International Bryozoology Association - 13th Conference, Concepción, 2004 - Abstracts

The Antarctic bryozoan Melicerita obliqua: Skeletal morphogenesis and growth check lines

     Beate Bader & Priska Schäfer
Institut für Geowissenschaften, Abt. Geologie/Paläontologie,
Universität Kiel, Ludewig-Meyn-Strasse 14,
D-24118 Kiel, Germany

      Melicerita obliqua is a common endemic species on Antarctic shelves. The colonies are bifoliate-flattened and sabre-shaped, and attached with organic rootlets in the sediment. The most significant feature of the skeletal morphology is the segmentation of the colony which indicate annual growth check lines. Formation of check lines occurs as a combination of thickened walls in the proximal part of the nodal autozooid and an oblonged, thin-walled distal part. Both are separated by a back-fold of the frontal cryptocyst resulting in a narrow slit on the cryptocyst. Changes in the length of autozooids and segments indicate seasonal and inter- annual variations in the environmental factors.

     The length of segments varies within a single colony but shows a slight decrease with colony age. Individual colonies of Melicerita obliqua reveal a maximum age of about 45 yr. Seasonal growth patterns in the colony are expressed in segment formation with nodes and internodes and differences in autozooid length. Variations in colony growth rate calculated from length increment of segments depict a pattern of longer and shorter segments corresponding between individual colonies that suggests a high-order cyclicity in primary productivity and sea-ice cover.

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Life, death and fighting at high latitude

     David K. A. Barnes
British Antarctic Survey, High Cross, Madingley road, Cambridge, CB3 0ET
dkab@bas.ac.uk

In earth's history, having two frozen polar regions is unusual. Not only do these regions experience extreme light climates and associated primary productivity, but freezing sea temperatures and seasonally intense UV irradiation. At these high polar latitudes severe wind speeds, wave action, ice scour and anchor ice (as well as massive fresh water runoff and localised anoxia in the arctic) make the nearshore environment the most disturbed anywhere. On land, in fresh water and in the intertidal zone there are few colonist species but just a few meters deeper in the sea there can be rich, diverse and abundant benthos even in shallow water. The severity of the physical environment is reflected in the interactions in the biological sphere. Amongst the most abundant shallow water benthos are lithophyllic polychaetes, bryozoans and sponges and in these communities its overgrow or be overgrown. The organisation of sessile animals is extremely hierarchical: at any given locality one species is overgrown by all others and one species overgrows all others - everyone else occupies a rank in between.

With no keystone predators to remove competitive dominant species only the catastrophically destructive power of ice and waves prevents monoculture of certain species. In ice-sheletered areas, such as crevices the end point of classically envisaged 'succession' can be seen. In these shallow water environments many animal populations display exactly the converse of characters typically associated with the polar regions. The most abundant species of many clades are the rarer broadcast spawners with pelagic larvae, that grow and reproduce fast (for polar animals) are small and have but brief lifespans. Many of these contrasts can be seen in the representatives of just one phylum - the Bryozoa. Rather than the predicted K selected species of deeper waters the shallows are ruled by lightly calcified pioneers. Here ecological and evolutionary success have become very much decoupled. A ~2°C rise, predicted in polar waters, could be enough to transform this unique zone

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Bipolar patterns of intraspecific competition and prevalence of homosyndrome

     David K. A. Barnes
British Antarctic Survey, High Cross, Madingley road, Cambridge, CB3 0ET
dkab@bas.ac.uk
     Piotr Kuklinski
University Courses on Svalbard (UNIS), P.O.Box 156, Longyearbyen N-9171, Norway.
Institute of Oceanology, Polish Academy of Sciences, Marine Ecology Department, ul. Powstanców Warszawy 55, Sopot 81-712, Poland

Polar shores probably represent the most dynamic and intensely disturbed environments on the globe. Nevertheless intense battles amongst sessile organisms for space are commonplace on hard substrata. Typically assemblages are very young and competitively inferior but fast growing species occupy most space. As a result most competition tends to be intraspecific, though it is interspecific competition which has received the great majority of scientific attention. In this study we examine intraspecific encounters in numerically dominant, encrusting species at the high arctic latitudes of the Faeroes (63°N), Hornsund (77°N) and Ny Alesund (79°N) in Spitsbergen. We also measured the same data at the Antarctic localities of Signy (60°S), Dobrowlski (65°S) and Adelaide islands (68°S). Earth's two polar regions differ substantially in geological history, geography, bathymetry and biodiversity, but have some important similarities - competition structure seems to be one of them. We found strong similarities in the importance, nature and outcome of intraspecific interactions. In each location the vast proportion of intraspecific interactions occurred in just one species. Most encounters involved colonies meeting 'head-on' and resulted in ties rather than decided (win or loss) results. The relatively fast growth of the pioneers dominating all six study sites meant that many individuals experienced crowding, restricted growth and accelerated ovicell production due to intraspecific competition. Homosyndrome (fusion) was rare but their frequency differed significantly between competitor identities. We found that aggressive species (with high interspecific rankings) were more likely to tie in interspecific meetings and undergo homosyndrome. Furthermore species in higher water flow environments were less likely to show tied interactions or homosyndrome. We interpret these findings and their implications in the context of changing disturbance and climate patterns.

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Late Neogene bryogeography of southern Spain

     Björn Berning
Geologisch-Paläontologisches Institut, Universität Hamburg, Bundesstr. 55, 20146 Hamburg, Germany
     Pierre Moissette
Université de Lyon I, UFR Sciences de la Terre/UMR CNRS PEPS, 27 Bd du 11 novembre, 69622 Villeurbanne Cedex, France
     Christian Betzler
Geologisch-Paläontologisches Institut, Universität Hamburg, Bundesstr. 55, 20146 Hamburg, Germany
Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, UK

     Even 30 years after the discovery of the latest Miocene desiccation of the Mediterranean Sea, an episode known as the Messinian salinity crisis, the circumstances of this impressive event are a subject of much debate. Since Mediterranean bryozoans show a great diversity and are well preserved in late Neogene sediments, they have already played an important role in deciphering evolutionary and biogeographical consequences of the crisis to Mediterranean faunas. Nevertheless, many questions concerning the paleoceanography or paleobiogeography remain as yet to be answered. One major drawback lies in the fact that, while bryozoans from the Mediterranean Sea itself are quite well known, there are merely few data from regions west of the Strait of Gibraltar, i.e. of the Atlantic realm, which might have served as a refuge for the Mediterranean biota during the crisis. Here we present hitherto undescribed bryozoan faunas from southern Spain of the Atlantic Guadalquivir Basin (late Tortonian) as well as the Mediterranean Agua Amarga Basin (Tortonian/Messinian) and Carboneras Basin (Lower Pliocene) to record environmental and biogeographical disparities in these different regions before and after the crisis.
     Our new systematic and biogeographic data allow us to address several aspects. (1) The occurrence of species of Mediterranean affinity (e.g. Myriapora truncata) in sediments of the Guadalquivir Basin provides evidence for a westerly direction of surface water flow through the Rifian and/or Betic gateways before the onset of the crisis. This observation is confirmed by Tortonian-Messinian current-indicative sedimentary structures preserved within the Spanish straits. The late Miocene dispersal of taxa thus differs from the modern situation in that today a westwards migration of shallow-water Mediterranean biota is prevented by the constant inflow of surface water from the Atlantic through the Strait of Gibraltar. (2) The fact that several species endemic to the Mediterranean survived the salinity crisis indicates the presence of a contemporary refuge providing environmental conditions sufficiently similar to the Mediterranean Sea. Since the bryozoan fauna of the Guadalquivir Basin is comparable to the Mediterranean ones, this basin qualifies to have served as such a retreat. However, some species of the Guadalquivir Basin differ significantly in colonial or zooidal morphology from their neighbouring Mediterranean representatives which might suggest environmental boundary conditions for the existence of some Mediterranean species in this peripheral region. (3) The huge set of data resulting from past and ongoing research on Neogene bryozoans from the Mediterranean realm, in which over 400 species are known to occur, provides a sound basis to statistically evaluate the processes and patterns of evolution, extinction and migration. In particular, the valuable information that we have gained from a marginal part of the late Miocene Mediterranean Sea, the Guadalquivir Basin, allows us to reconstruct the evolutionary history of Neogene to Recent Mediterranean bryozoans and the impact of the Messinian salinity crisis on the biota of the region.

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Upper Devonian to Lower Carboniferous Bryozoa in Central Hunan (S. China)

     Françoise P. Bigey
Université P. & M. Curie (Paris VI)
Département de Géologie Sédimentaire, Laboratoire de Micropaléontologie
C. 104. 4, Place Jussieu F-75252 Paris-Cédex 05, France

The considered region of Central Hunan Province is located in the Lengshuijiang-Xinshao-Qiyang area. It consists of a part of the tectonic South China Fold System. Continuous sedimentation during late Paleozoic has to be underlined. Consequently this area is especially valuable for study of Frasnian-Famennian event and Devonian-Carboniferous transition beds. Conodonts and foraminifers chiefly give precise biostratigraphical data. The littoral and shallow environment favours widely the development of benthic fauna: rugose corals, stromatoporoids, brachiopods, echinoderms in addition to bryozoa. According to the preservation conditions, accuracy in identification of zoaria remains unsteady.
     From the four sections studied by Chinese and Belgian biostratigraphers, three of them have yielded bryozoa. Laojiangchong section: Frasnian to Famennian, Oujiachong section: Famennian and Malanbian section: Famennian to Tournaisian. Within the Laojiangchong section, the late Frasnian Laojiangchong Formation corresponds to the richest levels for bryozoa: numerous ramose trepostomate colonies are present. Higher in the section, within Tuzitang Limestone (base of the Famennian Xikuangshan Group), bryozoa are more diverse: trepostomes (branched or encrusting) and rhabdomesids. Within the Oujiachong section the last levels of the Magunao Limestone (top of the Famennian Xikuangshan Group) bryozoa are chiefly rhabdomesids, but some encrusting trepostomes exist. Within the Malanbian section, bryozoa are known from nearly all Formations. Within the Famennian Shaodong Formation, the scarce bryozoa are trepostomes and rhabdomesids. Within the uppermost Famennian Menggongao Formation are found some encrusting trepostomes and rhabdomesids; fenestellids appear. Within the Tournaisian (Hastarian) Tianeping Formation, fenestellids dominate ramose trepostomes. Within the Tournaisian (Ivoirian) Doulingao Formation, the last Carboniferous one of the area bryozoa are more diverse, fenestellids, rhabdomesids and few trepostomes and fistuliporids.
     Value is attached to occurrence of bryozoa in Famennian levels, as in South China, because rare elsewhere. Meanwhile bryozoa are not directly involved in the Frasnian-Famennian event and the Devonian-Carboniferous boundary because lacking in the concerned strata.

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Basal attachment structure of Nudicella cribriforma Schmidt & Bone in the Miocene of Tasmania, Australia, and its similarity to Modern Adeona spp. from southern Australia

     Yvonne Bone
Geosciences, School of Earth & Environmental Sciences, University of Adelaide, Australia, 5005
     Rolf Schmidt
Museum Victoria, PO Box 666E, Melbourne, Victoria, Australia, 3001

     The large onychocellid cheilostome bryozoan, Nudicella cribriforma Schmidt & Bone (in press) has until early 2003, only been found as fragments of the feeding portions of the erect bilaminar foliaceous colony, i.e. pieces with zooids on a cribrate fenestrate sheet. This architecture is reminiscent of Modern Adeona Lamouroux species, with this similarity particularly relating to the large fenestrules (up to 2 mm in diameter) present in both taxa.
     There had been no recovery of definite basal attachment structures of any Nudicella until recently. Adeona spp. are amongst the most successful bryozoans in terms of weight and/or volume in their preferred environment of mid- to outer-shelf unconsolidated sediments, particularly at depths of 60 - 120m. This success probably stems from its complex attachment structure, which is a robust, trunk-like, kenozooidal basal section that is usually buried up to 5 cm into the sediment. This trunk becomes branch-like in the section of the colony above the sea floor, whereas the subsurface section is root-like, with the tips of the roots extending into the sediment to a depth of a metre or more, as mucilaginous strings. The kenozooids in the trunk and root sections are articulated by connecting chitin-like material. The whole attachment process allows the bryozoan to be a pioneer exploiter of an open-shelf, high-energy environment that is otherwise poorly colonised by sea-floor biota. It then acts as a substrate for secondary settlers, e.g. the cellariids and articulated zooidal forms. The presence of a hardground, however, sees Adeona cement itself to the substrate, even if it is only a few cm in length, thereby not producing the bulk of the roots nor the mucilaginous strings, but still retaining the flexibility of the articulated kenozooids of the trunk section.
     The cribrate growth form occurs commonly and convergently throughout the bryozoan fossil record, so we assumed that N. cribriforma had a rigid basal structure similar to most other fossil cribrate species, although it had never been found. Then in February, 2003, at Marramah on the western coast of Tasmania, in the Miocene limestone, a large specimen of N. cribriforma was found, complete with the kenozooidal attachment structure and the anastomosing roots. It was associated with the same diverse fauna that we find it associated with in South Australia, which is similar to the modern southern Australian shelf.
     Adeona uses aragonite to produce its skeletal structure today. N. cribriforma did not appear to use aragonite, as the specimens are generally not dissolved like those other co-occurring organisms which did use aragonite, even when the colony has fragmented into individual zooids. This raised the question of why none of the basal trunk sections have previously been found, as they should survive diagenesis, unless the colony was bi-mineralic and used aragonite for the basal section.

