Epizoans on late Ordovician brachiopods from Southeastern Indiana

The frequency of epizoans (cornulitids, inarticulate brachiopods, bryozoans, solitary and colonial rugosan corals) on over 8000 specimens of articulate brachiopods (four strophomenids, five orthids, one rhynchonellid) was calculated for four stratigraphic horizons in the Dillsboro Formation of southeastern Indiana. Frequency of shells encrusted correlates significantly with the surface area of the valves. Punctae in brachiopod shells (Onniella meeki) may have deterred larval settlement of epizoans. Coarse ribbing on articulates deterred encrustation by the inarticulate brachiopod. The horn coral shows a preference for attachment to the anterior of Hiscobeccus capax. Bryozoans show a preference for the incurrent lateral margins of inferred living hosts, suggesting rheotropic behavior by settling larvae. Inarticulate brachiopods are concentrated around the sloping commissure of the brachial valve of strophomenids, suggesting geotropic behavior and/or selective survival of settling larvae. Inarticulates deterred overgrowth by bryozoans. High frequencies of encrustations of the medial region of pedicle valves of orthids and strophomenids probably reflect post‐mortem encrustations. Alternating intervals of slow sediment accumulation punctuated by tropical storms and rapid shell burial may account for the high frequency of shells with either their entire surface veneered or only a very small area encrusted by bryozoans.     

Survival and repair of surgical and natural shell damage in the articulate brachiopod Terebratulina retusa (Linnaeus)

Living specimens of Terebratulina retusa from the Firth of Lorn, Scotland, were surgically damaged by drilling 2 mm diameter holes or narrow slits one cm long in the anterior portion of one valve, by bevelling the anterior margin of both valves, or by amputation of the anterior third of one valve. These injuries to the shell and mantle simulated the type of repaired shell damage seen in Paleozoic species, i.e., scalloped, divoted, cleft, and embayed valves. Less than ten percent of the 200 damaged specimens survived until the 25th week after surgery. Specimens of T. retusa showed the ability to repair drill holes, slits, and bevelled anterior shell regions, but not the most severe damage, i.e., amputations of the anterior third of one valve. Shell‐repair was initiated in the fourth week after surgery by the development of a membrane across the wound. The development of caeca in the new shell layer secreted to plug the drill holes became apparent by the eighth week. The punctate pattern was complete in the new, translucent shell material of bevelled and drilled specimens by the 25th week following surgery. Failure of any specimens to survive amputation of the anterior portion of a valve for more than seven weeks after surgery, and the absence of initiation of the repair process, suggests that terebratulids do not have the tolerance for, nor the ability to repair, the severe injuries (embayed valves) which were sustained and mended by extinct strophomenids.     

Mechanical strength of shells of selected extant articulate brachiopods: Implications for paleozoic morphologic trends

Dried shells of Terebratalia transversa, Laqueus californianus, Hemithyris psittacea, and T. unguicula and alcohol‐soaked, tissue‐lined shells of Terebratulina retusa, Dallina septigera, Cryphus vitreus, and Liothyrella uva were crushed in an apparatus that facilitated measurement of the force (newtons) against the valves at the instant of fracture. The results revealed that the costate shells of T. transversa and T. retusa were the strongest. Force is correlated with valve thickness, but not with size (length). When normalized for valve thickness, the force required to fracture shells is correlated with shell biconvexity (height/length) among pooled species of dried specimens. Geniculate specimens of T. retusa were not stronger than the intraspecific variants with a constant radius of curvature to their valves.

The percent‐frequency of plicate, spinose, lamellose and rugate genera increase significantly in the successive stages, Caradocian (Late Ordovician) through Famennian (Late Devonian) at the expense of smooth to costellate genera. The percent‐frequency of rectimarginate (central fold lacking) genera also decreases appreciably in this time frame. These morphologic trends, in combination with the experimental crushing data, support the hypothesis that selection favored species with such anti‐predatory adaptations during a time of escalation of shell‐crushing predators.     

