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.     

The functional significance of aperture form in gastropods

Aperture form of marine prosobranch gastropods has evolved under the influence of a number of different selective forces, including: generation of shell form; protection from predation; accommodation of the foot during clamping behavior: and accommodation of water currents in and out of the mantle cavity. Aperture form correlates positively with foot shape in most gastropods and foot shape, in turn, correlates moderately well with substrate preference. Almost all gastropods that have non-round apertures elongate the aperture parallel to the foot so that water currenth tend to flow anteriorly to posteriorly. Fresh-water pulmonates have responded to somewhat different stresses. They exhibit clamping behavior and thus show correspondence between foot shape and aperture shape. They show less apertural strengthening as crab (or crayfish) predation is less of a factor and presumably because calcium carbonate is less available. They also lack anterior-posterior apertural elongation due to the absence of water currents through their mantle cavity. Due to the absence of mantle cavity water currents and clamping behavior, terrestrial gastropods do not show the apertural modifications associated with these two factors. In addition. few adaptations of apertural form are present to resist predation. Instead, many of the apertural modifications of terrestrial pulmonates seem to be concer-ned with the problems of water loss during estivation.     

Latest Helcionelloid Molluscs from the Lower Ordovician of Kazakhstan

The helcionelloid mollusc Chuiliella elenae gen. et sp. nov. is described from the Lower Ordovician of Kazakhstan. It represents the geologically youngest record of a group of mainly bilaterally symmetrical ancient molluscs which originated in the earliest Cambrian, flourished during the early–mid Cambrian and was thought to have become extinct during the late Cambrian. Chuiliella is a typical helcionelloid in terms of shell shape, although the comarginal ornamentation characteristic of many helcionelloids is lacking. Interpretation of the raised margin of the aperture adjacent to the earlier coiled whorl as exhalant channels favours reconstruction of helcionelloids as endogastrically coiled, i.e., with the apex posterior.     

Brachiopod shell thickness links environment and evolution

While it is well established that the shapes and sizes of shells are strongly phylogenetically controlled, little is known about the phylogenetic constraints on shell thickness. Yet, shell thickness is likely to be sensitive to environmental fluctuations and has the potential to illuminate environmental perturbations through deep time. Here we systematically quantify the thickness of the anterior brachiopod shell which protects the filtration chamber and is thus considered functionally homologous across higher taxa of brachiopods. Our data come from 66 genera and 10 different orders and shows well-defined upper and lower boundaries of anterior shell thickness. For Ordovician and Silurian brachiopods we find significant order-level differences and a trend of increasing shell thickness with water depth. Modern (Cenozoic) brachiopods, by comparison, fall into the lower half of observed shell thicknesses. Among Ordovician–Silurian brachiopods, older stocks commonly have thicker shells, and thick-shelled taxa contributed more prominently to the Great Ordovician Biodiversification but suffered more severely during the Late Ordovician Mass Extinction. Our data highlight a significant reduction in maximum and minimum shell thickness following the Late Ordovician mass extinction. This points towards stronger selection pressure for energy-efficient shell secretion during times of crisis.     

Variability in the sculpture of the shell aperture of Ruthenica filograna (Rossmassler, 1836) (Gastropoda: Clausiliidae) in specimens from natural populations and from laboratory breeding

An analysis of variability in the sculpture of shell apertureswas carried out for 1,087 specimens of Ruthenica filograna from33 localities and for 20 laboratory-bred individuals rearedfrom parents from a single locality. Nine patterns of aperturesculpture were distinguished. There are usually two kinds ofsculpture: with one plica in the aperture being an extensionof the inferior lamella, and without plicae in the aperture.Sporadically, the subcolumellar lamella may be connected withthe lip of the aperture by a plica. Additional plicae occurin the aperture only rarely. All populations examined showedvariations in the number and arrangement of plicae. The coefficientof variation in the number of plicae in the shell aperture rangedfrom 32.6% to 121.8% within localities. No consistent relationshipwas found between the geographical location and the shell sculpturepattern. Specimens bred in the laboratory developed shell apertureswith a variety of sculptural patterns, including one presentneither in the parents, nor in the population from which itcame. This characteristic is probably phenotypically plastic.It cannot be used for taxonomic purposes, and it is suggestedthat great caution is needed before using similar charactersin other species. (Received 27 March 2007; accepted 4 February 2008)     