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Cyclostome bryozoans collected during historical Antarctic expeditions are now providing geochemically-determined oceanographic temperature data.

     Elizabeth M. Campbell & Yvonne Bone
Geosciences, School of Earth & Environmental Sciences, University of Adelaide, Australia, 5005

     Geochemical analyses of δ¹⁸O values from calcareous fossils are widely used to calculate the paleotemperatures of the ambient ancient ocean water. Brachiopods and suitable foraminfers are commonly used, but these are not always available. It is also possible, however, to use low-Mg calcite (LMC) bryozoans as proxies for oceanographic parameters. Cyclostome bryozoans predominantly use LMC for their skeletal elements so this makes them especially useful.
     None of the expeditions to Antarctica in the period 1901 to 1937 had the collection of bryozoans as a major goal. These expeditions were financed for entirely different reasons, e.g. the BANZARE expeditions (Mawson et al.), the Discovery cruises, Scott's cruises, were investigating krill for the whaling industry, investigating and mapping onshore areas, producing navigation charts and Empire-building. Nevertheless, these leaders had the foresight to dredge sea-floor for biota and sediments whenever the opportunity arose. They lived at a time when the collection and diligent archiving of the material obtained was considered a "proper pastime" for all educated gentlemen, and indeed, also a fascinating one.
     So, the world has repositories of historically-important biota collections that have been subjected to at best, a first-pass sorting. Few of the collections have been studied to the species level. Instead, they have been sitting in such locations as the basements of Museums around the world, awaiting the right time to be accorded their place in the scientific assessment and cataloguing of the diversity, distribution and density of the marine biota.
     It is in such Antarctic collections we have found a treasure-trove of cyclostome bryozoans. Our research has been:

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Ecological observations on shallow water marine bryozoans in King George Island, Antarctica

     Juan M. Cancino
Departamento de Ecología Costera, Facultad de Ciencias, Universidad Católica de la Santísima Concepción. Alonso de Ribera 2850, Concepción, Chile
     Hugo I. Moyano
Departamento de Zoología, Facultad de Ciencias Naturales y Oceanográficas, Universidad de Concepción
     Patricio H. Manríquez
Estación Costera de Investigaciones Marinas, Facultad de Ciencias Biológicas, Pontificia Universidad Católica de Chile

     With the aim of contributing to the knowledge of both, bryozoan species diversity and reproductive biology of some selected species of shallow water bryozoans in Antarctic waters, different substrata were sampled in Fildes Bay (62°, 11' 52" S, 58° 55' 57"W). Samples were collected by scuba diving in January 2003, at depth ranging between 7 and 20m, brought to the laboratory and kept in running seawater until studied under the binocular microscope.
     A total of 28 species, 7 Cyclostomes and 21 Cheilostomes were found. Bryozoans occurred on red algae (2 to 8 species, depending on algal species); on foliose Bryozoa (3 to 11 species), on invertebrates (Porifera, Brachiopods, solitary ascidians and Hydrozoa, (2 to 7 species) and on rocks (9 species). Dominant species were Inversiula nutrix on rocks, Celleporella bougainvillei and Osthimosia milleporoides on animals and algae, Celleporella antarctica on thin filamentous substrata. Celleporella bougainvillei was widely distributed on red seaweed, while Celleporella dictyota was found mainly on the stipes of Desmarestia sp. Fourteen out of the 21 Cheilostome species were endemic to Antarctica (66.6% endemism), while the other 7 are also present in the Magellan area.
     Size at first reproduction was smaller in Celleporella antarctica (8 autozooids) than in Celleporella bougainvillei (19 autozooids). Fecundity (sexual zooids/autozooids) was under 0.2 for Nematoflustra flagellata; ranged between 0.05 and 0.37 for C. bougainvillei and between 0.1 and 0.75 for C. antarctica. These results are discussed in relation to the nature of the substratum used by the different species and compared with published information of similar species in temperate waters. INACH Project 05/97.

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Survival to total overgrowth in encrusting bryozoans

     Juan M. Cancino & María Cristina Orellana
Departamento de Ecología Costera, Facultad de Ciencias, Universidad Católica de la Santísima Concepción. Alonso de Ribera 2850, Concepción, Chile.

     Overgrowth is common among colonies of encrusting bryozoans. Total colonial overgrowth occurs mainly when small colonies of species get into contact with the fast growing edges of larger colonies. The totally overgrown colony has usually been regarded as dead. In the present study we determined whether the colonies totally smothered by others are actually dead.
     Encrusting bryozoans were grown in 132 cm² glass slides kept during 3 months in the subtidal, allowing natural settlement and overgrowth to take place. The glass slides were photographed weekly and afterwards all the colonies that had been totally overgrown were exposed by carefully removing the overgrowing colonies. We registered the initial number of totally overgrown colonies per species and the number of such colonies having active zooids 1 week after the experimental removal of the smothering colony.
     Three hundred thirty five total overgrown colonies, of the 7 commonest encrusting bryozoans species in central Chile, were studied. Between 28 and 80% of the colonies of these species survived to total overgrowth. Ability to survive total overgrowth was present in species of a wide range of competitive hierarchies, suggesting that this is not a mechanism present only in species that are more likely to be overgrown due to their low competitive hierarchy. According to the present results, totally overgrown colonies should be regarded as dormant rather than death. More information is required to assess the role of such dormancy in encrusting bryozoans community structure.

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Bryozoan Species and Possible Sedimentologic Roles in Small Waulsortian-Like Mud-Mound Biohermsin the Mississippian (Lower Carboniferous) of the American Mid-West

     Roger J. Cuffey
Department of Geosciences (412 Deike Bldg.), Pennsylvania State University, University Park, Pennsylvania 16802, USA

     Bryozoan reefs have a lengthy geologic history (Ordovician - Recent), during which their importance has fluctuated greatly. The earlier Mississippian (earlier Lower Carboniferous) was one time when such structures were conspicuous, in the form of mud-mounds in part possibly baffled or stabilized by fenestrate bryozoans, deposits long known in western Europe as the Waulsortian facies.
     Similar, but not identical (hence instead often termed "Waulsortian-like"), bioherms have been studied in the Mid-West (from Kentucky to Missouri, and Illinois to Tennessee). These American build-ups, in contrast to the European ones, are smaller-sized, coarser-grained (calcisiltite), in shallower paleoenvironmental settings, and include not only carbonate but also argillaceous compositions.
     Bryozoan fossils are scattered through these mounds, mostly fragmentary, lying frontal-surface downward and horizontally parallel to bedding, but locally intact and preserved still standing upright within the entombing mud sediment. In total, 62 bryozoan species (out of the several hundred described from the region in similar-age rocks) have been identified from the mounds examined, mostly delicate fenestrates, in places accompanied by bifoliate fistuliporoids, and occasionally by tiny rhabdomesids. Two species in particular occur in all (Exfenestella exigua) or most (Banastella regalis) of the mounds investigated. Several more species are rare but found in many of the mounds, while others are both rare and in few of the bioherms. Sedimentologic or constructional roles envisioned for these bryozoans include possible baffling or trapping mud from suspension, stabilizing or anchoring already-deposited unconsolidated sediment, and probable locally contributing minor skeletal debris to the accumulating carbonate sediment.
     These small mud-mounds furnish interesting comparisons and contrasts to other bryoherms, especially Ordovician crust-mounds and Permian frame-thickets.

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Actualized biological definition of the Bryozoa

     Jean-Loup L. d'Hondt
Muséum National d'Histoire Naturelle, USM 0403 « Biodiversité et dynamique des communautés aquatiques », Département « Milieux et peuplements aquatiques », (Biologie des Invertébrés marins), 55, rue de Buffon, F - 75231 Paris Cedex 05

     Reactualized definition of the phylum Bryozoa, after critical discussion of their main biological characters by the light of the more recent knowledges on this taxon :
     Colonial animals which are morphologically and anatomically unsegmented, triploblastic, with embryonic endomesoderm and generally abortive endodermic macromeres; endocoelomates without archimery, in the larva as in the adult; two-layered - ectodermic and mesodermic - tegument, each layer with peculiar morphological and ontogenetical capacities, on the one hand in the larva, on the other hand in the adult; with chitinous or chitino-calcareous exoskeleton; embryologically with protostomian larva, but atypical deuterostomian ancestrula - and by the way it is also the case for the other adult individuals, the autozoecia - ; neither hyponeurian nor epineurian in the adult stages; of which some epidermical cell lineages, true stem cells, keep a given and permanent totipotence from the larval metamorphosis to the death of the zoarium, this latter proceeding by clonal development from a single larva; presenting, according to the different phylogenetic trends, various types of ontogenetic capacities of morphogenetical substitutions; of which capacities of epidermic cells are polarized; where each neopolypidial morphogenesis is an automatic biological phenomenon, probably under hormonal influence; devoid of some physiological systems (respiration, circulation, excretion); with adult tentacles from ectodermic origin; ectoprocts; with generally an intermittent digestive system (= main part of the polypid), periodically degenerating and renewed from an epidermic proliferation to the inside (consequently : 1 - the adult epidermis has kept some morphogenetical capacities of a gastrula; 2 - the budding - by disappearing of an inhibition factor - from an ectodermis, of an organ with a digestive vocation, is an infraction to the classical theory of the embryological layers; with digestive system U-shaped, without digestive gland; without coelom in the larva, the coelomic cavity appearing by schizocoelie only during the metamorphosis.
     Some evolutive trends genetically « programmed » and predetermined, materialized by embryological apoptosis and pre-differentiations, events obvious from the embryological stages. Elaborated coloniality (pore plates and funicules-rosettes system); presenting both capacities of asexual and sexual reproduction and a very complicated metamorphosis. The species are mainly sedentary and benthic, exceptionally free-living; always aquatic, generally marine; filter-feedings with prevalent vegetalian diet. Often an interzoecial polymorphism corresponding to a functional specialisation of the individuals.

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The Historical collections of Recent Bryozoa in the French National Collections

     Jean-Loup L. d'Hondt
Muséum National d'Histoire Naturelle, USM O403 « Biodiversité et dynamique des communautés aquatiques », Département « Milieux et peuplements aquatiques » (Biologie des Invertébrés Marins), 55, rue de Buffon, F - 75231 Paris Cedex 05

     The collections of the Muséum National d'Histoire Naturelle of Paris contain hundreds lots of specimens regarded as precious from a historical point of view, because they have been collected by ancient and prestigious naturalists (as Vaillant, Lamarck, Lamouroux, Péron, Lesueur, Quoy, Gaimard, Henri Milne Edwards or Jullien), and often containing numerous type-specimens from the XVIIIth or XIXth centuries, or arising from famous later circumnavigations (for example, voyages of Baudin, Dumont d'Urville or Freycinet). This historical material is presented here under the names of the authors having collected, offered or valorized the collections.