Predatory boreholes in Tournaisian (Lower Carboniferous) spiriferid brachiopods

A brachiopod fauna from the uppermost part of the Tournaisian Tournai Formation (Belgium) contains an undetermined species of Crurithyris (Spiriferida, Ambocoeliidae), which displays numerous bored shells. About 8% of the 432 specimens with conjoined valves display single, small (≤ 1 mm) boreholes, which are smooth-sided, cylindrical or weakly conical, circular to slightly elliptical in plan view, perpendicular to the shell surface and generally complete. Of the 35 bored articulated specimens, 27 were drilled on the ventral valve. Most of the boreholes are located in the posterior half of the shell, and no case of edge-drilling has been observed. The boreholes were drilled by a predator, or possibly a parasite, which selected individuals greater than 2.5 mm long. Crurithyris sp. may have represented an attractive (in terms of energy cost) and easy target for a small-sized predator because of its thin shell and ornament of minute spines.     

Population paleoecology and comparative taphonomy of two edrioasteroid (echinodermata) pavements: Upper ordovician of kentucky and Ohio

Two strophomenid brachiopod shell pavements bearing abundant edrioasteroid epizoans occur stratigraphically within the top 4.5 m of the Corryville Formation (Upper Ordovician, Maysvillian) in Boone County, Kentucky, and Hamilton County, Ohio. Both assemblages are dominated by Isorophus cincinnatiensis and contain Carneyella pilea and Streptaster vorticellatus as lesser constituents. Pedicle exterior valves of Rafinesquina “alternata”; are the preferred substratum for all species at both sites, but the Kentucky population occurs mainly on unabraded, articulated shells and the Ohio population occupies abraded, disarticulated shells. The Kentucky population includes a greater size range of Isorophus (at least three cohorts) than the Ohio population (at least two cohorts), suggesting a greater time span of colonization for the Kentucky site. Host shells from Kentucky have anterior‐posterior axes strongly aligned NE‐SW whereas the Ohio shells are unoriented. Isorophus in Kentucky show a preference for the anterior margin of the Rafinesquina pedicle exterior valve, whereas Isorophus in Ohio show no preferred location on host shells. The Kentucky edrioasteroids are inferred to have attached to mainly live Rafinesquina in brachial‐valve‐up position. Host shells were inverted and oriented by a NE‐SW storm surge, then buried rapidly. Ohio edrioasteroids occupied mainly dead, already inverted host shells not oriented by storm surge possibly because of their more downslope (distal) location. Comparative taphonomic relationships between the two coeval populations are consistent with a model of proximal to distal distribution along a gentle shallow subtidal N‐S paleoslope subject to episodic storm activity.     

Sexual dimorphism in shells of Cochlostoma septemspirale (Caenogastropoda, Cyclophoroidea, Diplommatinidae, Cochlostomatinae)

Sexual dimorphisms in shell-bearing snails expressed by characteristic traits of their respective shells would offer the possibility for a lot of studies about gender distribution in populations, species, etc. In this study, the seven main shell characters of the snail Cochlostoma septemspirale were measured in both sexes: (1) height and (2) width of the shell, (3) height and (4) width of the aperture, (5) width of the last whorl, (6) rib density on the last whorl, and (7) intensity of the reddish or brown pigments forming three bands over the shell. The variation of size and shape was explored with statistical methods adapted to principal components analysis (PCA) and linear discriminant analysis (LDA). In particular, we applied some multivariate morphometric tools for the analysis of ratios that have been developed only recently, that is, the PCA ratio spectrum, allometry ratio spectrum, and LDA ratio extractor. The overall separation of the two sexes was tested with LDA cross validation.The results show that there is a sexual dimorphism in the size and shape of shells. Females are more slender than males and are characterised by larger size, a slightly reduced aperture height but larger shell height and whorl width. Therefore they have a considerable larger shell volume (about one fifth) in the part above the aperture. Furthermore, the last whorl of females is slightly less strongly pigmented and mean rib density slightly higher. All characters overlap quite considerably between sexes. However, by using cross validation based on the 5 continuous shell characters more than 90% of the shells can be correctly assigned to each sex.     

