Behavioral stereotypy and some ecological consequences of entrance-shaft placement of the domichnium Sanctum laurentiensis in Ordovician trepostomate Bryozoa

Abstract

The ichnofossil Sanctum laurentiensis from the Late Ordovician (Cincinnatian) of Laurentia is a domichnium tunneled into live colonies of ramose and frondose trepostome Bryozoa. A preferred entrance location was chosen by the trace-making endoskeletozoan. The organism chose to tunnel in positions near growing tips of branches where one of two conditions prevailed. At stereotypical sites, here considered “determinate”, domicile entrances were made where a trepostome branch was in the growth process of bifurcation; sites considered “indeterminate” were chosen where one blade or branch was impinging on another, or on some other nearby substrate, thus interfering with normal colony growth in a random, or non-ontogenetic, manner. This stereotypy is a rare example of demonstrable entrance-siting behavior by an unknown invertebrate organism. A skeleto-structural condition was present at both determinate and indeterminate locations on ramose trepostome colonies. Stereotypical entrance-site choices included a location where bryozoan freewall membrane was compromised or thinned and exozone was incompletely developed. Sanctum producers preferred incomplete exozone in order to access the endozone for excavation to form a domichnium. Middle and Late Ordovician diversification of ramose bryozoan forms stimulated escalation of bioeroder and symbiont taxa in living zoaria thus encouraging further trepostome evolution.     

Bioerosion in Late Permian Rugosa from reefal blocks (Hawasina Complex,Oman Mountains): implications for reef degradation

Summary Rugose corals are known from allochthonous Late Permian reefal blocks of the A1 Jil and Ba’id Formation (Hawasina Complex), Oman Mountains. In contrast to many Late Permian Rugosa found elsewhere in the Tethys, they occurred in sponge reefs and contributed to reef construction. The waagenophyllid warm water coral fauna is moderately diverse comprising cerioid, thamnasterioid, and fasciculate taxa. In contrast to sponges, chaetetids, and low-growing reefbuilders, the corals secreted diagenetically stable, most probably Mg-calcitic skeletons. Borings in coral skeletons are consequently well preserved providing important data for the interpretation of reef destructive processes. Thin-section analysis revealed three taxa of infaunal borers includingEntobia Bronn 1837, uncertain thallophyte borings, and borings of unknown bioeroders. Macroborers were more important than microborers, because of the dominance of clionid sponges. Good evidence exists also for the occurrence of two types of undetermined grazers which destroyed the coral surfaces. The amount and distribution of bioerosions is variable among different coral taxa. The fasciculate coralPraewentzelella regulare Flügel 1995 was the favorate substrate. Up to 33% of the calices were bored. Dendroid and compound corals were bored subordinately. Bioerosion of these colonies does not exceed 2%. There is good evidence for substrate preference amongst the borers. Major controlling factors affecting borer distribution are believed to be variations of skeletal density and gross morphology. The borer assemblage could not limit reef accretion significantly. Factors controlling boring activity might have been quality of substrate, sedimentation rate, rapid incrustation of substrates, and competition for food with reef constructors including sponges, chaetetids, and rugose corals.     

Boring by Macro-organisms in the Coral Montastrea annularis on Barbados Reefs

The fauna boring into Montastrea annularis includes sponges, bivalves, sipunculid and polychaete worms and barnacles. Sponges are most important in hard tissue destruction and account for more than 90% of the total boring in most heads. Bivalves and barnacles are locally important. Sipunculids and sabellids account for less than 4% of the total boring. The volume removed from coral samples by boring ranged from 3–60% and samples from a deeper bank reef were more highly bored than fringing reef samples. An average of 20% of the volume of bank reef corals, and 5% of the volume of fringing reef corals, was removed by boring. The distribution of individual borers is not a function of depth. The density and variety of borers and the extent of boring in coral heads is greater in older heads. The ratio of living coral surface to dead encrusted areas on colonies also influences borer density and the extent of boring.     