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Bryozoan-sponge associations to bryozoan stable carbonate lithoskels and silica spicules accumulations, to friable limestones with flint layers

     Linda Deer & Yvonne Bone
Geosciences, School of Earth & Environmental Sciences, University of Adelaide, Australia, 5005

     The Cretaceous chalk of Europe has long been noted for its extensive stratigraphic-like flint horizons. This phenomenon is also observed in many other limestones throughout the Cenozoic, e.g. the Eocene Wilson Bluff Limestone of South and Western Australia and the Gambier Limestone of Victoria and South Australia. All the limestones have some notable features in common:
     they are invariably friable, indicating minimal carbonate diagenesis,
     they are bryozoan-rich, indicating they were formed in a cool-water or polar environment,
     the majority of the bryozoans present are genera that today use low-Mg calcite (LMC) for their skeletons,
     the majority of these bryozoans today live in deeper water, i.e. below 100m. These flint layers are not present in all bryozoan-rich limestones. Some may have irregularly-spaced chert nodules, but have not developed spectacular flint horizons. These "limestones" are often more marls than true limestones, and will also often contain partially-silicified sponge body fossils. Yet other bryozoan-rich limestones do not have any such features but these are usually relatively shallow-water mollusc-rich units, with a diverse assemblage of accessory biota.
     We proposed that the spicules from siliceous sponges were the source of the silica that forms the flint, not quartz-rich terrigenous input during low-stands or silica-rich groundwaters during diagenesis. Our research looked at the percentage of silica spicules contained in living sponges, on a weight/weight basis. The living sponges were collected from a range of depths from the southern Australian continental margin. The sponge material was frozen upon collection so that no changes would occur prior to laboratory testing. Ascidians were also collected from the same sites, as these also contain spicules, but these silt-sized mace-shaped spicules are aragonitic. Aragonite is metastable and readily dissolves during early diagenesis.
     The results showed that there is an increase in spicules in sponges (weight/weight) as the water depth increases. Thus, as the organic fraction of the sponge decays after death, there is a greater source of silica spicules incorporated into the accumulating sediment. These same sediments coincide with those with the higher percentage of bryozoan fragments. Furthermore, these bryozoans belong to genera that use LMC for their skeletal elements. The ascidian spicules show the same depth behaviour.
     Silica is metastable in the presence of high pH waters. The ascidian spicules and the minor bryozoans using metastable carbonates for their skeletons would buffer any circulating waters, thus maintaining a high pH, enabling the dissolution of the sponge spicules.
     We propose that later in the geological record, the alkalinity of the water changes and the silica precipitates at that horizontal level as a layer of flint. So, the association of living bryozoans and sponges, and ascidians to a lesser degree, has important implications for the rock record.

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Environmental impact of the river Nile on the Recent bryozoans of the northern coast of Egypt

     Yasser A. El Safori
Department of Geology, Faculty of Science, Ain Shams University, P.O. Box 11566 Cairo, Egypt

     Two marine biofacies for the Recent bryozoans of the northern coast of Egypt are recorded. The negative impact of the fresh waters and clastics of the River Nile downstreams is recorded from the coastal sediments of the Delta and to the East (East biofacies). This effect is completely minimized to the West of the Nile Delta (West biofacies) for being far from the Nile downstreams and their lateral water drifts. The coastal sediments of the West biofacies are composed of carbonate grains rich in erect and liberated encrusting bryozoans. Also, encrusting bryozoans grow upon different substrates of molluscan shells, rock fragments, and plant stems. However, encrusting anascans bryozoans of low diversity, characterize the East biofacies. The coastal sediments of the East biofacies are composed of quartz sands with rare/or no erect bryozoans. Also, the encrusting bryozoans are collected from the molluscan shells and wave barriers. Across the downstream straits (brackish waters), only encrusting colonies of Membranipora lacroixii are recorded on the wave barriers. The population abundance of the encrusting bryozoans is related to other encrusters as calcareous tubeworms and barnacles.

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Oligocene bryozoans From al Jabal Akhdar, Libya

     Yasser A. El Safori
Geology Department, Faculty of Science, Ain Shams University, P.O. Box 11566 Cairo, Egypt
     Ahmed M. Muftah
Geological Department, Geological Laboratory, Arabian Gulf Oil Company, P.O. Box 263 Benghazi, Libya

     Fifteen bryozoan species are separated from the Oligocene sediments (Faidiya Formation) of al Jabal Akhdar, East Libya. They represent one of the minor records of the marine Oligocene bryozoans of North Africa. The studied assemblage are mostly erect cheilostomatous species, associated with reefal fauna of benthic foraminifers and ostracods.
     The paleobiogeography of the studied bryozoans indicates their East Atlantic/Mediterranean affinity. The paleoecology of the studied bryozoans is discussed along with the studied lithofacies.

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Bryozoans from the Artinskian (Lower Permian) Great Bear Cape Formation, Ellesmere Island (Canadian Arctic)

     Andrej Ernst
Institut für Geowissenschaften der Christian-Albrechts-Universität zu Kiel,
Ludewig-Meyn-Str. 10, D-24118 Kiel, Germany
     Hans Arne Nakrem
Geological Museum, University of Oslo, PO Box 1172 Blindern, NO-0318 Oslo, Norway

     During the 1898-1902 «Fram» expedition to the Canadian Arctic islands, scientific material was collected and in part described in more than 27 reports. Geological material was collected by P. Schei, and some Upper Palaeozoic bryozoans from the expedition were described by T. Tschernyschew and P. Stepanov in 1916. The current investigation is based on a small collection housed in the Geological Museum (Oslo) from Great Bear Cape, Bjorne Peninsula, SW Ellesmere Island. The stratigraphic unit has for a long time been known as «Unknown Fm.», of uncertain age. Recent fieldwork and conodont investigations by B. Beauchamp and C. M. Henderson have made it possible to revise the stratigraphy here, and the lithostratigraphic unit is now known as the Great Bear Cape Formation of Artinskian-Earliest Kungurian (Early Permian) age. Sedimentation took place on a carbonate shelf under regressive conditions, - the lithology consists of resistant, yellowish-weathering, pure to locally sandy, variably cherty, highly fossiliferous limestone. Remnants of bryozoans as well as other animals reveal numerous borings by unknown organisms. The bryozoan fauna comprise the following taxa: Cystoporida: Fistulipora volongensis Nikiforova 1938, Fistulipora sp., Cyclotrypa distincta Morozova 1986, Ramiporidra variolata Shul'ga-Nesterenko 1933. Trepostomida: Hinaclema sp., Tabulipora spp., Rhombotrypella ? amdrupensis Ross & Ross 1962, Dyscritella vulgata Gorjunova 1972, Dyscritella tenuis Kruchinina 1973. Ulrichotrypa ramulosa Bassler 1929. Cryptostomida: Streblascopora vera Morozova 1986, Streblascopora germana (Bassler 1929), Clausotrypa monticola (Eichwald 1860), Primorella ? superba Morozova 1981, Primorella tundrica Kruchinina 1986, ?Rhombopora sp., Bashkirella operculata Shul'ga-Nesterenko 1952. Cryptostomida (Intraporidae): Phragmophera sp. n. Fenestrida: Alternifenestella bifida (Eichwald 1860), Alternifenestella crassiseptata (Shul'ga-Nesterenko 1941), Alternifenestella cyclotriangulata (Eichwald 1860), Alternifenestella cf. invisitata Kruchinina 1986, Fabifenestella cf. subvirgosa (Shul'ga-Nesterenko 1952), Fabifenestella cf. virgosa (Eichwald 1860), Fabifenestella tortuosa (Trizna & Klautsan 1961), Fenestella akselensis Nakrem 1995 [1994], Rectifenestella microporata (Shul'ga-Nesterenko 1939), Rectifenestella robusta (Shul'ga-Nesterenko 1936), Polypora confirmata Kruchinina 1986, Polypora kossjensis Ravikovich 1948, Polypora kutorgae Stuckenberg 1895, Polypora martis Fischer 1837, Polypora voluminosa Trizna & Klautsan 1961, Penniretepora invisa (Trizna 1939), Acanthocladia cf. sparsifurcata Shul'ga-Nesterenko 1939, Acanthocladia cf. rhombicellata Shul'ga-Nesterenko 1955.
     Beside these taxa, two more bryozoans are problematical in their taxonomic assignment belonging apparently to trepostomes and cryptostomes respectively. Phragmophera sp. n. is until now the youngest known representative of bifoliate cryptostomes.
     The taxonomic composition bears a striking similarity with Sakmarian-Artinskian faunas known from Timan-Pechora area (Western Siberia, Russia) and the Artinskian-Kungurian of Svalbard, the Early Permian of NW Greenland, as well as some Early Permian faunas previously described from Ellesmere Island.

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Lower Carboniferous Bryozoa from some localities in Sauerland (Germany)

     Andrej Ernst
Institut für Geowissenschaften der Christian-Albrechts-Universität zu Kiel, Ludewig-Meyn-Str. 10, D-24118 Kiel, Germany

     Lower Carboniferous bryozoans from Germany are scarcely investigated because of their mostly poor preservation. Only few papers were published about this fauna, with restricted use of oriented thin sections. The most extensive study has been performed by Nekhoroshev (1932). However, some newly investigated localities in Sauerland (Westenfeld, Hellefeld, and Frenkhausen) revealed a number of bryozoans in excellent, mostly calcitic preservation. Material was sampled by Dieter Weyer and Dieter Korn (both Berlin) from rocks dated by Goniatites crenistria and Arnsbergites gracilis zones of the Lower Carboniferous. Lithology is characterized by turbidid sediments, bearing numerous organic remnants (bryozoans, brachiopods, calcareous algae etc.).
     About 50 thin sections were prepared from the available material. Following bryozoan taxa could be identified during the present investigation: Fistulipora incrustans (Phillips 1836), Evactinopora sp., Stenopora sp., Nipponostenopora ? sp., Stenodiscus redesdalensis (Lee 1912) (= S. tumida; Wyse-Jackson, pers. comm.), Rhabdomeson progracile Wyse Jackson & Bancroft 1995, Rhabdomeson regulare Nekhoroshev 1932, Nematopora hibernica Wyse-Jackson 1996, Streblotrypa pectinata Owen 1966, Rhombocladia cf. dichotoma (M'Coy 1844). The rock contains also numerous unidentified remnants of fenestellid bryozoans.
     The faunal composition shows a close relation to the Lower Carboniferous of the British Isles. This is the first record of the genus Evactinopora Meek & Worthen 1865 from the Lower Carboniferous of Europe. The genus Evactinopora is a typical fossil of the Lower Carboniferous of Northern America and Western Australia. Elsewhere it was reported from the Middle Carboniferous of SE Russia (Morozova 1981). The species Rhabdomeson regulare Nekhoroshev 1932 was also reported from the Lower Carboniferous of Russia (Morozova 1955)

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Paleobiogeography of Middle to Late Permian Bryozoans

     Ernest H. Gilmour
Department of Geology, Eastern Washington University, Cheney, WA, 99004, USA
     Iraida P. Morozova
Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117647, Russia

     Middle to Late Permian bryozoans, which belong to 119 genera, 29 families, and 7 orders, lived in all climatic zones. These bryozoan genera occur in 19 provinces of 6 regions. New unpublished data from New Zealand, Pakistan, Cordilleran of western U.S., and Texas are included in this summary. The majority of bryozoan genera in the Permian are cosmopolitan, but several are confined to boreal, tropical, or notal climatic zones. Some genera normally found only in the Euro-Canadian boreal zone occur in the Cordilleran province; whereas the bryozoan fauna of the New Zealand province includes both notal endemic genera as well as genera that are in common with or very close to bryozoans of the boreal province. These genera represent a bipolarity in the evolution of cold-water faunas during the Middle to Late Permian.
     The peak of taxonomic diversity occurred in the Wordian (Early Kazanian) with a considerable number of highly specialized genera emerging and then becoming extinct by the end of the Capitanian (Late Kazanian). Extinction of genera continued during the Wuchiapingian and Changhsingian Stages with only four genera of bryozoans (trepostomes) surviving into the Triassic. This gradual reduction of bryozoan genera during the Middle and Late Permian and the survival of at least four genera into the Triassic does not support a sudden extinction event at the end of the Changhsingian for this group of organisms.