Comparative hydrodynamic stability of brachiopod shells on current-scoured arenaceous substrates

Selected brachiopod specimens were placed on a well-sorted medium grained sand (0.5 mm) in a recirculating flume and subjected to a constant unidirectional current of 26–27 cm/sec. The specimens were placed in each of six feasible life-orientations. In three orientations, the valves were reclining with either the lateral, anterior or posterior profile upcurrent. In three orientations, the commissural plane was perpendicular to the substrate with either the pedicle or brachial valve upcurrent, or a lateral margin upcurrent. Destabilization indices were calculated for each specimen in each orientation based on the elapsed time until the specimen reoriented. Nonstrophic biconvex brachiopods are most stable with the valves reclining on the substrate. Among orientations with the valves erect, the pedicle valve upcurrent and the brachial valve upcurrent are the most and least stable orientations, respectively. Shell stability increased among nonstrophic specimens that were more equi-biconvex. Biconvex strophic brachiopods are also more stable if the valves were equi-biconvex, cither compressed (biplanate) or inflated, provided that the specimen was propped up on its hingeline with a lateral margin upcurrent. Catacline interareas afforded greater stability than procline, apsacline or orthocline interareas when the valves were perpendicular to the substrate. Accentuated plication reduced shell stability. Alate biconvex forms were often more stable if the valves were vertical to the substrate rather than reclining. Alate biconvex geometries are more stable than nonstrophic and strophic biconvex forms if the commissural plane is perpendicular to the substrate and parallel to the current. Broad catacline interareas increased stability in all orientations. Plano-convex and dorsi-biconvex alate forms are usually less stable than equi-biconvex or ventri-biconvex alate geometries. Concavo-convex geometries are stable in all orientations except with the valves nearly vertical to the substrate and parallel to the current. Spines greatly retard sediment-scour and maintain concavo-convex specimens in orientations with the valves elevated above the substrates. D Brachiopoda, shell shape, hydrodynamic stability.     

Microbilobata,a new genus of earliest terebratulid brachiopod from the lower Silurian of Northwestern Canada: Implications for the origin of higher taxa

Microbilobata avalanchensis n. gen. and n. sp. from the Lower Silurian (upper Wenlock) carbonate rocks of the lower Delorme Group in the Avalanche Lake area, northwestern Canada, is described here as the earliest known terebratulid brachiopod. These small shells (less than 2 mm long) are subtriangular, anteriorly emarginate, possibly punctate, with both valves being sulcate at their anterior halves, the ventral sulcus bearing one prominent plica, and the dorsal sulcus marked by two plicae. Internally, M. avalanchensis has a centronelliform loop extending for about three fifths of the total shell length. The shells are silicified in carbonate rocks of mid to outer shelf origin. M. avalanchensis is relatively rare, with about 40 specimens so far found from samples collected at 58–60 m above the base of section AV5 in the Avalanche Lake area. Its presence in rocks of Wenlock age extends the earliest known terebratulids back about 16 million years from the oldest previously recorded terebratulids (earliest Devonian age). The small size and simple form of the new species suggest that heterochrony (progenesis) could have played a role in the origin of the Terebratulida. M. avalanchensis serves as a good example of Cope's Rule, indicating that the terebratulids evolved from a very small, unspecialized ancestor. The unusually small size of this taxon also offers one explanation as to why some ancestors or transitional forms of major taxonomic groups are extremely difficult to find in the fossil record.     

Structure and formation of the adventitious tube of the Japanese watering-pot shell Stirpulina ramosa (Bivalvia, Anomalodesmata, Clavagellidae) and a comparison with that of the Penicillidae