Bioerosion of gastropod shells: with emphasis on effects of coralline algal cover and shell microstructure

Organisms boring into fifty nine species of gastropod shells on reefs around Guam were the bryozoan Penetrantia clionoides; the acrothoracian barnacles Cryptophialus coronorphorus, Cryptophialus zulloi and Lithoglyptis mitis; the foraminifer Cymbaloporella tabellaeformis, the polydorid Polydora sp. and seven species of clionid sponge. Evidence that crustose coralline algae interfere with settlement of larvae of acrothoracian barnacles, clionid sponges, and boring polychaetes came from two sources: (1) low intensity of boring in limpet shells, a potentially penetrable substrate that remains largely free of borings by virtue of becoming fully covered with coralline algae at a young age and (2) the extremely low levels of boring in the algal ridge, a massive area of carbonate almost entirely covered by a layer of living crustose corallines. There was a strong negative correlation between microstructural hardness and infestation by acrothoracian barnacles and no correlation in the case of the other borers. It is suggested that this points to a mechanical rather than a chemical method of boring by the barnacles. The periostracum, a layer of organic material reputedly a natural inhibitor of boring organisms, was bored by acrothoracican barnacles and by the bryozoan. The intensity of acrothoracican borings is shown to have no correlation with the length of the gastropod shell.     

Trepostome bryozoans from the Zahořany Formation (Upper Ordovician) of Loděnice,Prague Basin,Czech Republic

Three trepostome bryozoan species are described from the Upper Ordovician Zaho?any Formation of Loděnice, Prague Basin, Czech Republic. One genus is new—Lodenicella gen. nov. One species is described in open nomenclature. The described fauna contains ramose colonies or ramose branched projections from encrusting tubular-shaped colonies which inhabited shallow environment with moderate wave energy and significant influx of clastic material.     

The Fossil Record of Shell Boring by Snails

The predatory boring habit common to many recent snails probablyarose first in the Polinicinae (Naticacea) during Upper Cretaceous(Cenomanian) times (100 million years B.P.) . In the fossilrecord the frequency of bored shells increasesgreatly in rocksof latest Cretaceous age and becomes more widespread duringearly Tertiary times coincident with the major diversificationof the primary groups of boring snails. The borings in these Cretaceous and Tertiary shells show thesame characteristics of preference of penetration in one pelecypodvalve rather than the other or in position of the boring siteon the shell that are found in recent shell assemblages. Borings in Paleozoic brachiopod shells (230–550 millionyears old) that have previously been attributed to gastropodpredation are herein attributed to other but unknown boringorganisms. In part these borings are not accepted as evidence of Paleozoicgastropod predation because it necessitates: (1) Postulationof the separate development of a boring habit ith its concomitantdevelopment of an accessory boring organ in a groupwhose descendantsare all herbivores, and (2) The development of such a habithundreds of millions of years before the appearance of any relativesof present day borers.     

Boreholes in Devonian blastoids and their implications for boring by platyceratids

Complete, circular holes found on the thecae of 51 specimens of Devonian nucleocrinid blastoids are interpreted as biogenic in origin. Their morphology, their stereotyped position on the theca, and the lack of multiple borings, suggest that they represent predation or parasitism. Platyceratid gastropods are presumed to be the borers because (1) they occur in the same localities as the bored blastoids, (2) they have been found on thecae of other blastoids, and (3) their boring abilities have been previously demonstrated. □ Paleozoic boreholes, platyceratids, blastoids, gastropod drilling.     

