine hard substrata and result from the long-term stable position of the oyster on the seafloor. The cryptic habitats support a species assemblage of crustose algae and foraminifera that, on exposed surfaces, would occur in much deeper water.     

Drugs from the seas - current status and microbiological implications

The oceans are the source of a large group of structurally unique natural products that are mainly accumulated in invertebrates such as sponges, tunicates, bryozoans, and molluscs. Several of these compounds (especially the tunicate metabolite ET-743) show pronounced pharmacological activities and are interesting candidates for new drugs primarily in the area of cancer treatment. Other compounds are currently being developed as an analgesic (ziconotide from the mollusc Conus magus) or to treat inflammation. Numerous natural products from marine invertebrates show striking structural similarities to known metabolites of microbial origin, suggesting that microorganisms (bacteria, microalgae) are at least involved in their biosynthesis or are in fact the true sources of these respective metabolites. This assumption is corroborated by several studies on natural products from sponges that proved these compounds to be localized in symbiotic bacteria or cyanobacteria. Recently, molecular methods have successfully been applied to study the microbial diversity in marine sponges and to gain evidence for an involvement of bacteria in the biosynthesis of the bryostatins in the bryozoan Bugula neritina.     

Two levels of spacing and limits to local population density for settled larvae of the ascidian Clavelina moluccensis: a nearest-neighbour analysis

Among sessile marine invertebrates, the dispersion patterns of settling larvae are likely to strongly affect the nature and strength of subsequent intraspecific interactions. We use a recently developed graphical analysis technique to examine in situ spacing among settled larvae of the colonial ascidian Clavelina moluccensis. Larvae were mapped at settlement each day for 1 month at the time of their release from adults. Experimental plots measuring 90×90 mm were located on naturally occurring bare patches of wood on pier pilings which were extensively encrusted with sponges and ascidians. An inclusive one-tailed nearest-neighbour analysis applied at the time of the maximum number of individuals per plot revealed consistent fine-scale regular spacing between settled larvae. Furthermore, a two-tailed goodness-of-fit analysis indicated two modes of spacing: one at about 4.5 mm and the other at around 9 mm. No contact between larvae was observed at settlement and the spatial regularity is interpreted as a response of settling larvae to water-borne cues released from settled individuals. The two modes of spacing indicate the possibility of kin recognition as has been demonstrated for bryozoans. Water-borne cues may also limit local population density, as those plots with a high density of settlers showed a much greater decrease in settlement rate over time than low-density plots. Our findings are consistent with the hypothesis that intraspecific competitive interactions shape the patterns of dispersion of these invertebrates on hard substrata in the field.     

Sub-rubble communities of Curaçao and Bonaire coral reefs

The distribution and abundance of sessile organisms under coral rubble has been studied at Bonaire and Curaçao, Netherlands Antilles. Species richness under rubble is extremely high with at least 367 species of which sponges, tunicates and bryozoans are the most important. Shallow sub-rubble communities can be considered refuges as the majority of these species are crypt-obligate. Sub-rubble communities may also have a preserve function for sponges, but do not harbour enough corals to ensure a quick coral recolonization of the reef surface after a major disaster. Cryptic community composition is affected by depth and pollution, and differs substantially between the two neighbouring islands, possibly as a result of different bottom characteristics. Biomass of the sub-rubble communities may contribute considerably to total reef biomass. Diversity varies inversely with increased depth and increased rubble size, possibly indicating abiotic control (e.g. physical disturbance by wave action and reef slope substrate collapse).     

Movement patterns of Rhyssomatus lineaticollis Say (Coleoptera: Curculionidae) within and among Asclepias syriaca (Asclepiadaceae) patches in a fragmented landscape

Abstract.  1. Dispersal capabilities of organisms are critical in determining the landscape population structure of species as well as their likelihood of survival in fragmented landscapes. Using mark–recapture techniques on the monophagous weevil Rhyssomatus lineaticollis Say (Curculionidae), within- and between-patch dispersal capabilities, landscape level population structure, and the role of beetle density and host patch characteristics in setting distances, amounts, and timing of dispersal were studied.
2. The data indicate that R. lineaticollis is sedentary, with 50% of recaptured beetles moving < 1 m and the maximum distance moved < 1 km. Within- and between-patch movement of beetles was unrelated to host plant patch characteristics and beetle densities.
3. Despite limited dispersal, R. lineaticollis probably functions as a patchy population in east-central Iowa, U.S.A. because dispersals between patches are common and because all host patches surveyed contained this herbivore, indicating a lack of suitable vacant patches, a prerequisite for metapopulation structure.
4. Between-patch distances are well within the dispersal capabilities of R. lineaticollis , although this may be the result of an increase in the density of patches of its host, Asclepias syriaca , in the landscape over the last 150 years as a result of human disturbance and this species' weedy habit.
5. Metapopulation structure in monophagous prairie herbivores may be most likely in species whose non-weedy host plants form highly predictable resources in space and time, but which are now widely scattered in habitat fragments.     