Length frequency and morphometric analysis of the non-indigenous red-rimmed melania (<Emphasis Type="Italic">Melanoides tuberculata</Emphasis>) populations in Slovakia

The red-rimmed melania Melanoides tuberculata (O. F. Müller) has been used in the aquarist trade and biocontrol programs, leading to its emergence outside its native range. Two populations of the red-rimmed melania occur in Slovakia. We investigated the morphometric features of mentioned populations and made a comparison. These two populations differ significantly in investigated features. Mean shell length of red-rimmed melania individuals from the Opatovce nad Nitrou was 14.7 mm, while the mean shell length of specimens from Pie??any was 24.4 mm (p?<?10^?6). Mean shell width, aperture length, and aperture width for specimens from the Opatovce nad Nitrou was 4.4 mm, 4.1 mm, and 2.3 mm, respectively. Mean shell width, aperture length, and aperture width of Pie??any specimens was 7.5 mm (p?<?10^?6), 7.1 mm (p?<?10^?6), and 3.8 mm (p?<?10^?6), respectively. The maximum shell length of specimens from the Opatovce nad Nitrou site was 22–23 mm, while shell length of specimens from Pie??any reached 32–33 mm. Regression analysis of the shell length and shell width, aperture length and aperture width, indicate equivalent relationship of the shell patterns for the two populations. Population structure analysis provided first inside of their population status, identifying a proliferating population at the Opatovce nad Nitrou site, while only adult individuals were observed at the site in Pie??any. The red-rimmed malania is a good candidate for possible biological invasions in thermal waters of temperate zone.     

Punctal density in the Ordovician orthide brachiopod Paucicrura rogata: anatomical and paleoenvironmental variation

In the orthide brachiopod Paucicrura rogata , from the Upper Ordovician of New York, the density of punctae (small perforations in the shell) increases from about 350 to about 450 per mm² of shell in the transition from shallow-water limestones to black graptolitic shales down the outer slope of the Taconic Trench. Punctal density is significantly correlated with water depth, as measured by distance downslope along individual volcanic ash layers, and by reciprocal averaging score for macroinvertebrate fossil assemblages ( p < O.01). Because punctal density increases with temperature in living brachiopods, the results suggest that water temperature increased with depth in the Taconic basin, corroborating similar conclusions based on Paucicrura's oxygen-isotopic composition. The increase in punctal density with increasing water depth may also reflect reduced shell growth rates of P. rogata in oxygen-deficient bottom waters, causing a reduction in spacing between punctae. Ontogenetic decrease of punctal density with increasing shell size reflects differential rates of puncta formation at the shell margin. □ Brachiopoda, Orthida, Ordovician, Taconic Basin, paleoceanography, paleoecology, punctae, caeca.     

Structure of the first-formed shell of the Middle Ordovician orthid-like brachiopods from the Leningrad Region

Shell structure of the first-formed shell of the Middle Ordovician orthid-like brachiopods from the Leningrad Region is described. The 190-μm-wide first-formed shell is composed of finely granular layer while 700-μm-wide first-formed shell is fibrous. Thus the order Orthida in the Early Paleozoic included brachiopods with both planktotrophic and lecithotrophic larvae in the ontogeny.     