Unusual skeletal morphology and systematic description of a new Devonian cryptostome bryozoan from the Western Sahara

Palaeozoic bryozoan colonies display a large variety of skeletal elements. Various types of rod-like styles are mainly built by laminated and non-laminated (hyaline) skeleton. Typical styles consist of hyaline cores and surrounding laminated sheaths. They usually protrude on the colony surface as spines. In a new bryozoan genus from the Middle Devonian of the Western Sahara described here, styles do not protrude on the colony surface and are embedded within the laminated skeleton. They consist of fibrous material. This new type of styles, unknown from other bryozoans, is here called a “cryptostyle”. A special characteristic of these structures is their intensive reddish-brown colouration. This colouration is apparently caused by the presence of ferric iron between the individual fibres. The function of cryptostyles was apparently weight reduction and stabilising the skeleton. The general morphology of the new bryozoan, Cryptostyloecia hexapuncta gen. et n. sp., implies its systematic position within ptilodictyine cryptostomes, specifically the Family Ptilodictyidae Zittel, 1880.     

Recent borings in limestone cobbles from Marloes Bay, southwest Wales

Limestone clasts from the beach at Marloes Sands, southwest Wales, contain slender, straight to sinuous borings cross-cut by younger, clavate borings. The former were probably produced by sipunculids or polychaetes; the latter preserve shells of the boring bivalve Gastrochaena dubia (Pennant). Unusually, the calcareous linings of the clavate bivalve borings extend into many of the slender worm borings. Such linings are considered part of the hard parts of the producing bivalve, but the chance association of the two morphologies of borings has led to the lining becoming intimately associated with both of them. The modified linings of the bivalve borings have a similar morphology to the crypt of certain clavagellid bivalves, perhaps presenting an analogue for the morphology of a pre-clavagellid, boring ancestor.     

Palaeocology of Hard Substrate Faunas from the Cretaceous Qahlah Formation of the Oman Mountains

Skeletal encrusters and carbonate hardgrounds are rare in siliciclastic sands and gravels because of high levels of abrasion and sediment movement. An exception to this is the Maastrichtian Qahlah Formation of the Oman Mountains, a sequence of coarse siliciclastic sediments deposited on a shallow marine shelf above wavebase and at an equatorial palaeolatitude. This unit contains intercalated carbonate hardgrounds and other hard substrates which were encrusted and bored. The hard substrates, comprising carbonate and silicate clasts, calcareous bioclasts (mollusc shells and coral fragments) and wood, supported a diverse encrusting and boring fauna dominated in biomass by the oyster Acutostrea . There are twelve bryozoan species and at least two serpulid worm species, most living cryptically. Other encrusters on exposed surfaces include the agglutinated foraminiferan Placopsilina and several species of colonial corals. Borings in the carbonate clasts and shells are predominantly those of bivalves ( Gastrochaenolites ), with subsidiary clionid sponge ( Entobia ) and acrothoracican barnacle ( Rogerella ) borings. The woodgrounds are thoroughly bored by teredinid bivalves ( Teredolites ). Of the common substrate types, carbonate hardground clasts support the greatest number of taxa, followed by chert clasts, with limestone rockground pebbles being depauperate. Clast composition and relative stability probably explain these differences. Individual clasts probably had variable and typically long colonisation histories. Detailed palaeoecological interpretation is constrained by taphonomic loss, time-averaging and clast transportation and reorientation. Evidence from the Qahlah Formation shows that tropical rocky-shore biotas in the Cretaceous were not impoverished as previously believed.     

Symbiosis, competition, and physical disturbance in the growth histories of Pliocene cheilostome bryoliths