Larger foraminifera as a substratum for encrusting bryozoans (Late Oligocene,Tethyan Seaway,Iran)

Considering the diversity and abundance of larger foraminifera examined from a wide range of Late Oligocene to Early Miocene palaeoenvironments in the Tethyan Seaway, encrusting bryozoans make extremely little use of their tests as substratum. Significant encrustations by bryozoans were exclusively found on large (ø c. 6 cm), undulating tests of Lepidocyclina spp., on which, however, a remarkable 34 taxa of encrusting bryozoans were recorded. This shallow-water fauna of Chattian age was analyzed in respect of the bryozoan taxa present, colony growth type, and mode of budding, colony size, as well as onset of reproduction. Taxic and morphological similarities between the fossil assemblage and modern faunas encrusting mobile substrata indicate a long history of bryozoans as part of the interstitial habitat, while the tests of certain larger foraminifera may have played a significant role in the evolution of shallow-water bryozoans by providing substrata for encrusting species in otherwise unfavorable environments.     

Space as a limiting resource in freshwater systems: competition between zebra mussels (Dreissena polymorpha) and freshwater sponges (Porifera)

Species interactions between two types of sessile benthic invertebrates, the zebra mussel (Dreissena polymorpha) and freshwater sponges (Porifera), were evaluated in Michigan City IN Harbor in southern Lake Michigan during 1996. The study objective was to define whether competition plays a role in structuring benthic communities using experimental techniques commonly employed in marine systems. Sponges were uninhibited by zebra mussel presence and overgrew zebra mussel shells on hard vertical substrata. In contrast, zebra mussels did not overgrow sponge colonies, but did show an ability to re-capture hard substrata if relinquished by the sponge. The negative affect of sponges on zebra mussels through overgrowth and recruitment suggests interactions that could eventually displace zebra mussels from these benthic communities. However, seasonal reduction of sponge biomass from autumn through winter appears to allow the zebra mussel a periodic respite from overgrowth, preventing exclusion of zebra mussels from the community and allowing these two taxa to co-exist.     

Within- and between-patch variability of predation intensity on the mussel Mytilus trossulus Gould on a rocky intertidal shore in Oregon, USA

Variability of predation intensity is an important cause of spatial differences of community structure and organization in the intertidal rocky shore. Field experiments were conducted to evaluate the within- and between-patch variability of the effects of different types of predators (small invertebrates and birds) on Mytilus trossulus Gould, which occupies an intermediate position in the competitive hierarchy among sessile organisms in disturbance patches within a California mussel (Mytilus californianus Conrad) bed community on the central Oregon Coast. Predation by birds did not significantly affect the mortality of M. trossulus. On the contrary, predation by small invertebrate whelks (Nucella spp.) had a significant effect on M. trossulus mortality. Predation by whelks also caused between- and within-patch variability of mortality of M. trossulus. Within patches, M. trossulus mortality at patch margins was significantly higher than at patch centers only when invertebrate predators were present. Wave exposure did not cause between-patch variability of predation intensity. Between-and within-patch variability of predation intensity may be caused by the variability of supply of whelks from the surrounding mussel mat. The movement of predators between patches and surrounding matrices may play an important role in the patch dynamics of these communities.     

Oviposition site preference inNeacoryphus bicrucis (Hemiptera: Lygaeidae): Responses to the density and dispersion of a single host-plant species

Caged females of Neacoryphus bicrucis(Hemiptera: Lygaeidae) were permitted to settle upon and oviposit within a small or large ragwort patch, each of which contained both maturing (= adult food/water source) and dehisced (=oviposition substrate/resource for nymphs) flower heads. The proportion of females settling in the large patch was significantly greater than for the small patch but not significantly greater than expected based on the proportion of maturing flower heads contained in each patch. Also, the number of egg masses laid in each patch was proportional to the number of females that settled there. However, egg mass size was significantly skewed toward larger numbers of eggs in the larger patch. Within the large patch egg mass size was significantly larger where the number of dehisced flower heads was greater and accounts for the between-patch difference in number of eggs per mass. Thus, female settlement does not demonstrate a resource concentration effect but females do appear to evaluate patches after settlement with regard to suitability for oviposition.     