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The cheilostomatous genera of Alcide d'Orbigny - nomenclatural and taxonomic status

     Dennis P. Gordon
National Institute of Water Atmospheric Research, P.O. Box 14-901 Kilbirnie, Wellington, New Zealand
     Paul D. Taylor
Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

     Alcide Dessalines d'Orbigny was the author of 75 genera of Cheilostomata, 73 of which were introduced in his volume on Bryozoa in the Paléontologie française series Terrains crétacés. A recent SEM study of as many type and other specimens pertaining to these genera has made possible the clarification of the nomenclatural and taxonomic status of the genera. Discarding those genera for which type species and specimens are lost and which may never be properly understood, we regard 40 as having validity. These include 16 previously overlooked or ignored in Bassler's (1953) Treatise revision, viz Filiflustrina, Latereschara, Lateroflustrella, Multescharellina, Multescharipora, Pyriflustrina, Reptescharella, Reptoflustrella, Reptolunulites, Reptoporella, Reptoporina, Reptescharinella, Semiescharipora, Semiflustrella, Semiflustrina, and Reptescharipora.
     These genera are reinstated, based on identifiable type species. We give new diagnoses for each of these genera and classify them to family level according to current phylogenetic concepts of the order.

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A new genus of Umbonulidae (Bryozoa: Cheilostomata) from the northwest Pacific

     Andrei V. Grischenko & Shunsuke F. Mawatari
Laboratory of Systematics and Evolution, Division of Biological Sciences, Graduate School of Science, Hokkaido University, Sapporo 060-0810, Japan

     A new genus and species of Umbonulidae is described from two areas of the northwestern Pacific: Ptichii Island (Western Kamchatka shelf of the Sea of Okhotsk) and Bering Island (Commander Islands, the Bering Sea). The colony is encrusting. Zooids are umbonuloid with a solid umbo and the secondary orifice is cormidial. There are no suboral avicularia, ovicells or oral spines. The new genus is close to Umbonula in having an umbonate frontal shield with marginal areolae separated by elongated ridges; however the new species differs from U. ovicellata Hastings, 1944 (the type species of Umbonula) in the cormidial nature of the secondary orifice and the absence of a suboral avicularium and ovicells. Discovery of the new genus and species justifies splitting the genus Umbonula Hincks, 1880. In a newly suggested classification, each segregated genus includes several species, avoiding monotypy. Accordingly, U. littoralis Hastings, 1944 and U. inarmata Kluge, 1962, together with the newly described species, are included in the new genus. The nominate genus is here taken to include U. patens (Smitt, 1868), an undescribed species from the Commander Islands as well as the type species. Whereas Arctonula Gordon & Grischenko, 1994 was segregated from Umbonula and removed to the Romancheinidae, the new genus is accepted as a member of the family Umbonulidae even though it lacks ovicells. The new taxon occurs predominantly intertidally but ranges to 25 m, on hard substrata, and can be categorized as a Pacific high-boreal species.

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Diversity, evolution and paleoecology of the Tertiary bryozoan assemblages of western Kachchh, Gujarat, India

     Asit K. Guha & K. Gopikrishna
Department of Geology & Geophysics, Indian Institute of Technology, Kharagpur-721 302, India.
Email: aguha@gg.iitkgp.ernet.in

     Ninety-nine bryozoan species (eight cyclostomatids under six genera and five families, and 91 cheilostomatids under 56 genera and 33 families) have been retrieved from rock formations belonging to the Lutetian - Burdigalian sequences of western Kachchh, Gujarat. Cyclostomatids are restricted to the lower part of Lutetian sequence (Harudi Formation). Only 24 percent of genera have more than one species under them. Among cheilostomatids, the ascophorans are more diverse at different hierarchical levels than the anascans. The present information on number of the Tertiary bryozoan species from Kachchh has considerably enlarged the old checklist of nine fossil species reported from this area about four decades ago. The diversity and richness of bryozoan colonies are highly variable within and between formations, reaching their maximum during the deposition of the Khari Nadi Formation (Aquitanian).
     Fifteen fossil species of Thalamoporella Hincks, 1887, recognized in these sequences, have doubled the number of fossil taxa under this genus from fifteen to thirty, and have enriched our knowledge about the Tertiary evolutionary history of this common extant genus of tropical and warm temperate waters. The presence of three steginoporellid genera in these formations, out of a total of five Tertiary genera under the family, makes Kachchh one of the main centers of radiation in the spatial evolution of this family. The species under Therenia David & Pouyet, 1978, till unknown in the Indo-Pacific realm, may take a conspicuous place in the phylogeny of the genus.
     The Tertiary basin of Kachchh, being a pericratonic rift basin located at the western margin of India, was generally shallow with shell banks of various magnitudes and argillites that supported bryozoan colonies associated with shells of foraminifera, bivalves (chiefly oysters), algae, gastropods and barnacles. Encrusters (both uni- and bilaminar) placed under 65 species and represented by over 61 percent of colonies examined, are far in excess of sum total of colonies under other six types of growth habits, namely, erect-rigid-foliaceous, erect-rigid-delicate, free-living, erect-foliaceous, erect-rigid-fenestrate and erect-rigid-robust. An analysis of paleoenviron-mental significance of relative abundance of growth habits in different formations has been made.

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Bryozoan fauna of Green Island, Taiwan - six new species for science

     Tea Gluhak
Rudjer Boskovic Institute, Center for Marine Research, G. Paliaga 5, 52210 Rovinj Croatia
     Peter J. Hayward
School of Biological Sciences, University of Wales, Swansea, Singleton Park, Swansea SA2 8PP
     Ivan Cvitkovic
Institute of Oceanography and Fisheries, Laboratory for benthos, Setaliste Ivana Mestrovica 63, 21000 Split, Croatia
     Jane E. Lewis
PO Box 7-18, Keelung, Taiwan
     Aleksandar Popijac
Laboratory for Animal Ecology, Department of Zoology, Faculty of Science, University of Zagreb, Rooseveltov trg 6, 10000 Zagreb, Croatia

     This poster reports on 6 species from Green Island, Taiwan. These are result of two workshops that processed and identified collections from a five day field trip to Green Island, southeast Taiwan. The annotated list presented here includes Amastigia tricervicornis, Caberea sinensis, Catenicella marceli, Hemismittoidea taiwanensis, Parasmittina spiculata, Celleporina avicularidentata which are reported for science for the first time. The primary objective of this study is to document the distribution of species, and to describe morphological features by which they have been proclaimed as new for science.

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Hierarchical sources of environmental variation in a modern bryozoan, Electra pilosa

     Steven J. Hageman
Department of Geology, Appalachian State University, Boone, North Carolina, 28608, USA
     Christopher D. Todd
Gatty Marine Laboratory, School of Environmental and Evolutionary Biology, University of St. Andrews, St. Andrews, Fife KY16 >8LB, Scotland, UK

     Documentation of genetic versus environmental sources of morphological variation in skeletal hard parts is important for meaningful application of species concepts to broader studies of paleobiology. This study evaluates the hierarchical influence of large and small scale environmental variation on skeletal morphology of the anascan bryozoan Electra pilosa from western Scotland. Previous studies by the authors have established the degree to which morphological variation is induced by genetic variation among closely relatedcolonies in controlled environmental conditions. The data set generated for the present study consists of colonies of E. pilosa collected from a 1.5 km region of Clachen Seil Sound, western Scotland at three Stations (~0.5 km apart), from three Substation within each station (10 m apart), five colonies per substation on separate fronds of brown algae (Laminaria), and two patches of multiple zooids sampled from each colony. A suite of five morphometric characters was collected from each and analyzed with Nested ANOVA and Principal Components Analysis. Comparable data sets were included in a second phase of analyses using colonies from varied environments along the western coast of Scotland, and colonies grown under invarient environmental condition in the laboratory. Preliminary results indicate that there is : (1) no systematic variation among the three primary stations at Clachen Seil Sound, (2) no systematic variation among the three Substations within each station, (3) variation among colonies within substations is significant for all characters, accounting for 20-35% of the total variance for most characters and 48.5% for the character Zooecia Length. (4) Zooecia Length is most significant among colonies, but less significant than all other characters within colonies. Thus, there is difference between genetic and microenvironmental (within vs. among colony) effects on length vs. width in zooecial characters, but little systematic effect on any at the meso-environmental level of 10 to 500 m of separation.

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Miocene free-living bryozoans from Patagonia

     Eckart Håkansson
Geological Institute, University of Copenhagen
     Silvio Casadío & Ana Parras
Departamento de Ciencias Naturales, Universidad Nacional de La Pampa

     Cupuladriids are reported for the first time from Patagonia, southernmost South America, where at least two species - Cupuladria sp. and Discoporella sp. - have been collected from upper Miocene strata.
     Miocene deposits in Patagonia, southern Argentina, constitute a comparatively thin sedimentary wedge deposited in an epicratonic basin related to the passive continental margin bordering the Atlantic Ocean. These deposits include a series of spectacular mollusk-bryozoan-dominated shell concentrations comprising the Puerto Madryn Formation. Well-developed exposures of this unit are located around Puerto Pirámides, Penísula de Valdés. Here the preserved section shows a vertical succession of facies passing from open mid-shelf to more shallow, shoreface to foreshore environments characterized by upward-coarsening cycles capped by condensed shell-beds. Whereas bryozoans in general are plentiful throughout the entire section, the free-living bryozoans appear to be restricted to a number of distinct horizons through the middle part of the formation, where they occur in fair amounts in fine sands with minor mud content.
     The biogeographic and phylogenetic implications of the new find will be discussed.

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Biogeography and phylogeny of the free-living bryozoans in the Miocene of South America

     Eckart Håkansson
Geological Institute, University of Copenhagen
     Silvio Casadío
Departamento de Ciencias Naturales, Universidad Nacional de La Pampa
     Sven Nielsen
Geologisch-Paläontologisches Institut und Museum, Universität Hamburg

     The first finds of cupuladriids from Patagonia, southernmost South America, have provoked speculations as to the mode and tempo of the Neogene invasion of free-living bryozoans into South America.
     Two routes were utilized from a Caribbean center of evolution prior to the closure of the Isthmus of Panama, one along the east coast and one along the west coast. Migration along the east coast reached as far south as Península de Valdés in Patagonia whereas, on the west coast, the southernmost occurrence is around Navidad in central Chile. While the Atlantic trail apparently was utilized solely by cupuladriids - with both Cupuladria sp. and Discoporella sp. reaching all the way to Patagonia - the Pacific trail may also have provided a sanctuary for the last surviving members of the North American branch of the Otionellidae.

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Bryozoans from Zechstein (Upper Permian) of southwestern Poland

     Urszula Hara
Polish Geological Institute Rakowiecka, 4 00-975 Warsaw, Poland

     The rich and diverse Permian bryozoan fauna from the southwestern part of Poland has proved to be significant in respect of taxonomy and biogeographical connections. The bryozoans come from the marly calcareous series of the Lower Zechstein of the North Sudetic Basin accompanied by a rich shallow water biota (Raczynski 1996). In terms of the higher taxa the Zechstein bryozoans are dominated respectively by fenestriids which form a lyre-shaped colonies, however, the trepostomids also constitue an important epibiontic element of the studied fauna. The following fourteen taxa have been distinguished in the systematic account, making extensive use of SEM: Stenopora columnaris (Lonsdale), Rectifenestella retiformis (Schlotheim), Fenestella geinitzi (d'Orbigny), Fenestella minuta Korn, Fenestella sp., Penniretepora sp., Thamniscus diffusus Korn, Thamniscus geometricus Korn, Thamniscus siccus Dreyer, Thamniscus sp., Kingopora solida (Korn), Acanthocladia anceps (Schlotheim), Acanthocladia minor Korn, Acanthocladia laxa Korn (see Hara 2001). The species of Fenestella sp., Penniretepora sp., Thamniscus siccus Dreyer, Thamniscus sp and Kingopora solida (Korn) are the first time recorded from Poland. Thamniscus sp. and Penniretepora sp. should to be referred to the type material, because they may represent the new taxa . The relationship between the colony-form, growth pattern, inferred associated biota as well as sedimentary structures points to shallow-water environment with low to moderate current activity for Zechstein palaeoenvironment of the North Sudetic Basin.
     This recently studied fauna adds to the knowledge of the Permian taxonomy some important facts which previously was based on the fragmentary data what reduces its biogeographical value. From the zoogeographical point of view, the Polish Permian bryozoans accentuates the biostratigraphical links and a marked faunal affinity with the bryozoans of the West-European province of England and Germany as well as with the southern Baltic Region. This fauna shows a great importance of fenestellids and acanthocladiids during the Zechstein giving priority of such taxa as Rectifenestella, Penniretepora, Thamniscus, Kingopora and Acanthocladia (Korn 1930, Gilmour 1999).

