Redescription of Hexaconularia He and Yang, 1986 (Lower Cambrian, South China): implications for the affinities of conulariid-like small shelly fossils

Abstract:  Hexaconularia , a Lower Cambrian small shelly fossil (SSF) that has been allied with conulariids and scyphozoan cnidarians, is redescribed and refigured. A salient feature of this monospecific genus is the presence of distinct apical and abapical regions. The apical region probably represents an embryonic shell that apparently lacked a basal attachment structure. Comparisons of this feature with the apical end of the smallest known conulariids and with conulariids terminating in an apical wall (schott) reveal substantial differences in structure and ornamentation. Differences in apical anatomy between conulariids and Arthrochites , possibly the nearest SSF relative of Hexaconularia , are also apparent. Comparisons of Hexaconularia with Punctatus , an SSF taxon showing distinct apical and abapical regions in both posthatching specimens and prehatching embryos, suggest that the early development of Hexaconularia was direct. These results have important implications for hypotheses of a conulariid/scyphozoan affinity for Hexaconularia and its possible SSF relatives, and they suggest that Hexaconularia -bearing strata may yield prehatching embryos of this genus.     

Haldane's rule in Carabus: interspecific mating between Carabus punctatoauratus and Carabus splendens using experimental tests and molecular markers

Carabus splendens Olivier and Carabus punctatoauratus Germar (Coleoptera: Carabidae: Carabini) are ground beetles found in sympatry in forests in the eastern Pyrénées. We investigated potential interspecific mating using crossing experiments under controlled laboratory conditions and paternity analyses based on microsatellite markers. We show that under laboratory conditions, interspecific mating produces viable and fertile offspring. Although hybrid males were sterile, females were fertile and thus reproductive isolation between the two species is far from complete. We discuss the results in relation to species integrity due to pre- or post-zygotic barriers, and compare them to a previous study quantifying hybridization between the two species in natural populations.     

New data on the morphology of Sphenothallus Hall: implications for its affinities

Van Iten, H., Cox, R. S. & Mapes, R. H. 1992 04 15: New data on the morphology of Sphenothallus Hall: implications for its affinities. Lethaia , Vol. 25, pp. 135–144. Oslo. ISSN 0024–1164.
Recent speculation on the phylogenetic relationships of Sphenothallus Hall, 1847 has focused on two alternatives: (1) affinity with hydrozoan or scyphozoan cnidarians, or (2) affinity with annelids or other 'worms'. We have found that some species of Sphenothallus formed branching clonal colonies, and that others produced a thin transverse wall, similar to the conulariid schott. Sphenothallus tests are composed of carbonate apatite and are built of numerous, extremely thin lamellae (< 1 μm) that parallel the surface of the test. The holdfast, long interpreted as consisting of a pair of nested cups, actually consists of a single closed, broadly conical expansion floored by a thin basal membrane. Many thecate hydrozoan and scyphozoan polyps form branching clonal colonies and produce a thin transverse wall, similar to that produced by Sphenothallus . Further, thecae of coronatid scyphozoans are built of submicron-thick lamellae that parallel the outer surface of the theca, and coronatid thecae possess a closed, broadly conical apical expansion that serves as a holdfast. No such combination of characteristics exists among annelids or other non-cnidarian taxa. The recent discovery of paired tentacles in Sphenofhallus from the Early Devonian Hunsrück Slate by Fauchald et al ., thought to indicate that Sphenorhallus was wormlike, does not militate against a hypothesis of affinity with cnidarians. We therefore favour the hypothesis of a cnidarian affinity for Sphenothallus. * Sphenothallus, vermiform problematica, phylogenetic inference, coloniality, Cnidoria, Hydrozoa, Scyphozoa, Annelida, conulariids, byroniids, Bear Gulch Limestone .     