Abstract. Stirpulina ramosa is the only extant endobenthic representative of the Clavagellidae and is restricted to the waters of Japan. A single intact adventitious tube of this species has been obtained and its structure is described. The right valve is 16 mm long and located within the adventitious tube. It has an opisthodetic ligament located on resilifers. There are anterior and posterior adductor muscle scars, a thick pallial line, and pallial and pedal gape (right valve only) sinuses. The left shell valve is but 9 mm long and is united into the fabric of the adventitious tube via the intermediary of a shelly saddle. Internally, only the anterior adductor muscle scar and a small element of the pallial line scar are identifiable on the left valve. The posterior adductor and the rest of the pallial line scar (including a pallial sinus) are, remarkably, located on the adventitious tube beyond the shell valve margin. The adventitious tube of S. ramosa is formed in a manner wholly dissimilar from that of Brechites vaginiferus (Penicillidae). In B. vaginiferus, the tube is secreted as a single entity from the general outer mantle surface, including the siphons, covering the body. As a consequence, both shell valves are incorporated into the structure of the tube and the watering pot is bilaterally symmetrical. In S. ramosa, the tube and watering pot are secreted from the mantle margin and surface surrounding and extending from the left shell valve, so that only the left valve is incorporated into its structure. A dorsally derived mantle element is progressively extended over to the right side of the body, meeting a ventrally derived counterpart that passes beneath it, forming a pleat in the calcareous structure of the right side of the tube that they secrete. This pleat extends into the complex of watering‐pot tubules and forms the pedal gape. The watering pot is thus Ω shaped. The ventrally derived mantle element forms a sinusoidal crest on the right‐hand base of the watering pot, creating a pedal gape sinus scar on the right valve. The Clavagellidae radiated widely in the Mesozoic, leaving behind a rich fossil record for Stirpulina. Only S. ramosa, however, has survived until the present. In contrast, the Cenozoic Penicillidae has a poor fossil record, but there is a rich variety of extant endobenthic watering‐pot shells. It has been argued hitherto that the two families represent a remarkable example of convergent evolution. In view of the success of the Penicillidae and thus the endobenthic, tube‐dwelling lifestyle, however, it is hard to understand why Stirpulina has largely died out—even S. ramosa being known by but one or two specimens. A study of the anatomy of S. ramosa might one day answer this question.     

Operculum regeneration following failed predation in the Silurian gastropod Oriostoma

Regeneration of the calcareous rigiclaudent operculum following severe, non‐lethal, predatory attacks is described in two specimens of the characteristic Silurian gastropod Oriostoma from the English Midlands. This is the first record of the regeneration of opercula in Palaeozoic gastropods and of opercula conjoined with shells that have been repaired after significant, failed, durophagous attacks. In one specimen, a series of repaired injuries culminates in a repaired aperture closed by the operculum. In the second specimen, new shell growth has failed to recover the full extent of the original broken whorl, and the new aperture and operculum, though well developed, are smaller than the originals. Both opercula are unusually thin centrally when compared to other, strongly domed, Oriostoma opercula.     

我国硅藻类的新记录属——微舟藻属及其一新种(英文)

利用透射电镜对采自东黄海海域典型站位的微型硅藻(nanodiatom,<20mm)进行了研究。观察到我国微型硅藻类的一个新记录属微舟藻属NanoneisR.E.Norris以及该属的一个新种长微舟藻Nanoneislongtasp.nov.。该属的主要特征是壳面两侧对称,具有不完全壳缝,壳面除中轴区外都有横肋纹或长室孔分布。本文描述该属的两个种,其中海斯微舟藻(NanoneishasleaeR.E.Norris)为我国的新记录种,长微舟藻为新种,两种之间主要的区别特征是壳面外形、长宽比例以及壳面横肋纹的密度。并较详细地描述了该属种类的分类特征、生态习性和分布。     

A CAUTIONARY TALE: RAZOR SHELLS, ACORN BARNACLES AND PALAEOECOLOGY

Abstract:  The extant, introduced razor shell Ensis americanus (Binney) is a burrowing inhabitant of sandy, shallow-water substrates off the North Sea and Wadden Sea coasts of The Netherlands and adjacent areas. Three articulated shells with broken valves, collected from the strandline at Zandvoort, Noord-Holland, in April 2006 have dense skeletozoan infestations of the barnacle Balanus crenatus Brugière on the outer and inner surfaces of all valves. Such infestations must have occurred after death of the bivalve, decomposition of the soft tissues (but not the ligament) and disinterment of the shells. Larvae of B. crenatus settle in the spring, suggesting that these infestations are perhaps less than a year old, and testifying to the post-mortem persistence of the ligament and the density of skeletozoan infestation after a geologically brief duration. Such specimens would be a palaeoecological conundrum if fossilized; however, the ligament is likely to rot before final burial and the valves break further. Fossil specimens would probably be interpreted as valves that became encrusted on their inner and outer surfaces (balanuliths) during a long post-disarticulation residence on the sea floor.     