Free-living (unattached) subspherical bryozoan masses (bryoliths) in Pliocene tidal channel deposits of the Imperial Formation of southeastern California show complex intra- and interspecific interactions during their accretionary growth. Ranging up to 10 cm in length, the bryoliths are composed almost exclusively of the anascan cheilostome Biflustra commensale. Approximately 50% of the bryoliths are nucleated on cerithiid or muricoid gastropod shells; secondary occupants (presumably pagurid crabs) determined the subspherical growth and associated epibionts of all of these specimens. Evidence for crab occupation includes the thick and relatively symmetrical bryozoan overgrowths that form short tubes extending from the aperture, thinning and pinchout of laminae on the undersides of bryoliths (wear facets), and the distinctive borings (Helicotaphrichnus) of symbiotic worms. In some instances, shells were infested by bryozoans and other encrusters before death of the gastropod, but these overgrowths are thin relative to hermit-associated bryozoan colonies. Episodic hermit abandonment, indicated by extensive erosion of the bryolith and/or its colonization by a more diverse epibiont assemblage including oysters and serpulids, was more frequent among bryoliths nucleated on the largest and most fouled gastropod shells; it was also more frequent among bryoliths in the relatively high-energy tidal channel thalweg than among those associated with oyster thickets on muddy channel margins. Bryoliths nucleated on other shell substrata are similarly thick, but have more irregular stratigraphies including more sedimentary inclusions, more borings, and fewer encrusting epibionts. Pebbles of crystalline basement rock are also encrusted by B. commensale, but only thinly. All of these bryoliths not inhabited by crabs are limited to the channel thalweg. As many as four distinct colonies of B. commensale could coexist on a single bryolith; lines of competitive standoff between colonies are marked by mineralized walls and topographic ridges on the bryolith exterior, and by teepee-like structures in cross-section. These standoffs were preferred sites of infestation by other epibionts and were remarkably stable in position on bryoliths with continuous hermit occupation. Bryoliths that suffered repeated abandonment by hermits, or that depended entirely upon chance reorientation, are characterized by highly unstable standoff positions, reflecting scramble competition under less predictable conditions. These circumstances were most common among large bryoliths and among those in channel thalwegs.     

中国湘南早侏罗世双壳类壳体内的遗迹化石--兼论遗迹化石属Talpina Hagenow, 1840

论文描述的双壳类壳体内的钻孔遗迹组合发现于湖南省东南部宜章县下坪附近的心田门组上部(早侏罗世,早辛涅缪尔期)浅海沉积地层中。这个遗迹化石群中,帚虫类动物制造的钻孔Talpinahunanensisichnosp.nov.(新遗迹化石种)明显地占据了大部分,偶尔共生的其它钻孔包括:Rogerellaichnosp.(蔓足亚纲尖胸类的遗迹)、Calcideletrixichnosp.(可能是藻类的遗迹)、藻或菌造的遗迹(暂定为"Mycelites"ichnosp.)、以及一种可能被海绵动物制造的未定钻孔。生物侵蚀者侵害了海底上较大的双壳类壳体。钻孔保存为天然印模(石核),它们是文石质壳体溶解后留下的空间中的充填物。此外,文中还讨论了遗迹化石属TalpinaHagenow,1840,尤其讨论了有关遗迹种颇为复杂的鉴别问题,同时校正了该遗迹属的属征。     

The effects of ambient flow velocity,colony size,and upstream colonies on the feeding success of bryozoa. I. Bugula stolonifera Ryland,an arborescent species

The effects of ambient flow velocity, colony size, and the presence of upstream colonies on the feeding success of the arborescent bryozoan, Bugula stolonifera (Ryland), were studied. Faster ambient flow velocities were found to reduce feeding of zooids of small colonies but not of large colonies. Zooids from different regions of colonies dominated in feeding at different ambient flow velocities: upstream zooids dominated in feeding from slow ambient flow: zooids from central regions dominated in feeding from fast ambient flow. These results are interpreted with respect to the branching morphology of colonies. Finally, evidence that upstream colonies interfere with the feeding success of zooids of colonies downstream was obtained.     

Three phyla—Two type specimens—One shell: History of a snail shell revealed by modern imaging technology

In a recent revision of the gastropod genus Bellardiella, it became evident that the neotype of B. crassilabris had numerous borings over its entire shell surface. Superficial and light microscopical observation of the shell showed that the borings were made by a bryozoan belonging to the genus Penetrantia. Conventional techniques such as endocasting with the dissolution of the shell are impossible for type material. The application of non‐destructive micro‐CT‐scanning of the shell revealed the three‐dimensional arrangement of the entire colony within the shell. Careful analysis of the structure of the autozooids of the colony revealed that the species could not be assigned to any known species; we accordingly describe it as Penetrantia bellardiellae Schwaha, n. sp. The application of micro‐CT not only allowed the depiction of the new bryozoan species but also revealed the remains of a hermit crab located within the shell. The shell of Bellardiella crassilabris thus represents the type specimen of two species and contains a member of a third phylum inside. To our knowledge, this is a unique case, which also emphasizes the usefulness of micro‐CT imaging for precious specimens as well as the importance of historical museum collections.     