Exceptionally preserved conulariids and an edrioasteroid from the Hunsrück Slate (Lower Devonian,SW Germany)

Abstract: Nineteen partial specimens of Conularia sp., together with an articulated agelacrinitid edrioasteroid and several discinid brachiopods, occur in close association with a probable biological substrate on a small slab of silty Hunsrück Slate (Lower Devonian, Emsian) from Bundenbach, Germany. Most of the conulariids occur in V‐like pairs or in a single cluster of 12 specimens arranged in a fan‐like radial pattern. Together with the edrioasteroid and (possibly) brachiopods, the conulariids probably were attached to the substrate in life and then were buried and possibly killed by a single influx of silty mud. The apertural end of many of the conulariids is partially covered by inwardly folded short lappets, which may have closed in response to rapid (but gentle) burial. Rock matrix in the apertural region of the peridermal cavity of nearly all of the conulariids exhibits irregular, variably dense concentrations of pyrite. The concentrations occur almost exclusively within the conulariids, where they probably formed as a result of the decay of retracted conulariid soft parts. Although the concentrations lack clearly defined anatomical features that can be unambiguously homologized with particular anatomical structures of any extant taxon, their form and distribution within the conulariids are consistent with the hypothesis that conulariids were polypoid scyphozoans.     

Calculating census efficiency for river birds: a case study with the White-throated Dipper Cinclus cinclus in the Pyrénées

Using the binomial law we modelled field data to estimate the probability ( ̂ ) of detecting pairs of breeding White-throated Dippers, and the population size ( ̂ ± confidence limits). The model was divided into two parts according to whether the actual size of the population under study was known or not; in the latter case the truncated binomial model was used. Dipper abundance data were collected from three 4-km-long river tracts in the Pyrénées (France) during the breeding seasons of different years. Goodness-of-fit tests indicated that the binomial model fitted the data well. For a given visit during the survey, the estimated probability of detecting any pair of Dippers if they were present was always high (0.63–0.94) and constant from year to year but not between sites. Estimations ( ̂ ) of the size of the population provided by the binomial model were very close to that derived from mapping techniques. This study provides the first ever quantification of the number of visits required to detect birds on linear territories: three visits were necessary to detect the whole breeding population.     

A hybrid zone between Chorthippus parallelus parallelus and Chorthippus parallelus erythropus (Orthoptera: Acrididae): morphological and electrophoretic characters

Two subspecies of the grasshopper Chorthippus parullelus have distributions that abut in the Pyrenees. We have studied morphological and electrophoretic variation in II populations near Mont Louis, Pyrénées-Orientales, France. The two subspecies differ in several morphological characters and at one enzyme locus, esterase-2, and in this area they hybridize forming a cline in morphology less than 5 km wide. Examination of museum specimens suggests that this hybrid zone extends for most of the length of the Pyrenees, possibly breaking down in the extreme west. As the two subspecies are known to differ in mating behaviour this hybrid zone is well suited to the study of reinforcement.     

Insights into the Skeletonization,Lifestyle, and Affinity of the Unusual Ediacaran Fossil Corumbella

The Ediacaran fossil Corumbella is important because it is hypothesized to be a scyphozoan cnidarian, and thus might be one of the rare examples of bona fide Neoproterozoic animals. Unfortunately, its mode of life, style of skeletonization, and taxonomic affinity have been very controversial. Here, we use X-ray micro-CT, SEM, and taphonomic analysis to compare preservational modes of Corumbella, in order to better understand the symmetry, mode of construction, preservational style, and taxonomy of this group. Results suggest that articulated and disarticulated specimens of Corumbella from the Ediacaran of Brazil, Paraguay, and the United States, although sometimes preserved very differently, represent the same taxon—Corumbella werneri. Corumbellids had a thick but flexible theca and probably lived with their basalmost part anchored in the sediment, much like Conotubus. When considered together, these results suggest that Corumbella was one of the first animals to build a skeleton, employing a lamellar microfabric similar to conulariids.     