Biology of the Hyolitha

Hyoliths are Paleozoic fossils that have a calcareous exoskeleton consisting of an elongate, usually bilaterally symmetrical cone, a close fitting operculum, and a pair of curved appendages. Their skeletal ultrastructure resembles the crossed-lamellar shell layers of some molluscs. Several specimens from the Ordovician of France and the Cambrian of Antarctica have parts of the gut preserved by infilling matrix, showing that both mouth ad anus were located near the cone aperture. Muscle scars in other hyolith shells indicate that the animal had a series of dorsoventral and longitudinal, or longitudinal and circular muscles, which operated through a hydrostatic skeleton to protract and retract the head, to open and close the operculum, and to move the appendages. Although the shell form and skeletal ultra-structure of hyoliths are of a molluscan type, the muscle insertions suggest that the hyolith cone is not homologous with the dorsal exoskeleton of primitive molluscs. Hyoliths probably constitute a small extinct branch of phylum size, related to the Mollusca and the Sipunculoidea. All three groups may have had common ancestors in the late Precambrian.     

Shell structure and affinities of the enigmatic Lower Ordovician articulate brachiopod Lycophoria Lahusen, 1886

Wright, A. D. 1992 04 15: Shell structure and affinities of the enigmatic Lower Ordovician articulate brachiopod Lycophoria Lahusen. 1886. Lethaia . Vol. 25, pp. 125–129. Oslo. ISSN 0024–1164.
The enigmatic Baltoscandian Lower Ordovician brachiopod Lycophoriu has a combination of morphological characters that makes it difficult to place taxonomically. The more recent assignments of the genus have been with the Porambonitacea, the Triplesiacea and the Orthacea. A basic character in articulate brachiopods is the differentiation of the secondary shell into either stacked fibres or laminar sheets. The hitherto unknown shell structure in Lycophoria has been examined under the electron microscope and is shown to be fibrous, which is taken as ruling out any close affinity with the lamellar shelled Triplesiacea. Despite superficial similarities, features of the shell interior are not compatible with the pentameride Porambonitacea and although there are differences from the typical orthacean, these are no greater than those of the accepted orthid Producrorrhis. Lycophoria , in the monotypic family Lycophoriidae, is accordingly best regarded as a specialized offshoot of the basic orthacean stock. * Shell microstructure, Lycophoridae, Orrhacea, Porambonitacea, Triplesiacea, Lower Ordovician, Baltoscandia .     

Early growth-stages and classification of orthoceridan Cephalopods of the Darriwillian (Middle Ordovician) of Baltoscandia

Five apices of orthoceridan cephalopods from the early Middle Ordovician Holen Limestone of Öland, Sweden that where collected in the late 19th Century by G. Holm provide information on cephalopod evolution in the early Palaeozoic. The apices belong to specimens of the genus Hedstroemoceras Foerste, 1930 and Archigeisonoceras Chen, 1984. The apices are small in comparison with apices of other cephalopods of the Ordovician; the initial chambers of the shells of both genera are hemispherical and approximately 1 mm and 1.5 mm in cross-section diameter, respectively. The apical 2–3 mm of the shell are free from growth-lines and possess no cicatrix, though distinct longitudinal wrinkles are present. There is a slight variability of siphuncle position during early growth in Archigeisonoceras. It can be shown that the structure of the connecting ring of Hedstroemoceras is similar to that of other Orthocerida. Additionally, the hemispherical apex of Lituites perfectus Wahlenberg, 1821 gives evidence for the orthoceridan affinity of lituitidans. The investigation shows that early Middle Ordovician Orthocerida display a characteristic connecting ring structure, a characteristic apex morphology and variable siphuncular positions that differs significantly from other cephalopods of the Ordovician. Based on this evidence it is concluded that a small spherical apex is an autapomorphy of the Orthocerida. Moreover, this evidence supports a splitting of the order Orthocerida in two taxa of different affinities. The Orthocerida sensu stricto comprises orthocones with a tubular siphuncle nearly without endospiphuncular deposits, and a spherical apex. Embryonic shell, orthoceridan ancestry, orthoceridan classification.     