Sedimentation and composition of wall communities in Alaskan fjords

The effects of suspended sediments on the composition of wall communities in Alaskan fjords were investigated by quantitative assessment of underwater photo-quadrats. In fjords with actively retreating tidewater glaciers, suspended sediment levels were extremely high at the heads and were exponentially lower at the mouths. Fjords without glaciers had low suspended sediment levels throughout. The per cent cover and number of species were lowest where sedimentation was highest, at the heads of glacial fjords. Here the wall communities were dominated by a sparse cover of small serpulid worms. Richer communities comprising mostly algae, sponges, hydroids, bryozoans, tunicates and brachiopods were found in fjords without glaciers, and at the mouths of glacial fjords. There was a positive correlation between water clarity and community composition along a gradient from heads to the mouths of fjords with high suspended sediment levels. Accepted: 11 January 1999     

Competitive dominance among sessile marine organisms in a high Arctic boulder community

In most hard substrate environments, space is a limiting resource for sessile organisms. Competition for space is often high and is a structuring force within the community. In the Beaufort Sea’s Boulder Patch, crustose coralline red algae are major space occupiers. This research determined if coralline algae were competitively dominant over other sessile organisms. To test this hypothesis, overgrowth was documented in terms of “winners” and “losers” on the contact borders between different species. Crustose corallines occurred in over 80% of the observed interactions but were only winners in approximately half of them. Most frequently, bryozoans, tunicates, and sponges were superior competitors over crustose corallines, while at the same time these invertebrate groups were among the least abundant space occupiers.     

Bryozoan assemblages on hard substrata: species abundance distribution and competition for space

Bryozoans are colonial invertebrates that often dominate epibenthic assemblages on the lower surfaces of hard substrata. Competition among neighbouring organisms is usually a critical process regulating biodiversity, and hard substrata have proved to be a suitable model habitat to analyse spatial interactions. We explored the relationships among abundance, species richness, diversity, competitive ability, coverage, available surface, depth and substratum size in an assemblage of bryozoans encrusting pebbles and cobbles in a bank off the eastern mouth of the Strait of Magellan. We also tested whether overgrowth competition can be regarded as hierarchical, and whether the species abundance distribution shows a mode of rare species and a decreasing frequency of increasingly abundant species. Abundance, species richness, diversity and overgrowth competition were highest on the largest substrata. Smaller pebbles tended to be encrusted by the commonest bryozoans, while the rarest species were mainly found on relatively larger clasts. A high proportion of the lower surfaces of most substrata was available for growth. Diversity values of relatively shallow stations were lower than expected under Caswell’s neutral model. Interspecific competition was hierarchical, but the abundance of colonies was not related to the competitive ability of each species. The species abundance distribution was bimodal, with a main mode of rare species and a second one partly composed of relatively abundant bryozoans with poor competitive abilities. This study shows that even in an encrusting assemblage where competition is hierarchical, the weakest competitors persist and the dominant species are far from being capable of monopolizing space.     

Biomedicinals from the phytosymbionts of marine invertebrates: a molecular approach

Marine invertebrate animals such as sponges, gorgonians, tunicates and bryozoans are sources of biomedicinally relevant natural products, a small but growing number of which are advancing through clinical trials. Most metazoan and anthozoan species harbour commensal microorganisms that include prokaryotic bacteria, cyanobacteria (blue-green algae), eukaryotic microalgae, and fungi within host tissues where they reside as extra- and intra-cellular symbionts. In some sponges these associated microbes may constitute as much as 40% of the holobiont volume. There is now abundant evidence to suggest that a significant portion of the bioactive metabolites thought originally to be products of the source animal are often synthesized by their symbiotic microbiota. Several anti-cancer metabolites from marine sponges that have progressed to pre-clinical or clinical-trial phases, such as discodermolide, halichondrin B and bryostatin 1, are thought to be products derived from their microbiotic consortia. Freshwater and marine cyanobacteria are well recognised for producing numerous and structurally diverse bioactive and cytotoxic secondary metabolites suited to drug discovery. Sea sponges often contain dominant taxa-specific populations of cyanobacteria, and it is these phytosymbionts (= photosymbionts) that are considered to be the true biogenic source of a number of pharmacologically active polyketides and nonribosomally synthesized peptides produced within the sponge. Accordingly, new collections can be pre-screened in the field for the presence of phytobionts and, together with metagenomic screening using degenerate PCR primers to identify key polyketide synthase (PKS) and nonribosomal peptide synthetase (NRPS) genes, afford a biodiscovery rationale based on the therapeutic prospects of phytochemical selection. Additionally, new cloning and biosynthetic expression strategies may provide a sustainable method for the supply of new pharmaceuticals derived from the uncultured phytosymbionts of marine organisms.     