References
Hara, U. 2001. Bryozoa - The Lower Permian, In: Structural Geology of Poland, Atlas of the Index Fossils, Permian, Polish Geological Institute, (In Polish), 3: 43-49.
Raczynski, P. 1996. Palaeontological and sedymentological indicators for the development of the sedimentary facies in the Zechstein of the North Sudetic Basin. Archives of the Institute of the Geological Sciences. University of Wroclaw. Wroclaw.
Gilmour, E.H. and Morozova I.P. 1999. Biogeography of the Late Permian Bryozoans. Paleontological Journal, 33: 36-51.
Korn, H., 1930. Die Cryptostomen Bryozoen des Deutschen Perms, K. Deutsche Akad. Naturforscher zu Halle. Nova Acta Leopoldina, 6: 141-377.

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Morphological description of the setae of four species of the family Cupuladriidae from both sides of the Isthmus of Panama

     Amalia Herrera-Cubilla & Felix Rodriguez
Center for Tropical Paleoecology and Archaeology, Smithsonian Tropical Research Institute P.O. Box 2072, Balboa/Ancon Panama Rep. of Panama

     In this work we describe the morphology of the setae or mandibles of four species of the family Cupuladriidae. Three of them are from the Caribbean: Cupuladria biporosa Canu and Bassler 1923, Cupuladria surinamensis Cadée 1975, and Discoporella 2. While one is from the Pacific: Cupuladria 5 biporosa.
     C. biporosa and Cupuladria 5 biporosa have three kind of seta or mandibles. While C. surinamensis and Discoporella 2 presented only one type of seta. The morphology of the seta varies greatly between species and gives valuable information, which could help for a better taxonomical understanding of this family and probably its live history.

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Taxonomy of the genus Cupuladria from both sides of the Isthmus of Panama. I. Morphology and Systematics

     Amalia Herrera-Cubilla
Center for Tropical Paleoecology and Archaeology, Smithsonian Tropical Research Institute P.O. Box 2072, Balboa/Ancon Panama Rep. of Panama
     Matthew H. Dick
Department of Biology, Middlebury College Middlebury, VT 05753 USA
     JoAnn Sanner
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia 20650
     Jeremy B.C. Jackson
Center for Tropical Paleoecology and Archaeology, Smithsonian Tropical Research Institute P.O. Box 2072, Balboa/Ancon Panama Rep. of Panama Geoscience Research Division, Scripps Institution of Oceanography, La Jolla, California 92093

     Up to 28 morphological characters were used to analyze phenetically colony specimens of the genus Cupuladria collected from both coast of the Isthmus of Panama. These characters were used first to Cluster colonies and obtain discrete groups that were later entered in a series of Direct Discriminant Analyses that permitted us to identify morphospecies with high confidence. The taxonomy, so obtained, showed a clear cut off between specimens of the C. surinamensis clade vs. C. biporosa clade, as has been pointed out in the phylogeny of the genus Cupuladria. Also revealed a greater morphospecies diversity, from both sides of the Isthmus of Panama, than that documented in the literature, and demonstrated no evidence of the presence of C. canariensis in this part of Tropical America. Despite that more rigorous phenetical analyses still to do, this first attempt prove that morphometric analyses of cheilostomes should be pushed to their limit, to obtain meaningful taxonomies of particularly difficult groups like Cupuladria.

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Taxonomy of the genus Cupuladria from both sides of the Isthmus of Panama. II. Description of the species

     Amalia Herrera-Cubilla
Center for Tropical Paleoecology and Archaeology, Smithsonian Tropical Research Institute P.O. Box 2072, Balboa/Ancon Panama Rep. of Panama
     Matthew H. Dick
Department of Biology, Middlebury College Middlebury, VT 05753 USA
     JoAnn Sanner
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, District of Columbia 20650
     Jeremy B.C. Jackson
Center for Tropical Paleoecology and Archaeology, Smithsonian Tropical Research Institute P.O. Box 2072, Balboa/Ancon Panama Rep. of Panama Geoscience Research Division, Scripps Institution of Oceanography, La Jolla, California 92093

     The identification of eight species of the genus Cupuladria collected from both sides of the Isthmus of Panama was done using up to 28 morphological characters entered in a rigorous phenetic analyses described in a previous paper. Six species Cupuladria multesima, Cupuladria incognita, Cupuladria cheethami, Cupuladria pacifiensis, Cupuladria exfragminis and Cupuladria panemensis are described as new. Two species Cupuladria biporosa and Cupuladria surinamensis have a wider distribution in the Caribbean. This morphological description provides further documentation of the greater morphospecies diversity, of the genus Cupuladria from both sides of the Isthmus.

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A novel technique for the assessment of the distribution of Lophopus crystallinus, a rare phylactolaemate within the U.K

     Samantha Hill
School of Animal and Microbial Sciences, University of Reading, PO Box 228, Reading, RG6 6AJ, United Kingdom
     Beth Okamura
School of Animal and Microbial Sciences, University of Reading, PO Box 228, Reading, RG6 6AJ, United Kingdom

     At the Rio Earth Summit in 1992, the delegates ratified the Convention of Biological Diversity which committed the nations to the maintenance of biological diversity. This agreement led the U.K. to generate the Biodiversity Action Plan (B.A.P.) in 1994. The U.K. B.A.P. consists of Habitat Action Plans (H.A.P.) and Species Action Plans (S.A.P.). The H.A.P.'s and the S.A.P.'s respectively outline the current status and necessary management to ensure the continuation of threatened habitats and species within the U.K. Lophopus crystallinus is the only bryozoan listed within the U.K. B.A.P. and it is given a classification of 'RARE'. The objectives of the S.A.P. for L. crystallinus include the maintenance of all long-term populations in the U.K., and to increase the number of populations within the U.K. by 2010. The protection of sites with known populations is also discussed, as is the possibility of designating these sites as Sites of Special Scientific Interest (SSSI), which may help to safeguard the populations. The Invertebrate Site Register of L. crystallinus by English Nature lists 10 sites within England, but L. crystallinus has only been observed at four of these sites since the 1970's. O'Dea (unpublished report) surveyed various sites throughout the U.K. in 2001/2002 and found evidence of populations in only two sites. We have developed a technique to quickly ascertain the presence of L. crystallinus within a watershed by the inspection of flood or floating debris for L. crystallinus' floatoblasts. Flood debris gathers on banks after high waters and is normally composed of fine-grained organic detritus. Floating debris gathers in slow moving areas of lakes and rivers and can be composed of any floating material; often twigs, leaves, seeds, and grass are picked up along with statoblasts. We describe techniques to sample flood and floating debris for L. crystallinus statoblasts and present results of collections from over 50 sites. This approach has allowed us to confirm the presence of L. crystallinus in river systems from which it has previously not been recorded, and leads to the conclusion that its distribution is much more widespread than formerly thought. This work may have an impact upon the Rare status given to L. crystallinus within the U.K. Red Data Book and will certainly have an impact upon the S.A.P. of L. crystallinus.

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A review of the effects of heavy metal toxicity in freshwater Bryozoa

     Victoria B. Holmes
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom
     Mary E. Spencer Jones
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom

     Work on the effects of heavy metal contamination in fresh water Bryozoa has been very limited with the main bulk of investigations carried out in the 1980's. The studies carried out have concentrated on the mortality rates of the Bryozoa, with very little work carried out on the effects of the heavy metals on the internal structure of the organism or the statoblasts produced. Early work has mainly hinted at the possible effects of specific heavy metals on the structure of statoblasts with no continuation of the study. This review is an attempt to access the work carried out so far and to look at the areas that have not been covered in the experimental work carried out.

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Phylogeography and Sibling Speciation in Celleporella hyalina

     Roger N. Hughes, A. Gómez, P. J. Wright, D. Lunt, & G.R. Carvalho,
School of Biological Sciences, University of Wales, Bangor, Gwynedd, LL57 2UW, UK.
     Juan M. Cancino
Departamento de Ecología Costera, Facultad de Ciencias, Universidad Católica de la Santísima Concepción, Casilla 297, Concepción, Chile.
     Hugo I. Moyano G.
Departamento de Zoología , Universidad de Concepción; Casilla 160-C; Concepción.
hmoyano@udec.cl

     Celleporella hyalina is reported from cold-temperate to polar oceans circum-globally. We have sampled populations from around the world and investigated the genetic diversity, at both mitochondrial and nuclear loci, found in this 'cosmopolitan' species. Laboratory mating studies to determine mating compatability have accompanied this molecular approach.
     Both mtDNA and nuclear gene sequences strongly support C. hyalina as a sibling species complex. Mating compatibility trials indicate that some 11 major lineages are reproductively isolated. These lineages are not recent but represent millions of years of independent evolution, perhaps since the Miocene (~10myrs). Phylogeographic substructure is evident within each of these major lineages. The NE Atlantic contains several geographically structured lineages which are probably of Pleistocene or Pliocene origin.
     The C. hyalina species complex seems to have originated in the northern hemisphere, possibly in the N Atlantic, with at least two episodes of colonization of the Pacific predating the opening of the Bering Strait. At least three more recent long-distance colonization events were revealed. Two of these events are trans-tropical, involving colonization of the South Pacific by northern clades, one from the north Pacific to Chile and Argentina and other from the North Atlantic to the Magellanic region. A third event involved the recent colonization of Svalbard by genotypes from coastal Alaska.

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Discussions on the benefit of commensalism in solitary entoprocts (Entoprocta: Loxosomatidae)

     Tohru Iseto
The Kyoto University Museum, Kyoto University, Yoshida, Sakyo, Kyoto 606-8501, Japan.

     Entoprocts are suspension feeders that create water currents using tentacular cilia and catch food particles in the currents. Most of the solitary entoprocts (=loxosomatids) are known to live on bodies or tubes of other animals such as polychaetes, bryozoans, sponges, and sipunculans. The commensal species has been believed to be "energy commensals" that partly depend on host-made currents to get enough foods. In fact, some species have very short tentacles that seem to be incapable of creating sufficient water currents by themselves.
     Recently, ten non-commensal loxosomatids were described from Okinawa, Japan. The ten species live on non-living substrata and get foods through self-made water current only. However, there is no considerable difference in number and length of tentacles between commensal and non-commensal species. It suggests that most commensal species, except some short-tentacle species mentioned above, are also able to make sufficient water currents by themselves and not depend largely on the host-made currents.
     My observations on non-commensal species found in Okinawa, Japan suggest other benefits in their commensalism. Several times, I observed non-commensal species covered by bryozoans. Such coverage by other organisms is obviously fatal for entoprocts. The non-commensal species in Okinawa tend to settle on "naked" substrate that is less covered by other organisms: this may be to avoid covering by neighboring animals. On the other hand, some benthic animals such as nudibranches and flat worms are known to eat entoprocts. The tubes of host animals or protection mechanism of host animals may keep the predators away from loxosomatids.
     These observations and speculations suggest that the major benefit of commensalism in loxosomatids is avoiding coverage and predation by other animals. The host bodies or tubes seem to be very safety habitat for them.