Conical Thecae of Precambrian Macroorganisms

On the basis of new extensive collections, made by the present authors, a group of unsegmented three-dimensional fossil remains from the Late Precambrian of Namibia and the southeastern White Sea area, including such genera as Protechiurus, Vendoglossa, and Vendoconularia, were restudied. It is established that the fossils belong to two morphologically close genera (Protechiurus and Vendoconularia), composing the new family Protechiuridae. It is possible that they represent the same genus or even species, but the incompleteness of the preservation of the Namibian member of the group prevents their integration. The fossils initially represented conical hollow elastic objects, hexagonal in the cross-section, which were open at the wide end and closed at the pointed end. The sculpture of their surface consisted of longitudinal ridges, both single and double, bounding the rows of the short transverse ridges, scalloped along one margin. Vendoconularia had wide plates which were attached to the cone outwardly along the longitudinal unpaired ridges. In general morphology and several details the fossils resemble the thecae of conulariids and anabaritids, known from Late Precambrian and Paleozoic and classified as the scyphozoan cnidarians. The main difference lays in the theca mineralization, which consists of phosphate in conulariids, carbonate in anabaritids, and is absent in the case of protechiurids. It is proposed that the protechiurids, which appeared in the fossil record first, may belong to the basal group of ancient scyphozoans, ancestral to the conulariids on the one hand, and to the anabaritids on the other.     

A hybrid zone between Chorthippus parallelus parallelus and Chorthippus parallelus erythropus (Orthoptera: Acrididae): behavioural characters

Two subspecies of the grasshopper Chorthippus parallelus have ranges that abut in the Pyrenees. Males produce two types of song—calling song and courtship song. We have examined the use and structure of these songs in males from nine sites near Mont-Louis, Pyrénées-Onentales, France. These sites were previously used to identify the position of the hybrid zone on morphological characters. The subspecies differ in the use and structure of both types of song. Clines in these characters correspond in position with the morphological cline. At least one character may be relevant to mate choice but none of the characters show evidence of reinforcement.     

Asymmetry in an Ordovician conulariid cnidarian

Sendino, C., Zágor?ek, K. & Taylor, P.D. 2012: Asymmetry in an Ordovician conulariid cnidarian. Lethaia, Vol. 45, pp. 423–431. Conulariids are fossils of the presumed polyps of an extinct scyphozoan cnidarian group. Their cone‐shaped skeletons normally show perfect tetraradial symmetry. However, in the Ordovician species Metaconularia anomala ( Barrande 1867 ) from Drabov (Czech Republic), tetraradial symmetry is compromised in three ways: (1) the skeleton often shows torsion; (2) the four sides may vary in width at the same level within one individual; and (3) one side may be deleted to give a triradial skeleton. Almost 2000 specimens were studied in museum collections. About 56% of analysed specimens are twisted in an anticlockwise direction (sinistral) when viewed from the apex towards the aperture, 28% show no torsion, 1% exhibit clockwise torsion (dextral) and the remaining 15% cannot be classified. Maximum measured torsion rate was 1.5°/mm. A significant negative correlation between torsion rate and length suggests that more highly torted individuals may have survived less well. Almost 5% of individuals show loss of one side for at least part of their lengths. Although many individuals have four sides of equal width, in a significant proportion the sides are of unequal width, up to a maximum/minimum side width ratio of 2 (i.e. widest face twice the width of the narrowest). In the absence of a satisfactory taphonomic model to explain the asymmetries, they are regarded as mirroring asymmetries in the living conulariids, with the strong preference for sinistral torsion interpreted as an example of a fixed asymmetry that was genetically controlled and heritable. It is speculated that the signalling protein Nodal as well as Hox‐like genes were involved in controlling the asymmetries described in M. anomala.     

Genetic diversity of dog breeds: within-breed diversity comparing genealogical and molecular data

The genetic diversity of 61 dog breeds raised in France was investigated. Genealogical analyses were performed on the pedigree file of the French kennel club. A total of 1514 dogs were also genotyped using 21 microsatellite markers. For animals born from 2001 to 2005, the average coefficient of inbreeding ranged from 0.2% to 8.8% and the effective number of ancestors ranged from 9 to 209, according to the breed. The mean value of heterozygosity was 0.62 over all breeds (range 0.37–0.77). At the breed level, few correlations were found between genealogical and molecular parameters. Kinship coefficients and individual similarity estimators were, however, significantly correlated, with the best mean correlation being found for the Lynch & Ritland estimator ( r  =   0.43). According to both approaches, it was concluded that special efforts should be made to maintain diversity for three breeds, namely the Berger des Pyrénées, Braque Saint-Germain and Bull Terrier.     