A Middle Devonian symbiotic relationship involving a gastropod, a trepostomatous bryozoan, and an inferred secondary occupant

Morris, P. J., Linsley, R. M. & Cottrell. J. F. 1991 01 15: A Middle Devonian symbiotic relationship involving a gastropod, a trepostomatous bryozoan. and an inferred sceondary occupant. Lethaia , Vol. 24. pp. 55–67. Oslo. ISSN 0024–1164.
The high-spired gastropod Palaeozygopleuru (Loxonematacea) of the Hamilton Group (Middle Devonian) in New York State is often entrusted by the trepostomatous bryozoan Leptotrypella (Hetero-trypidae). The form of this enerustation leads us to infer a secondary occupant that dwelled in empty shells of Palaeozygopleura while the bryozoan grew upon them. Encrusted specimens usually have an open aperture, thin or no enerustation on the apertural side of the gastropod shell. and thick encrustation on the abapertural side. While the aperture is invariably open. the columellar and parictal lips of the aperture, and the apertural face of the first whorl are usually encrusted. While the gastropod is alive, this area rests upon the dorsal surface of the foot and the remainder of the shell rests upon the substratc. These encrustation patterns suggest that an occupant of the shell. other than the gastropexl. prevented overgrowth of the aperture and oriented the shell aperture-down. This allowed ahapertural growth of the bryozoan. The presence of thin encrustation on the apertural side of the shell is problematical. It requires either the presence of a secondary occupant capable of holding a thinly encrusted shell off the substrate, or the ability of the juvenile bryozoan colony to extend onto portions of the shell that were in contact with the substrate. The presence of a secondary occupant. such as a sipunculan worm. capable of lifting the shell, but usually resting it aperture-down on the substrate seems the most suitable explanation for the nature of the encrustation. Gastropoda. bryozoa. symbiosis. Devoniun. hermit crab, Sipuncula     

Finite element analysis of a double membrane tube (DMS-tube) and its implication for gastropod shell morphology

Molluscan shells, including those of Gastropoda, are formed by accretionary growth at the mantle edge. The mantle is a thin membrane of skirt-like shape, which extends minutely beyond the aperture, and its edge adds a shell increment to the aperture margin so that each increment copies a configuration of the mantle edge at that time. Thus, regulation of shell morphogeny is almost equivalent to the factors which control the mantle form at the moment of shell growth. Form of the mantle skirt is considered to be kept in a state of balance between the force of its internal stress and forces acting on it such as fluid pressure or muscle contraction. The expansion behavior of the mantle skirt has been numerically analyzed by using an elastic model (DMS-tube), which represents the fundamental structure of the mantle tissue as a double membrane structure with internal springs (DMS). Four characteristic expansion patterns of the DMS-tube have been detected: (1) general outward expansion; (2) developing a ridge-like fold on an initial longitudinal protrusion of the tube edge; (3) drastic shift of the expanded state from a uniformly curved to an elliptical shape in outline, owing to the existence of a fixed boundary condition on the tube wall; and (4) constricted protrusion on the open region of the shell wall surrounding the DMS-tube. These results have the potential for answering the following questions relating to the morphogenesis of gastropod shells. How does the mantle skirt usually make contact with the inner surface of the shell wall so as to ensure continuous accretion of shell materials to the aperture margin? What is the cause of spiral ridges? Why do open coiling or minimally overlapping shells have generally circular apertures, while shells with apertures overlapped by whorls have non-uniformly curved apertural lips? What is the cause of long closed spines and why do they always appear on spiral ridges?     

Evolution,palaeoecology, and palaeobiogeography of the Late Ordovician–Early Silurian brachiopod Epitomyonia