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Bryozoans and Bryozoa-associated fungi from Galápagos Deep Sea

     Jürgen Kaselowsky & Joachim Scholz
Research Institute Senckenberg, Section ME III, Frankfurt am Main, Germany

     The major objective of the 1999 cruise of the R/V Sonne (SO 144-3) was the systematic sampling of volcanic rocks in the area between the Galápagos Islands and Central South America. In 2001, the R/V Sonne cruise 158 (SO 158) was the systematic sampling of rocks from submarine volcanoes ("seamounts") and an east-west trending volcanic ridge, the so-called Galápagos spreading center, located north of the Galápagos archipelago. Both cruises have documented a highly diverse bryozoan fauna, collected by P. Götz. B. Neuhaus and C. Lüter (Berlin). There are about 50 species, many of them new and belonging e.g. to Catenicella, Chaperiopsis, Escharina, Notoplites, Pleurocodonellina, and Talivittaticella. Furthermore, a strange interior walled cyclostome species was found that represents at least a new genus. In a colony Columnella cf. graminea collected from a depth of 1558 m, the frontal membrane is colonized by hyphae and yeast cells of fungi. This is one of the very first reports of fungi from the deep sea (see Kaselowsky et al 2003).

References:
Kaselowsky, J., Hamamoto, M., Nagahama, T., Scholz, J. & K. Sterflinger (2003): Marine Darwinfinken und Schwarze Pilze. - Biologie in unserer Zeit 33/1: 11. Weinheim.

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Comparison of selected ascophorine bryozoans from Red Sea, Philippines, and Socotra (Yemen)

     Jürgen Kaselowsky
Research Institute Senckenberg, Section ME III, Frankfurt am Main, Germany

     Ever since Harmer's Siboga Reports, type material from the Red Sea (e.g. Waters) has been used to identify bryozoans from the tropical western Pacific. On the other hand, in their monography of smittinids of Hawaii, Soule & Soule (1973) have demonstrated that bryozoans show a distinct regionalism (as marine "Darwin Finches"). A re-evaluation of types from some Indo-Pacific collections (e.g. Waters, Red Sea, Doederlein / Ortmann, Japan), and modern state-of-the-art re-illustration of type material has now become necessary to have a new data base about the comparative morphology of indopacific bryozoans. Some selected ones, which are known for extraordinary wid-ranging distribution patterns, should be revised. Comparison of type material with colonies from the Philippines, Japan and New Zealand, which have been collected within the scope of earlier research grants projects (HI 273/3, 446; SCHO 581/6; JAP-113/216/0; GE 64/8) helps to outline the true geographical range of several circumtropical or indopacific species. One holotype which was re-examined was Smittia tropica = Parasmittina tropica. The type-material from the Hartmeyer collection was described by Waters (1909). Parts of the holotype stored at the Humboldt-Museum of Natural History in Berlin have been bleached and prepared for SEM. Comparison of P. tropica specimen from New Zealand described by Gordon (1984), P. tropica from Mauritius reported by Hayward (1988), and colonies determined by Scholz (1991) from the Philippines revealed, that these are different species. In the collection of H. Ristedt (Bonn) which is now kept at the Senckenberg-Museum, additional new Parasmittina species similar to P. tropica are documented.

References:
Gordon, D. P. (1984): The marine fauna of New Zealand: Bryozoa: Gymnolaemata from the Kermadec Ridge. - N.Z. Oceanogr. Inst.Mem. 91: 1-198.
Hayward, P. (1988): Mauritian cheilostome Bryozoa. - J. Zool. 215: 269-356.
Scholz, J. (1991): Die Bryozoenfauna der philippinischen Riffregion Cebu. - Mitt.Geol.-Paläont.Inst.Univ.Hamburg 71: 253-403.
Soule, D. & Soule, J.D. (1973): Morphology and Speciation of Hawaian and eastern Pacific Smittinidae (Bryozoa, Ectoprocta). - Bulletin of the American Museum of Natural History 152 (6): 365-440.
Waters (1909): Reports on the marine bioloy of the Sudanese Red Sea, Part 1 - Cheilostomata. Journal of the Linnean Society. Zoology 31: 123-181.

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C and O isotopic test of the algal symbiosis hypothesis for gigantism in Permian Trepostomes from Greenland

     Marcus M. Key, Jr.
Department of Geology, Dickinson College, Carlisle, PA 17013-2896, U.S.A.
     Patrick N. Wyse Jackson
Department of Geology, Trinity College, Dublin 2, Ireland
     Eckart Håkansson
Institute of Geology, Øster Voldgade 10, DK-1350 København, Denmark
     William P. Patterson
Department of Geological Sciences, University of Saskatchewan, 114 Science Place, Saskatoon, SK S7N 5E2, Canada
     M. Dustin Moore
Department of Geology, Dickinson College, Carlisle, PA 17013-2896, USA

     Gigantism in organisms can be caused in various ways: by disease, under ideal growing conditions, or symbiosis. It is well documented that symbiotic zooxanthellae algae can cause gigantism in Recent hermatypic corals. However, such symbiotic relationships have never been documented for extant or fossil bryozoans. Unusually large colonies of the trepostome bryozoan Tabulipora sp. have been recovered from the Kungurian (Early Permian) Kim Fjelde Fm. in eastern North Greenland. Branches reach 7 cm in diameter in this species, while other bryozoans in general and other Tabulipora species in particular in the fauna are an order of magnitude smaller. Disease can be ruled out as a causal factor for this example of gigantism, as all individuals in the species exhibited gigantism. Ideal growing conditions can not be completely ruled out as not all other species in the fauna exhibited normal growth sizes. Håkansson and Madsen (1991) hypothesized that the gigantism was caused by algal symbiosis. Their conclusion was based on carbon and oxygen isotopic values derived from coarse sampling that required ≥1.5 mg carbonate.
     In this study using a colony from the same formation and location, a more precise test of their hypothesis was conducted by sampling with 10 µm spatial precision using a computer-driven micromilling device that required only ≥ 20 µg carbonate. Skeletal results indicate a mean δ¹³C value of 3.85 ‰VPDB and a mean δ¹⁸O value of -6.52 ‰VPDB. Diagenetic effects were evaluated by separately sampling the cements contained within zooecial chambers; skeletal values are significantly heavier than the surrounding cements. Taking into account the inferred isotopic composition of the Permian ocean, it is concluded that these isotopic results fail to reject the algal symbiosis hypothesis for gigantism in these bryozoans. Other potential supporting information is lacking or ambiguous. Other than size and the isotopic signature, the remaining morphologic and paleoenvironmental evidence is equivocal. For example, marine organisms with symbiotic algae typically are restricted to simple phyla, have thin tissue layers, filter feeder, grow toward light, have high surface area-to-volume ratios, high skeleton-to-body ratios, and high levels of colonial integration. They are most common in tropical, oligotrophic, shallow water, low turbidity, reef environments. The giant Tabulipora colonies from the Permian of Greenland do not meet most of these criteria.

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From Plate tectonics to bryozoan evolution

     Marcus M. Key, Jr.
Department of Geology, Dickinson College, Carlisle, PA 17013-2896, U.S.A.
     Abigail M. Smith
Department of Marine Science, University of Otago, P. O. Box 56, Dunedin, New Zealand

     The hypotheses of biologists, paleobiologists, ecologists and paleoecologists often focus on biotic interactions as they can play important roles in evolution. Often less obvious are abiotic evolutionary forcing mechanisms, particularly large-scale global geochemical cycles. Stanley and Hardie (1998) synthesized a theory of secular geochemical variation over the Phanerozoic, describing oscillations in the mineralogy of marine inorganic and biogenic carbonates, with periods dominated by aragonite/high-Mg calcite (aragonite seas) and low-Mg calcite (calcite seas). They argued that temporal variations in the Mg/Ca ratio of sea water were driven by changes in plate tectonic spreading rates at mid-ocean ridges. Their supporting data included marine cements, ooids, evaporites, and tropical hypercalcifying organisms (e.g., calcareous algae, forams, corals).
     If seawater chemistry changes are global in scope, then geochemical signals should be expressed beyond these dominantly tropical examples. Temperate bryozoans, with their wide mineralogical and stratigraphic ranges, offer the opportunity to ascertain the extent of the mineralogical influence of sea water on biogenic carbonate production.
     Here we report on the results of a literature review of bryozoan mineralogy over the Phanerozoic. Do bryozoans exhibit more aragonite/high-Mg calcite skeletons during times of aragonite seas and more low-Mg calcite skeletons during times of calcite seas? Do bryozoans have the potential to delineate and refine this global model? And has bryozoan evolution itself been constrained by sea water chemistry?

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Bryozoans from the Jade Bight in Northwestern Germany and molecular biological analysis of their associated bacteria

     Sandra Kittelmann
Institute for Chemistry and Biology of the Marine Environment (ICBM), Environmental Biochemistry, Carl-von-Ossietzky University of Oldenburg, P.O. Box 2503, 26111 Oldenburg, Germany

     The Jade is an extended tidal polyhaline basin on the northwestern coast of Germany. The river Weser provides continuous freshwater supply. At high tide, the Jade covers an area of approximately 160 square kilometres.
     Preliminary screens of the site have shown the dominant occurence of the anascan species Conopeum reticulum, Conopeum seurati, Electra pilosa and Electra monostachys. These species predominantly occur on native mussel shells and will be collected by dredging.
     Based on previous research it is known that both the surface and the coelomic cavity of bryozoa may provide an ideal habitat for specific phylogenetic groups of bacteria depending on metabolism and texture of the host organism (Davidson et al. 2001, Gerdes et al., in prep). Host-specific bacterial community profiles are quite common in the marine environment, however they have been sparsely investigated on bryozoans so far.
     The main focus of the present study is the analysis of community profiles of bacterial associates on and within bryozoans.
     The surface topography and/or the chemical influence of bryozoan metabolites with nutritious or allelochemical effects may significantly shape bacterial communities on the host surface.
     In order to test these hypotheses, different locations for bacterial attachment and proliferation will be investigated, i.e. in and on the bryozoan, each with a "negative control":
     1. Coelomic cavity of living bryozoans
          vs. interior of immersed bryozoan skeletons
     2. Surface of living bryozoans
          vs. smooth substrate surfaces proximate to the bryozoan colonies.
     The bacterial community profile will be examined by means of molecular biological methods such as DNA-extraction, PCR, denaturing gradient gel electrophoresis and concluding sequence analysis. Furthermore a screening for antibacterial substances is carried out with extracted bryozoan material in order to explain the putative differences in community profiles. Therefore marine bacteria from the Jade bight will be cultivated, isolated and tested for antibacterial effects in contact with the examined bryozoan species (stand disc susceptibility assay).
     The results of the antibacterial screening in addition to the comparison between the viable and the non-viable conditions will allow conclusions for the reasons of potentially different bacterial community profiles on bryozoans.

References:
Davidson, S.K., S.W. Allen, G.E. Lim, C. Anderson and M.G. Haygood. 2001: Evidence for the biosynthesis of bryostatins by the bacterial symbiont "Candidatus Endobugula sertula" of the bryozoan Bugula neritina. Appl. Env. Microbiol. 67:4531-4537.
Gerdes, G., Kadagies, N., Kaselowsky, J., Lauer, A. & J. Scholz (in prep.): Bryozoans and microbial communities of cool-temperate and subtropical latitudes (Japan, New Zealand) - Paleoecological implications II. Diversity of microbial fouling on laminar shallow marine bryozoans.

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Biodiversity on coastal boulders at Spitsbergen

     Piotr Kuklinski
Institute of Oceanology, Polish Academy of Science, ul. Powstanców Warszawy 55, Sopot 81-712, Poland (mailing address)
University Center on Svalbard, P.O. Box 156, N-9171 Longyearbyean, Norway
     David. K. A. Barnes
British Antarctic Survey, High Cross, Madingley Road, Cambridge, CB3 0ET, United Kingdom

     Boulder communities were studied in summer 2002 in West Spitsbergen fiords. We examined multiple samples at three stations in Kongsfjorden (79° N, 12° E) and three stations in Hornsund (77° N, 16° E) fjord. Horsund is more influenced by Arctic water masses while Kongsfjorden is by Atlantic warmer waters. Altogether 2752 boulders (of 254 681 cm² surface area) were investigated in the intertidal, 6 m and 12 m. 73 taxa of Bryozoa were determined; 1 to phylum, 1 to order, 3 to family, 16 to genus and 52 to species level. ANOSIM a priori statistic (Global R=0.141, p=0.016) reveal no difference between the investigated sites.
     Cluster and multidimensional scaling analyses divided samples into two groups: intertidal samples and the rest of samples. Thirteen taxa (8 species) occurred in the intertidal zone whilst 73 taxa (52 species) were subtidal. All the intertidal taxa were also present in the subtidal samples. Communities were dominated by Harmeria scutulata (57% in of colonies in the intertidal and 54% in the subtidal). The next most abundant species, Cauloramphus intermedius and Cribrilina annulata represented just 5% of colonies each; the dominance values for the other taxa were below 5%. There was no diversity pattern along the depth gradient. In both fjords the highest values of Shannon-Wiener index were 2.7. Despite the distance between Kongsfjorden and Hornsund (over 200 km) and differences in hydrological features between the fjords no difference between their boulder communities.