Organization and Evolution of Animal Colonies

Based on the theory of organization and evolution of colonies in an extinct group of hemichordate graptolites (Urbanek, 1960, 1990) the relationship between the events in the late astogeny of bryozoan colonies and their somatic and reproductive cycle is proposed. The bryozoan colonies with simple organization are compared with graptoloid colonies and their structure is interpreted within the framework of the morphogen gradient theory. A morphogen produced by the founder-zooid (oozooid) diffuses along the long axis of the colony and controls the phenotypic expression of the size and structure of zooids. Evolutionary changes in the graptoloid colonies involve introduction of new characters and their spreading is also accompanied by gradient changes of their manifestation. Evolutionary mechanisms in bryozoan colonies are considered in terms of penetrance and expressivity of genes. In contrast to graptolites, many bryozoan colonies display multiple zones of astogenetic changes and repetitions.     

The effects of ambient flow velocity,colony size,and upstream colonies on the feeding success of Bryozoa. II. Conopeum reticulum (Linnaeus), an encrusting species

The effects of ambient flow velocity, colony size, and the presence of an actively-feeding colony upstream on the feeding success of the encrusting bryozoan Conopeum reticulum (Linnaeus) were studied. Zooids from both large and small colonies showed a reduction in feeding as flow velocity increased, however, the reduction in feeding was less for zooids from large colonies except at very fast ambient flow velocities. The greater pumping capacity of large colonies may result in a relatively greater per zooid feeding success from moving water. The presence of an actively-feeding colony upstream was found to enhance the feeding of zooids on downstream colonies. Diversion of flowing water by actively-feeding colonies upstream may account for the observed enhancement of feeding by zooids on colonies downstream.The results from this study on an encrusting species are compared with results from a previous study on feeding from flow by an arborescent bryozoan, and the feeding performances of these two colony types are related to their respective flow microhabitats.     

Paleontological evidence of a brief global sea-level event during the last interglacial

A gently undulating to flat erosion surface with shallow-water borings and burrows is present in the midst of a Sangamonian (Eemian; marine oxygen isotope substage 5e) reefal facies on the islands of San Salvador and Great Inagua, Bahamas. Precise U/Th dating of corals above and below this surface show that it formed around 125-124 ka, and that the sea-level regressive-transgressive cycle which produced it lasted for 1500 years or less. The surface occurs on entirely carbonate rocks and has a low relief punctuated by erosional channels and karstic caves formed during the sea-level lowstand. A terra rossa paleosol, developed during that lowstand, partially fills a set of large lithophagid bivalve borings ( Gastrochaenolites torpedo ), showing that they were excavated during the regression. Rhizomorphs formed by plant roots occur on the erosion surface at Great Inagua. Extensive boring of the upward-facing surfaces occurred during the ensuing transgression, including a smaller G. torpedo and a clionid sponge boring ( Entobia ovula ). The bored surface is encrusted by a variety of shallow-water corals and, eventually, the re-established bank-barrier coral reefs. A sparse assemblage of serpulid worm and vermicularid gastropod tubes encrusted the channel and cave walls. Robust Ophiomorpha burrow systems occur within pockets of sediment in the coral facies both below and above the erosion surface. The channels and caves are filled with transgressive calcarenitic sediments in which occur numerous Ophiomorpha and Skolithos burrows. The ichnofossils on, below, and above this erosion surface are prominent indicators of a short-lived but significant global sea-level event.     