Epitomyonia is characterized by various types of dorsal ridges, which may be transverse, longitudinal, or highly convoluted and probably served as skeletal supports for lophophores of various complexity. Multivariate analyses suggest that the Epitomyonia-bearing brachiopod associations lived in relatively shallow-water environment in the Late Ordovician, and inhabited mainly deep-water environments in the early Wenlock. The temporal and spatial change in the faunal distribution may be explained by three alternative scenarios: (1) Epitomyonia followed the broad evolutionary trend of the Palaeozoic Evolutionary Fauna to shift from shallow- to deeper-water settings over time; (2) the dicoelosiid communities could not compete with the large-shelled pentameride communities in continental shelf settings during the Early Silurian; or (3) only the shallow-water Epitomyonia died out in the Late Ordovician mass extinction event, whereas some poorly known deep-water Late Ordovician forms survived into the Early Silurian. Epitomyonia paucitropida n. sp. from the lower Whittaker Formation (late Katian) of the Mackenzie Mountains, northwestern Canada, is reported as the first known Ordovician species of Epitomyonia from the palaeocontinent of Laurentia, characterized by a small shell with weak, transverse dorsal ridges that are most primitive for the genus.     

Decline in diversity of early Palaeozoic loosely coiled gastropod protoconchs

Quantitative data on molluscan larval conch fossil assemblages of ages ranging from the Ordovician (Argentina and the Baltic region), through Silurian (Austria), Devonian (Poland) to Carboniferous (Texas) supplement knowledge of early planktonic gastropods communities transformations. They show that larval shells of the bilaterally symmetrical bellerophontids and dextrally coiled gastropods with a hook-like straight apical portion of the first whorl initially dominated. Their relative frequency, as well as that of the sinistrally coiled ‘paragastropods’, diminished during the Ordovician and Silurian to virtually disappear in the Late Devonian and Early Carboniferous. Already during the Ordovician, diversity of larvae with gently loosely coiled first whorl increased, to be replaced then with more and more tightly coiled forms. Both the aperture constrictions and mortality peaks, probably connected with hatching and metamorphosis, indicate that the Ordovician protoconchs with hook-like first coil represent both the stage of an embryo developing within the egg envelope and a planktonic larva. The similarity of the straight apex to larval conchs of hyoliths and advanced thecosome pteropods is superficial, as these were not homologous stages in early development.     

How to recognize in situ fossil cephalopods: evidence from experiments with modern Nautilus

Field and flume experiments with modern Nautilus pompilius establish two prerequisites to recognize in situ preservation of fossil cephalopod shells (soft parts were within body chamber in situ at the time of fossilization): occurrence of the upper jaw within the body chamber and the position of jaws within the body chamber. Morphology of shells and jaws in modern and fossil nautiloids is so similar that these prerequisites can be applied for fossil nautiloids and provide implications for ammonoids. The upper jaws of Nautilus start to move at a water velocity of > 0.2 m/s, when the shells are reoriented with the aperture downstream; jaws are therefore unlikely to be secondarily deposited near the shell aperture by bottom currents. The lower jaws, moved at the velocity of > 0.1 m/s, can be deposited around the shell aperture by weak current (0.1–0.2 m/s in velocity), but never enter the inside of body chamber. Neither jaw is likely to be separately and selectively displaced from the inside of the body chamber through scavenging of the soft parts by burrowing infaunal animals. An upper jaw preserved inside the body chamber, together with a lower jaw, is thus a reliable indicator of in situ preservation; a sole lower jaw preserved around the shell aperture is likely to be secondarily deposited. Sedimentary structures inferring rapid burial events and jaw size are useful as additional evidence. Smaller jaws were more likely to be displaced from the body chamber by scavenging by infaunal animals after in situ burial, so that smaller jaws preserved within the body chamber suggest less scavenging. These findings are crucial to interpreting the taphonomic history and palaeo-ecology of fossil cephalopods.     