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Bryozoan mode of life in the high Arctic dynamic fjordic environment

     Piotr Kuklinski
Institute of Oceanology, Polish Academy of Sciences, ul.Powstanców Warszawy 55, Sopot 81-712, Poland (mailing address)
University Center on Svalbard, P.O.Box 156, N-9171 Longyearbyean, Norway

     The aim of the study was to investigate the adaptations of bryozoans to life in dynamic environment of inner parts of arctic fjords. The study was carried on in West Spitsbergen fiords. The considered areas are characterized by high rate of sedimentation (up to 500g/m/24hr), high concentrations of total suspended matter (500g/dm³???) in water, fine unstable bottom sediments, fresh water discharge from glaciers and rivers, cold stagnant waters at the bottom. Majority of the coastline in the inner part of the considered fjords is occupied by tidal glacier. This investigation is based on material collected during the r/v Oceania and r/v Jan Mayen expeditions to Svalbard in years 1997-2002. Research was carried out in four fjords: Hornsund, Van Mijen, Isfjorden and Kongsfjorden. Four main types of adaptations to the dynamic fjordic environment were observed: free-living, colonisation of drop-stones, colonisation of algae, creation of roots. Free living bryozoans are only represented by one species - Alcyonidium disciforme Smitt. This species by its shape (disc like) and encapsulation of sand particles within the zooarium (acting as ballast) is very much adapted to live on unstable fine sediments and to thrive high rate of sedimentation. "Faunistic islands" created by drop stones are acting as oasis of rocky fauna on unfriendly for suspension feeders (bryozoans) soft sediment. Eucratea loricata Linnaeus and Dendrobeania murrayana Johnston are dominants here. Very often algae growing on the drop-stones are the substrate for bryzoans. During this study three species of algae were investigated: Desmarestia aculeata, Laminaria sacharina and Ptilota plumose. Harmeria scutulata Busk and Celleporella hyalina Linnaeus are the most numerous species present on that kind of substrate. Root creating bryozoan was recorded very rarely. Kinetoskias arborescens Danielssen was the only species observed.

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Boreholes, small drilling predators, and reparative growth in fossil cheilostome bryozoans

     Scott Lidgard
Department of Geology, Field Museum of Natural History, 1400 S. Lake Shore Dr., Chicago, Illinois 60605, USA

     One major evolutionary trend in cheilostome bryozoans is the broad secular increase of calcified armament of the frontal wall of the zooid, in all likelihood to protect the soft tissues inside from predators or mechanical stress. Reports of modern predators on bryozoans are dominated by omnivorous fishes and macroinvertebrate grazers, and by nudibranchs at the shallow extreme of bryozoan depth ranges. A survey of reported bryozoan predators reveals zooid-scale boring and demineralizing predators including turbellarians, nematodes, errant polychaetes, and both juvenile and adult prosobranch and opisthobranch gastropods. However, most taxonomic groups of small drilling predators on bryozoans are grossly underrepresented by collecting and observation biases, and thus their relative predation frequency is unknown. Some of these predators' feeding behaviors also circumvent skeletal armament by attacking fleshy everted polypides or boring through uncalcified zooid operculae without leaving skeletal boreholes. Skeletal/non-skeletal weight ratios and mechanical resistance to puncture suggest that for some drilling or demineralizing predators, dining on thickly calcified zooids may be overpriced energetically relative to consuming uncalcified zooids. Species with uncalcified frontal walls make up a disproportionately large percentage of all cheilostomes reported in these predators' diets. Fossil drilling predation is evidenced by small, centrally located and uniformly positioned boreholes. Yet this fossil evidence is actually quite sparse. Fossil cheilostomes from the Paleocene Vincentown Formation of New Jersey provide a rare example that meets both necessary and sufficient criteria to distinguish predatory borings from postmortem borings or other sources of holes in exterior skeletal walls. Fossil skeletal evidence of frequent reparative zooid budding in Lower Cretaceous taxa with uncalcified frontal walls supports the inference that zooid-scale predators were present from the earliest stages of cheilostome diversification.

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Cheilostomate Bryozoa from the Bellingshausen Sea (Western Antarctica): results of the BENTART 2003 Spanish expedition

     Carlos M^a López-Fé
Laboratorio de Biología Marina. Departamento de Fisiología y Zoología. Facultad de Biología. Universidad de Sevilla. Avda. Reina Mercedes, 6. 4102 Sevilla. Spain.

     The Spanish Antarctic expedition BENTART 2003 was carried out in January and February 2003 by the oceanographic ship "B.I.O. Hesperides", and collected benthic material from an area comprised between the Antarctic Peninsula and Thurston Island. Most stations yielded bryozoans, usually on pebbles and rocks from a bottom mainly constituted by iceberg debris. At the moment of writing this abstract not all the material has been identified. Comparisons will be done between stations and with data from other expeditions and previously published works, in order to find affinities and differences with other areas of Antarctica.

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Colonial fusion in relation to coancestry in Celleporella hyalina

     Patricio H. Manríquez
Estación Costera de Investigaciones Marinas, Las Cruces & Center for Advanced Studies in Ecology & Biodiversity. Departamento d Ecología, Pontificia Universidad Católica de Chile, Casilla 114-D, Correo 22, Santiago, Chile.
     Roger N. Hughes
School of Biological Sciences, University of Wales, Bangor, Gwynedd, LL57 2UW, UK.
     John D. Bishop
Marine Biological Association of the UK, The Laboratory, Citadel Hill, Plymouth PL1 2PB, UK and Department of Biological Sciences, University of Plymouth, Drake Circus, Plymouth PL4 8AA, UK

     Manipulated contact among fragments of adult colonies and early post-metamorphic colonies of Celleporella hyalina were used to investigate colonial fusion in relation to coancestry. Fusion trials demonstrated colonial fusion can occur after two colonies of C. hyalina make contact with each other at their natural growing edge. Experiments using colonies from the United Kingdom (Welsh populations) showed that coalescent zooids were strongly associated with inter-colonial fusion of manipulated contact among fragments of the same colony (isocontact). In the allocontact trials, fusion was mainly recorded in full-sib:full-sib and parent:F1 offspring contacts. Similar results were found in contact among early post-metamorphic colonies produced by full-sib and half-sib larvae. No colonial fusion was recorded in contact among unrelated fragments of colonies or among early post-metamorphic colonies originated by unrelated larvae. Moreover, some fusion between half-sibs trials were recorded.
     Experiments with Chilean specimens were conducted with colonies collected in two geographically distinct populations located ~1000 km apart (Antofagasta and Las Cruces). Within each geographic population, colonial fusion was recorded among early post-metamorphic colonies originated by larvae released from the same colony. In contrast, within each population no colonial fusion was recorded among pairs of postmetamorphic originated by two different colonies. These results, therefore suggest a strong association between colonial relatedness and fusion compatibility. Moreover, the total absence of both fusion among colonies from the two Chilean geographic populations and offspring of manipulated cross mating are in agreement with ongoing genetic studies by one of the authors (RNH). This suggests that each geographic population may represent a different species in the genus Celleporella.
     Funding: Natural Environment Research Council Grant GR3/1026 to RNH and JDDB. Chilean part of this study was funded by an A. Mellon Foundation project to J C Castilla.

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Measuring and analyzing morphological complexity in cheilostome bryozoans

     Virginia A. Miller
Committee on Evolutionary Biology, University of Chicago and
Department of Geology, Field Museum of Natural History, Chicago, IL, USA

     The dramatic macroevolutionary trend of increasing complexity has provoked much discussion and speculation as to its cause(s), yet it has rarely been studied in a rigorous fashion. Many studies lack an explicit definition of complexity and/or fail to test observed trends for evidence of a driving force necessitating an evolutionary explanation. A primary obstacle to studying complexity is that it is difficult to define and quantify within organisms. The number of parts or part types is commonly used as a quantitative measure of complexity, but parts can be difficult to define. At present, the question remains: how can biological complexity be meaningfully quantified? Here I present a method for measuring complexity and analyzing observed trends within the cheilostome bryozoans.
     Cheilostome bryozoans provide an excellent opportunity to investigate the evolution of complexity because their modular organization allows objective definition of parts. The modular units -zooids - of a bryozoan colony are developmentally equivalent and can objectively be considered equivalent parts of the colony. Following this logic, each morphologically distinct - polymorphic - zooid type is a distinct part type. The complexity of a cheilostome colony can therefore be quantified as a function of the number of polymorph types. Quantifying morphological complexity in this way provides a measure of whole-organism complexity, an improvement over previous complexity studies that measured complexity of a single component (e.g. the mammalian vertebral column) as a proxy for whole-organism complexity. The high degree of polymorphism within the cheilostomes and their extensive, well-sampled fossil record will permit comparisons of complexity within populations, within lineages, and among lineages through geologic time.
     The maximum degree of polymorphism (complexity) in cheilostomes increases from their origin in the latest Jurassic to the present. It is not known, however, whether the increase is passive (a random walk away from a lower bound) or driven (the result of an evolutionary force). This distinction is important because a driven trend implies an evolutionary mechanism (such as selection favoring an increase in complexity), whereas a passive trend requires no such explanation. To distinguish between these two types of trends within the cheilostomes, I will analyze the distribution of complexity values both for cheilostomes as a whole and within subgroups. A positive skew of the data provides evidence for an active evolutionary mechanism; a skew of zero indicates a trend not significantly different from a random walk.
     A preliminary study of the cheilostome bryozoans included in the Synopses of the British Fauna will be presented. The complexity of each genus will be measured by counting the number of described polymorphs. The complexity of these Recent cheilostomes will then be characterized by plotting the complexity values, both for the study group as a whole and within subgroups, and measuring skew. This study will determine whether the proposed measure of biological complexity will prove a fruitful avenue for studying complexity in cheilostome bryozoans.

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Bryozoans in Messinian carbonate buildups of western Sardinia, Italy

     Pierre Moissette
UFR Sciences de la Terre, UMR 5125 Paléoenvironnements & Paléobiosphère, Université de Lyon I, 27 Bd du 11 Novembre, 69622 Villeurbanne cedex, France
     Jean-Paul Saint Martin & Frédéric Garcia
Centre de Sédimentologie-Paléontologie, UMR 6019, Université de Provence, 3 place Victor-Hugo, 13331 Marseille cedex 03, France
     Jean-Pierre André
Laboratoire de Géologie, Université d'Angers, Bd Lavoisier, 49045 Angers cedex, France
     Jean-Jacques Cornée
UFR Sciences de la Terre, UMR 5125 Paléoenvironnements & Paléobiosphère, Université de Lyon I, 27 Bd du 11 Novembre, 69622 Villeurbanne cedex, France

     The Messinian (Late Miocene) Capo San Marco Formation of the Sinis Peninsula (Western Sardinia) contains numerous buildups that developed within a siliciclastic matrix (silty to sandy marls).
     The biogenic buildups are made of abundant microbial carbonates constantly associated with serpulid worms and bryozoans. Other sessile and vagile benthic organisms are also represented: coralline algae, foraminifers, calcareous sponges, solitary corals, bivalves, gastropods (notably vermetids), brachiopods, crustaceans (ostracodes and decapods), and echinoids.
     The microbialites constitute small bulbous and irregular masses, more or less anastomosed, or small pillars having a distinctive cauliflower-like shape. Serpulid tubes are often incorporated in the microbialite, but also occur within the sediment infilling. Bryozoans are always associated to the constructional processes. Erect rigid colonies, especially reteporiforms, but also small celleporiforms, first perform the important role of baffling and trapping mud-sized sediment. A complementary role is often played by numerous membraniporiform colonies coating the walls of cavities within the microbialite framework.
     A total of 28 species of bryozoans have been identified in the study buildups. The two species having a major frame-building role, Sertella septentrionalis (reteporiform) and Celleporina costazi (celleporiform), occur in all bioherms where they are always abundant to very abundant. Six vinculariiform, one adeoniform and another celleporiform species are represented; they are less ubiquitous, but some of them also contribute to the construction. Although diversified (13 species) the membraniporiforms are much less abundant and conspicuous. Numerous segments belonging to five cellariiform species are found within the infilling sediment.
     The major sedimentary and biogenic features of the serpulid-bryozoan-microbial buildups of western Sardinia are characteristic of a lower shoreface environment with intermittently agitated water, probably at depths of not more than 10-20 metres.
     No modern analogues for these Messinian carbonate buildups exist, but Upper Palaeozoic mud-mounds and bioherms show strong biotic and structural similarities. In both cases, the intimate association of microbial communities with a variety of filter-feeding invertebrates, especially reteporiform bryozoans, has been recorded. Another common feature with the Messinian carbonate platform of Sardinia is the notable absence or the scarcity of hermatypic corals and calcareous algae.
     Upwelling currents are suggested as contributing to the spectacular development of microbial carbonates, the abundance of filter-feeding metazoans and the dearth of corals. Not only cold deeper water are brought to the surface, but the increased nutrient levels are accompanied by the proliferation of microbial communities and numerous sessile invertebrates.