The tube-like structures on the juvenile shells of earliest strophomenides and billingsellides as evidence of their life cycles

Possible life cycle of some ancient plectambonitoids (order Strophomenida) from the Middle Ordovician of Russia is reconstructed based on the well-preserved specimens composing the ontogenetic series. Four regions may be distinguished on their shell surface: protegulum, brephic shell, neanic shell and adult shell. The posterior margin of ventral protegulum bears pedicle sheath, which is a tubular outgrowth with a 40-μm-wide aperture at the distal end. The protegulum and brephic shell have common type of microstructure that possibly is spherular; the neanic and adult shells are fibrous. The strophomenide ontogeny possibly was similar to that of recent discinides. The strophomenide life cycle possibly included the planktotrophic juvenile stage; the protegulum and brephic shell were formed in the water column. The aperture of the pedicle sheath was possibly used as an anal opening of the floating juvenile and as an attachment organ during the settlement; at early adult stages, the sheath erased, the anus closed, and the animal started to lie on the ventral valve. The origin of the order Strophomenida and its relative groups is possibly connected with the loss of the pedicle lobe; judging by some strophomenide morphological features, true pedicle was present in the strophomenide ancestors. The tubes on the ventral umbones of strophomenides and billingsellides are not homologous as pedicle sheaths of strophomenides are formed at the planktotrophic swimming stage, and the tubes surrounding the pedicles of billingsellides were formed by deltidial plates of almost adult shell after settling.     

Variability of shell repair in the Manila clam Ruditapes philippinarum affected by the Brown Ring Disease: a microstructural and biochemical study

For more than two decades, the Manila clam Ruditapes philippinarum has been regularly affected by Brown Ring Disease (BRD), an epizootic event caused by the bacterium Vibrio tapetis and characterized by the development of a brown deposit on the inner face of valves. Although BRD infection is often lethal, some clams recover by mineralizing a new repair shell layer, which covers the brown deposit and fully isolates it from living tissues. In order to understand this specific shell repair process, the microstructures of repaired zones were compared to those of shells unaffected by BRD. In addition, the organic matrix associated with unaffected shells and to repair patches were extracted and compared by biochemical and immunological techniques. Our results show that the repaired zones exhibit microstructures that resemble the so-called homogeneous microstructure of the internal layer, with some marked differences, like the development of crossed-acicular crystals, which form chevron-like patterns. In the three tested batches of repaired layers, the matrices exhibit certain heterogeneity, i.e., they are partially to widely different from the ones of shells unaffected by BRD, as illustrated by SDS-PAGE and by serological comparisons. Our results strongly suggest a modification of the secretory regime of calcifying mantle cells during the shell repair process. Polyclonal antibodies, which were developed against specific protein fractions of the shell, represent relevant tools for localizing by immunohistology the cells responsible for the repair.     

Belichnus Traces Produced on Shells of the Bivalve Lutraria lutraria by Gulls

In February 2011, after a storm, thousands of adult, articulated, and still-living common otter shells Lutraria lutraria (L. 1758) were stranded on the North Sea beach of the Island Texel (NL). These 9 to12 cm long bivalves were rapidly found and consumed by both herring- and lesser black-backed gulls. Holes, irregular in outline, were observed in some 10% of the articulated shells of these bivalves. These holes were always smaller on the outside of the valves than on the inside and varied in size from 1 to 20 mm (outside) to 4 to 22 mm (inside). Often the other valve was crushed indicating consumption by gulls. We concluded that these holes were made by the gulls probing the shells; in a few cases, we observed that valves were broken starting from such a hole. Such traces are described in the literature as the ichnogenus Belichnus and were until now attributed to Stomatopoda only. We also suggest that a separate ichnospecies name for two Belichnus holes in one shell should not be used, as we see them simply as a double injury due to two blows. Our findings stress once more the importance of avoiding premature phylogenetic interpretation of traces and the use of a separate ichnotaxonomy.     

New brachiopod subfamily Anechophragmiinae (Strophomenida) from the Ordovician of the Leningrad Region

A new strophomenid subfamily Anechophragmiidae is distinguished based on the peculiarities of cardinalia. The strophomenids of this subfamily lack cardinal process at all developmental stages; their socket ridges accrete and form a high plate, which closes the pedicle opening. Two genera are referred to the subfamily: Anechophragma Neuman, 1976 and Biseptata gen. nov. with new species B. briani sp. nov. from the Ordovician of the Leningrad Region. The shell structure, exterior, endoskeleton and ontogeny of A. rarum and B. briani were studied in detail due to the excellent preservation of the material. Two species are referred to Anechophragma: A. rarum Neuman, 1976 and A. alexandrae (Andreev, 1993).     