Schischcatella (Fenestrata,Bryozoa) from the Devonian of the Rhenish Massif,Germany

Abstract: The Devonian fenestrate bryozoan, Schischcatella Waschurova, 1964 , possessed colonies in the form of low, erect bifoliate fronds that grew from an encrusting sheet‐like base with autozooecia arranged in biserial, bifurcating rows. This growth habit is unique in fenestrates, which normally had unilaminate arborescent colonies. Originally, Schischcatella was described from the Lower Devonian of Tajikistan. This article describes a new species, S. heinorum sp. nov., from the Middle Devonian of the Eifel (western Rhenish Massif, Germany) with additional material from the Lower Devonian of the Kellerwald (eastern Rhenish Massif, Germany). External and internal morphologies of this bryozoan have been studied using abundant material. The growth habit of Schischcatella suggests a completely different pattern of feeding currents than that in the normal fenestrate colony. The outflow of the filtered water occurred only on edges of colonies between rami. In the absence of chimneys (areas of vertical water expelling), such a functional morphology may have restricted extension of the colony in a distal direction. The evolution of Schischcatella is apparently an example of paedomorphosis, the genus evolved from an unknown semicosciniid species by the early ontogenetic interruption of colony development and further changes in the mode of growth.     

Temporal variations of macroborers in massive Porites lobata on Moorea,French Polynesia

Massive colonies of Porites lobata on the barrier reef of Tiahura, Moorea, can be divided into four categories: living colonies, colonies consisting of 50% live coral and 50% dead skeleton, 100% dead coral and colonies which have been reduced to a basal plate. Replicate samples of each of these colony types were collected in the same vicinity of the barrier reef during October 1987. The macroborers were extracted, identified, counted and their volumes determined by displacement. Kruskal-Wallis tests showed that three different boring communities occur within these four categories of Porites colonies. Live colonies are characterised by only 3 species, the bivalve, Lithophaga laevigata; the vermetid Dendropoma maximun; and the non boring serpulid polychaete Spirobranchus. The completely dead colonies contain up to 17 boring species, with five to six individuals per 100 cm. Sipunculans are the dominant bioeroders with the most abundant species being Aspidosiphon elegans, sp A and sp B. Colonies of Porites which have been reduced to basal plates contain up to 18 boring species of which the bivalve Lithophaga hanleyana and the sipunculan Aspidosiphon sp. B are the most abundant.The cumulative volume of CaCO₃ lost by boring activity increases from 0.1 cm³ per 100 cm³ in a completely dead Porites colony to 1.4 cm³ per 100 cm³ in the residual basal plates of Porites. These can be extrapolated to minimum losses of 14.2 kg m^-3. We suggest that rates of boring increase with the time which has elapsed since the death of the colony and the dominant agents of boring also change with increasing age of the coral structure. There are significant additional losses of 5.25 kg m^-3 CaCO₃ caused by grazing echinoids and scarids.     

The inference of extrazooidal feeding currents in fossil bryozoan colonies

Taylor, Paul D. 1979 01 15: The inference of extrazooidal feeding currents in fossil bryozoan colonies. Lethaia , Vol. 12, pp. 47–56. Oslo. ISSN 0024–1164.
Previous studies on live bryozoan colonies have shown that the feeding autozooids in a colony may cooperate in differing ways to produce an extrazooidal watercurrent system organised on a colony-wide or subcolony-wide basis. The presence of an extrazooidal current system may be inferred in fossil stenolaemate bryozoans which exhibit either a differential spacing of open autozooecial apertures or a systematic variation across the marial surface in the orientation of automoecial distal portions. By inference, aggregations of autozooecial apertures represented loci of inhalant extrazooidal flow whereas zoarial protuberances (e.g. monticules) with outwardly leaning autozooecia acted as loci of exhalant extrazooidal flow. Bryozoans having automoecia opening obliquely into gaps or fenestrules in their zoaria probably drew a unidirectional extrazooidal current of water through the fenestrules. Extrazooidal water currents may function to accelerate colony clearance rate, decrease the chances of recycling filtered water, aid spermatoman and larval dispersal, and clear sediment from the colony surface.