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Microbial activity and frontal wall pore sieve plates in northeastern Pacific Microporellidae

     Penny A. Morris
University of Houston Downtown, 1 Main St., Houston, Tx. 77002, USA
     Dorothy F. Soule
Hancock Institute for Marine Studies, University of Southern California, Los Angeles, California 90089-0371, USA.

     Pore plates among eastern Pacific species of the Microporellidae genera Microporella and Microporelloides appear to be restricted to members inhabiting the southern and central California coastlines. Frontal wall pores are circular, but vary in size, and pattern, and placement depth within the frontal wall pores. The apparent occurrence of pore sieve plates within a restricted geographical locality is intriguing, but more intriguing is the presence of recently "fossilized" biofilms, filaments and ooid shaped calcium carbonate particles that appear to be concentrated either on or in close proximity to the pore plates. The existence of microorganisms associated with bryozoans have been described by other authors from both modern and fossilized localities (e.g., Scholz 1995, Scholz et al. 1995; Morris et al. 2002).
     The present study is different in that it is directed towards specific morphological structures of the bryozoan colony. Analyzed samples, some of which were pre-treated with sodium hypochlorite, are compared both morphologically, using a scanning electron microscope (SEM), and elementally with an elemental energy dispersive x-ray system (EDS). Preliminary EDS analysis indicates high levels of calcium, as would be expected, and low levels of magnesium, sulfur, and usually phosphorous. Morphological analysis of all bryozoan samples indicates the apparent absence or low numbers of microbes that occupy the frontal walls. The pore plates appear to possess higher levels of microbial activity as evidenced by the variety of their remains. For instance, Microporelloides setiformis frontal wall pore sieve plates contain filaments, mineralized biofilms and coccoid forms. Pore plates from another species, Microporella californica, possess a flotsam of probable microbial debris composed of a variety of shapes and sizes. The interior surface of a Fenestrulina farnsworthi ascopore has biofilms and diatoms. In the same specimen, the interior surface of a frontal pore is plugged with punctuate, egg-shaped calcium carbonate particles with evidence of small coccoid particles and biofilms. The question asked is why is the existence of microbes and pore plates important? First of all, we do not know the significance of pore sieve plates, which occur in an apparently restricted geographical area. Are they a protection against a specific type of predator? Pore plates would reduce the size of the pore and possibly restrict entry from predators above a specific size.
     Do microbes and their biofilms aid in the defense of a potential breach in the frontal wall membrane? Are the biofilms and their inhabitants active in preventing fouling of the surface and possibly part of an anti-predator brigade? Is there an evolutionary significance to these characteristics?

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Bryozoa of the CIMAR-7 Expedition to the Aysenian fjords and channels

     Hugo I. Moyano G.
Departamento de Zoología , Universidad de Concepción; Casilla 160-C; Concepción.
hmoyano@udec.cl

     Although studies on the bryozoans of the southern cone of South America have lasted more than 150 years and that have resulted in more than 200 species known so far, little is actually known of the bryozoans inhabiting the interior seas of Chiloé, Aysén and Magallanes. This lacking have been lessened during the last two decades by the collecting activities of Chilean, German and Italian cruices and expeditions to the Magellan straight areas and to channels and fjords situated north and south of it. Their faunal results allow to extend the geographical distributions of many species initially described from the Falkland and Burwood Bank areas up to Cape Horn and Magellan straight, what means a similarity of bryozoan marine faunas on both sides of the southernmost part of South America.
     The Aysenian region intermediates between the Magellanian and Chiloean ones to south and north respectively and some zoogeographers have assumed to have also an intermediary zoogeographical role among a southernmost Magellanic province and a northenmost Peruvian-Chilean zoogeographical region reaching Chiloean fjords and channels. In order to assess this and other issues resulting from the isolation and lack of knowledge of the Aysenian area it was carried out the CIMAR-7 Expedition.
     Bryozoans collected between 20 and 186 m depth in channels and fjords north to the Penas gulf , v . gr. Estero Elefantes near Laguna San Rafael National Park yielded typically Magellan species. Among these stand out Chondriovelum angustilobum, Beania maxilla, B. fragilis, Cellaria malvinensis, Callopora deseadensis, Aspidostoma giganteum, Nevianipora milneana, Smittina undulimargo and Romancheina labiosa. The presence of the genera Chondriovelum and Adeonella reveals both vicariant relationships with Antarctica , South Africa and Australia and also a bryozoan similarity among external and internal fjors, islands and channels of the Magellan Madre de Dios area and the interior sea of the Aysenian region.
     As a general conclusion the bryozoans collected by the CIMAR-7 Expedition indicate the existence of a typical Magellanic fauna in the inner Aysenian sea.

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Some Upper Permian bryozoans from Svalbard, Arctic Norway

     Hans Arne Nakrem
Geological Museum, University of Oslo, PO Box 1172 Blindern, NO-0318 Oslo, Norway

     The Upper Permian (Ufimian-?Kazanian) succession of Svalbard comprises the upper part of the Kapp Starostin Formation (above the Vøringen Member) (a c. 350 m thick unit) in western and central Spitsbergen, and the Miseryfjellet Formation (c. 110 m thickness) of Bjørnøya, - both units within the Tempelfjorden Group (?latest Artinskian-?Kazanian). The Ufimian-?Kazanian part of the Kapp Starostin Formation consists of silicified spiculitic shales ("cherts"), partly silicified limestones and locally developed glauconitic sandstones in the uppermost part. The Miseryfjellet Formation is made up of limestones (partly silicified) and more sandy units. A Permian hiatus separates the Tempelfjorden Group from the overlying soft Triassic shales.
     The fossils in the investigated sections are typical Late Permian cold water forms, - sponges (spicules), brachiopods, echinoderms and bryozoans. Corals and trilobites are rare, and other temperate to warm-water forms, such as fusulinid foraminiferans are absent. The age of the Upper Permian units of Svalbard is uncertain, and brachiopods, conodonts and palynomorphs have yet not given conclusive ages.
     The shaley parts of the Kapp Starostin Formation is dominated by abundant Ramipora hochstetteri Toula, finely branched trepostomes (Rhombotrypella, Stellahexaformis and Dyscritella), cryptostomes (Permoheloclema merum Ozhgibesov, Primorella polita Romanchuk & Kiseleva and Clausotrypa spinosa Fritz) and minute fenestellids like Rectifenestella pseudoretiformis (Morozova), Fabifenestella completa Morozova & Kruchinina, Alternifenestella greenharbourensis (Nikiforova), A. spitzbergenensis (Nikiforova), Lyrocladia vera Morozova & Kruchinina, and Polypora kossjensis Ravikovich. Timanodictyids are also present in this association, with Timanodictya nikiforovae Morozova and Gilmoropora heintzi (Malecki). The partly silicified limestone units are dominated by thick trepostomes (species of Tabulipora, Stenopora timanensis Morozova and Dyscritella parallela Morozova), robust species of Polypora, Acanthocladia and Reteporidra. This division may reflect the recurrent changing depositional environments (quiet/rough waters), or a selection through transport.
     The identified faunas resemble closely Ufimian faunas described from Ellesmere Island (Assistance Formation) and Novaya Zemlya (Gerke and Savina Groups).
     The Miseryfjellet Formation (?Kungurian-?Kazanian) of Bjørnøya shows a great variability and richness regarding both bryozoans and brachiopods. 30 bryozoan species have been identified. Typical taxa in the lower part of the Miseryfjellet Formation include Rhombotrypella alfredensis Morozova & Kruchinina, Tabulipora greenlandensis Ross & Ross, Dyscritella bogatensis Morozova, Rectifenestella retiformis (Schlotheim), Septopora synocladiaformis Nikiforova, Polyporella optima Morozova & Kruchinina, Kingopora micropora (Stuckenberg), Timanodictya nikiforovae Morozova, Gilmoropora heintzi (Malecki). Some new species appear in the upper part: Ramipora hochstetteri Toula, Stenopora grandis Morozova, Permoheloclema merum Ozhgibesov, Rectifenestella cf. gijigensis (Nekhoroshev), Lyrocladia cf. vera Morozova & Kruchinina and Polypora kossjensis Ravikovich.
     The investigated fauna shows a mixture regarding age assignment, as there are both Early and Late Permian species throughout this formation. There are species in common with the Lower Permian of Eastern North Greenland, the Urals and Timan, as well as with the Upper Permian of England (Zechstein) and the Urals. The lower part of this formation has species in common with the Vøringen Member (Spitsbergen) while some species in the upper part are common with the upper part of the Kapp Starostin Formation.

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Fossil Bryozoa from Svalbard (Arctic Norway) - a research history

     Hans Arne Nakrem
Geological Museum, University of Oslo, PO Box 1172 Blindern, NO-0318 Oslo, Norway

     The Svalbard Archipelago - 74°-81° N and 10°-35° E - was rediscovered by Willem Barents in 1596, and scientists began visiting the island group in the early 18. century. The first geological expedition was that by B. M. Keilhau in 1827. Keilhau visited Bjørnøya and Spitsbergen, and collected amongst other material Upper Permian bryozoans. Some of his brachiopods were described by L. von Buch in 1847 (for example Spirifer keilhavii), and he mentions Fenestella antiqua in the collection.
     British expeditions to Svalbard brought new fossil material, including some Upper Permian bryozoans from Spitsbergen (Stenopora and Fenestella) collected by J. Lamont in 1859, mentioned by J. W. Salter in 1860. He discusses in detail a possible "new genus" which may very well be identical to Ramipora hochstetteri later described by F. Toula.
     During an Austrian expedition in 1873 R. von Drasche brought back a collection of Upper Permian bryozoans from central Spitsbergen and Akseløya, subsequently described by F. Toula in 1875, and a new genus was erected, - Ramipora, with R. hochstetteri as type species. Toula's description was supported by illustrations, and the fauna also includes other new species: Polypora grandis and Phyllopora laubei.
     Material from Spitsbergen was collected during British expeditions in 1906 and 1907 by W. S. Bruce, and subsequently described by G. W. Lee in 1908. Two new bryozoan species were described: Stenopora cidariformis and S. brucei both from the Kapp Starostin Formation (Kungurian-Ufimian age). O. Holtedahl published in 1911 and 1913 the first Early Permian bryozoans (no new species), with strong affinity with Early Permian faunas from Western Siberia.
     Russian expeditions to Svalbard have taken place since the days of whaling and hunting, and during a Russian expedition to Bjørnøya in 1921 Upper Permian bryozoans were collected. P. I. Stepanov reported 11 bryozoan species (no new) from the "Spirifer Limestone". The first illustrated work based on internal characters in thin sections was published by A. I. Nikiforova in 1936. Four new species and one new subspecies described from Kongressdalen (Kapp Starostin Formation): Fenestella greenharbourensis, F. spitzbergenensis, Polypora reteporidraeformis, P. timorensis var. greenharbourensis, and Septopora synocladiaformis.
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