Shell abnormalities in scaphitid ammonites

The kinds incidence of shell abnormalities were investigated in over 2000 adult Late Cretaceous scaphitid ammonites. Most are species of Hoploscaphites Discoscaphites from the Maastrichtian Fox Hills Formation; for comparison, four older species of Scaphites were also studied. Four kinds of abnormalities vary in their incidence among the Fox Hills taxa but display no general trends. (1) Morton's syndrome, shell asymmetry unrelated to external injury, occurs in 3 to 6% of the Fox Hills scaphites. (2) Suture wander. veering-off of the ventral lobe of the suture from the midline of the venter, occurs in most Fox Hills taxa with an incidence ranging from 1 to 11%. (3) Stretch pathology, an attenuation of the shell on the posterior part of the adult body chamber, is confined to Hoploscaphifes nicolleti. (4) Externally inflicted injury, indicated by repaired damage on mature shells, has an average incidence of 10% in Fox Hills scaphites except in the stratigraphically youngest H. nebrascensis H. cheyennewis which approach 25 40% respectively. Good correlation between percent injuries adult size ornamentation is obtained only in Discoscaphifes; all morphs of the co-occurring Hoploscaphites display nearly the same incidence of injuries regardless of their morphologic variation. The stratigraphically youngest H. nebrascensis becomes larger more nodose simultaneously sustains more injuries. However, its dimorph H. cheyennensis remains morphologically unchanged although its incidence of injuries quadruples. These injury data, in themselves, do not support the hypothesis that predators were contributing more heavily to the mortality of Maastrichtian ammonites precipitated their extinction.     

Feeding behavior,ration and size structure of a population of the predatory gastropod <Emphasis Type="Italic">Cryptonatica janthostoma</Emphasis> in Vostok Bay,Sea of Japan

The relationships between the shell height of the predatory gastropod Cryptonatica janthostoma and the shell length of its typical prey, the bivalve Ruditapes philippinarum, and the diameter of the borehole on the prey shell resulting from a successful attack of the predator were experimentally found and assessed statistically. The shell height of C. janthostoma calculated retrospectively from the borehole diameter using the obtained relationships was 17–52 mm. The prey of C. janthostoma are burrowing bivalves, whose populations are affected by the predator to a varying degree. In populations of medium-sized mollusks (R. philippinarum, Protothaca euglypta, P. jedoensis, and others), C. janthostoma feeds on mollusks larger than 7–10 mm; in species with a shell length greater than 100 mm (Callista brevisiphonata, Saxidomus purpuratus), it eats specimens of 10–58 mm. C. janthostoma apparently has no effect on populations of small-sized mollusks (Anisocorbula venusta) and mollusks with an active avoidance response (Clinocardium californiense).     

ADAPTATION FROM RESTRICTED GEOMETRIES: THE SHELL INCLINATION OF TERRESTRIAL GASTROPODS

The adaptations that occur for support and protection can be studied with regard to the optimal structure that balances these objectives with any imposed constraints. The shell inclination of terrestrial gastropods is an appropriate model to address this problem. In this study, we examined how gastropods improve shell angles to well‐balanced ones from geometrically constrained shapes. Our geometric analysis and physical analysis showed that constantly coiled shells are constrained from adopting a well‐balanced angle; the shell angle of such basic shells tends to increase as the spire index (shell height/width) increases, although the optimum angle for stability is 90° for flat shells and 0° for tall shells. Furthermore, we estimated the influences of the geometric rule and the functional demands on actual shells by measuring the shell angles of both resting and active snails. We found that terrestrial gastropods have shell angles that are suited for balance. The growth lines of the shells indicated that this adaptation depends on the deflection of the last whorl: the apertures of flat shells are deflected downward, whereas those of tall shells are deflected upward. Our observations of active snails demonstrated that the animals hold their shells at better balanced angles than inactive snails.