SEXUAL DIMORPHISM OF NUCELLA LAPILLUS (GASTROPODA: MURICIDAE) IN NORTH WALES, UK

Multivariate statistical methods were employed to examine sexualdimorphism in size and shape of Nucella lapillus collected from 16sheltered sites along coasts of Anglesey and the Lleyn Peninsula, NorthWales, UK. Females were significantly larger than males in overallsize; among 12 relative measures of shell shape, two ratios (shellwidth/shell length and aperture length/shell length) were significantlydifferent between males and females, but these differences usuallydecreased with increasing age (shell length). The observed hypoallometricdimorphism could be a result of selection on increased femalefecundity, which may be positively correlated with shell sizein N. lapillus as in other gastropod species. (Received 22 November 1999; accepted 10 April 2000)     

Brachiopod morphology,life mode,and crushing predation in the Ordovician Arnheim Formation,Kentucky, USA

Predation on ancient shelled prey is an often-studied topic in paleoecology, but the early Paleozoic and the brachiopods that dominated the seafloor at that time are relatively underrepresented in the predation literature. We assessed predatory repair scar frequencies among the brachiopod genera from the Early Richmondian (Late Ordovician) Oregonia Member (Arnheim Formation) near Flemingsburg, Kentucky. We found higher repair frequencies on the concavo-convex Rafinesquina and Leptaena relative to the bi-convex genera. There were no trends in repair frequency through the stratigraphic section and no relationships between repair frequency and community diversity metrics. It is possible that concavo-convex brachiopods’ flat shape, thin shell profile, and free-lying (no pedicle attachment) lifestyle made them more likely or appealing targets of Ordovician crushing predators. It is also possible that concavo-convex brachiopods were better suited to survive crushing attacks than biconvex taxa. We also found differences in shell ornament that may influence the visibility of repair scars.     

Shape variation in the rough periwinkle Littorina saxatilis on the west and south coasts of Britain

Variation in the shape of the shell in Littorina saxatilis Olivi has been shown to be due largely to the same variables on both the west and the south coasts of Britain, and it exhibits various clines. Two important aspects are the size of the aperture, which becomes relatively larger from the Isle of Man southwards to Cornwall and eastwards from Devon to the Isle of Wight, and the jugosity of the shell, which increases with distance from Cornwall both northwards as far as the Isle of Man and eastwards as far as Kent. Superimposed on the clines are domains of shape, notably one in Lewis/Harris, where the shells have a relatively large aperture, which is long and narrow, coupled with a rather globose second whorl. The local and geographical aspects of shell shape variation are discussed.     

Infection of the mollusc Littorina saxatilis with parthenites of trematodes and their impact on a shell form: analysis of populations inhabiting the littoral coast of the White Sea

12 rocky shore populations of Littorina saxatilis from three islands of Chupa Inlet (Kandalaksha Bay, White Sea) were examined for infection with trematodes. Morphometric characters (6 indexes of the shell and aperture shape) of molluscs were investigated for all these populations. Exposed and sheltered sites were considered at every island and high and low littoral samples were fulfilled at every site. Seven species of trematodes, Podocotyle atomon, Cryptocotyle lingua, Renicola sp., Himasthla sp., Microphallus piriformes, M. pygmaeus, M. pseudopygmaeus, were found. Uneven distribution of trematodes was confirmed by log-linear analysis. Sheltered populations of L. saxatilis have the greater infection prevalence than exposed ones. This is due to the heavy infection with M. piriformes and M. pygmaeus. The prevalences by these trematodes are up to 52.97% and 27.16% respectively in sheltered populations of the host. The prevalence of M. piriformes tend to be higher at the upper shore level of sheltered sites. In a contrast, the prevalence of M. pygmaeus is significantly higher at the low part of such sites. Factor analysis shows a significant association of the indices of L. saxatilis shell shape with three factors. The first one is associated with the "elongation" of a shell and reveals L. saxatilis from the exposed rocky shore to be more elongated than the molluscs from sheltered sites. The second one is connected with the "aperture shape" index. There is an association of this factor with the shore level position of samples. The third factor reflects the affect of trematodes on the shell shape. The molluscs infected with M. piriformes show "elongated" shell shape and relatively smaller aperture. Shall peculiarities of the hosts infected with M. piriformes and M. pygmaeus are somewhat different. The results of the factor analysis is justified by the series of analysis of variances on the values of shell indices (MANOVA) according to the factors "exposure", "shore level" and "infection".