Microtaphonomy of bioclasts and paleoecology of microencrusters from Upper Jurassic spongiolithic limestones (External Prebetic,Southern Spain)

Microencrusters and microtaphonomic features of the Oxfordian spongiolithic limestones of the External Prebetic were studied using thin-section analysis. The spongiolithic limestone is a bioclastic-rich packstone with common echinoderm, mollusc and brachiopod remains. The bioclasts show a high fragmentation index and frequent microborings. The encrustation index (E _i) is higher for fragments of serpulids, ammonoids and bivalves, and increases with the initial grain-size of bioclasts. The main microencrusters consist of benthic microbial communities (BMC) and nubeculariids, as well as subordinate calcareous and siliceous agglutinated foraminifera, serpulids and bryozoans. BMC are usually the first colonizers, and encrusting foraminifera mainly appear on bioclasts larger than 2 mm. BMC dominate in well-developed encrustations on upward facing surfaces of larger bioclasts that are also colonized by foraminifera (nubeculariids and Subdelloidina). Bullopora, serpulids and bryozoans are more common on lower surfaces. The fact that the values of encrustation index, encrustation thickness and diversity of the microencrusters increase with the size of bioclasts is related to a higher stability and exposure time of the available bioclastic substrate. The microencruster distribution on upper and lower surfaces of large bioclasts may be related to photic control, space competition and/or predation avoidance.     

Cultivation-independent approaches to investigate the chemistry of marine symbiotic bacteria

Sessile marine animals like sponges, tunicates, and bryozoans are a rich source of bioactive natural products, many of which exhibit potent anticancer activities. However, most of these substances are available in very limited amounts only, which has prohibited further drug development. Recent evidence suggests that symbiotic bacteria might be the true producers of many animal-derived metabolites. In addition to revealing fascinating perspectives for research in marine chemical ecology, these findings suggest new solutions to the supply problem. Although most symbionts remain uncultivated, bacterial production systems might be created by isolating biosynthetic genes from marine metagenomes, and expressing them in culturable bacterial hosts. This review discusses cell-sorting, natural product visualization, and phylogenetic approaches to identify symbiotic producers. In addition, strategies to isolate genes and gene clusters from marine species consortia are described. These techniques have provided insights into the bacterial origin and biosynthesis of polyketides like the onnamides, swinholides, and bryostatins, of peptides including the patellamides, chlorinated dipeptides, and theopalauamide as well as of brominated biphenylethers.     

Seasonality and edge effect determine herbivory risk according to different plant association models

We report evidence of hierarchical resource selection by large herbivores and plant neighbouring effects in a Mediterranean ecosystem. Plant palatability was assessed according to herbivore foraging decisions. We hypothesize that under natural conditions large herbivores follow a hierarchical foraging pattern, starting at the landscape scale, and then selecting patches and individual plants. A between- and within-patch selection study was carried out in an area formed by scrubland and pasture patches, connected by habitat edges. With regard to between-patch selection, quality-dependent resource selection is reported: herbivores mainly consume pasture in spring and woody plants in winter. Within-patch selection was also observed in scrub habitats, influenced by season, relative patch palatability and edge effect. We defined a Proximity Index (PI) between palatable and unpalatable plants, which allowed verification of neighbouring effects. In spring, when the preferred food resource (i.e. herbs) is abundant, we observed that in habitat edges large herbivores basically select the relatively scarce palatable shrubs, whereas inside scrubland, unpalatable shrub consumption was related to increasing PI. In winter, a very different picture was observed; there was low consumption of palatable species surrounded by unpalatable species in habitat edges, where the latter were more abundant. These outcomes could be explained though different plant associations described in the literature. We conclude that optimal foraging theory provides a conceptual framework behind the observed interactions between plants and large herbivores in Mediterranean ecosystems.