Teratology and pathology of some Paleozoic conulariids

Conulariids exhibiting various pathologies and teratological conditions have been examined from Paleozoic rocks of North America, South America, Europe, and Africa. Published examples of teratological conditions in conulariids have been reviewed. To these cases we add a specimen of Paraconularia missouriensis (Swallow) from the Mississippian of Ohio which possesses six faces. The supposed three-sided conulariid species Conularina triangulata (Raymond) is based upon a specimen which is not a conulariid. This genus is removed from the phylum Conulariida and is considered incertae sedis. Pathologies include injuries to the exoskeleton which are grouped into patterns termed scalloped, cleft, and embayed. Scalloped injuries represent minor chipping at the aperture of the conulariid exoskeleton, and may have occurred accidentally or through predation. Cleft and embayed injuries, found only on post-Silurian taxa, indicate that conulariids suffered severe sublethal attacks more frequently after the rise of several types of durophagous predators in the middle Paleozoic. Some middle and late Paleozoic conulariid species strengthened the exoskeleton, perhaps to resist predation. Regeneration of injured integument or rods has occurred in conulariids exhibiting damaged exoskeletons.     

Embryos,polyps and medusae of the Early Cambrian scyphozoan Olivooides

The Early Cambrian organism Olivooides is known from both embryonic and post-embryonic stages and, consequently, it has the potential to yield vital insights into developmental evolution at the time that animal body plans were established. However, this potential can only be realized if the phylogenetic relationships of Olivooides can be constrained. The affinities of Olivooides have proved controversial because of the lack of knowledge of the internal anatomy and the limited range of developmental stages known. Here, we describe rare embryonic specimens in which internal anatomical features are preserved. We also present a fuller sequence of fossilized developmental stages of Olivooides, including associated specimens that we interpret as budding ephyrae (juvenile medusae), all of which display a clear pentaradial symmetry. Within the framework of a cnidarian interpretation, the new data serve to pinpoint the phylogenetic position of Olivooides to the scyphozoan stem group. Hypotheses about scalidophoran or echinoderm affinities of Olivooides can be rejected.     

A hydrogeomorphological context for ecological research on alpine glacial rivers

1. This paper presents a framework for the analysis of glacial stream ecosystems based upon the hierarchy of physical processes that operate over timescales varying from diel to millennial. Linked conceptual models propose interrelationships between climate, hydrology and fluvial geomorphology in alpine catchments.
2. These conceptual models are illustrated using data from the Taillon/Gabiétous catchment, French Pyrénées. Secondary sources provide information concerning the long-term sequence of climatic and geomorphological controls on contemporary catchment processes. Detailed hydrogeomorphological field data collected over three consecutive summer melt seasons (1995–97) permit identification of marked changes in shorter-term (diel, seasonal and inter-annual) physical processes.
3. Clear differences in the response of water quality and quantity variables were observed between years as climatic conditions varied. In two of the three study years, a precipitation-driven regime was imposed upon the typical ablation-driven river discharge pattern in alpine streams. Clear changes in water quality and quantity were evident with increasing distance from the glacier: (i) discharge increased although specific discharge decreased markedly, (ii) the mean and variability in water temperature increased, and (iii) base concentrations of suspended solids decreased.
4. The physical processes incorporated in the conceptual models presented in this paper have ecological implications because they underpin a nested suite of disturbance regimes operating over timescales from diel to millennial.     

CLADISTIC ANALYSIS OF THE SUBORDER CONULARIINA MILLER AND GURLEY, 1896 (CNIDARIA, SCYPHOZOA; VENDIAN–TRIASSIC)

Abstract: Results of a cladistic analysis of the suborder Conulariina Miller and Gurley, 1896, a major extinct (Vendian–Triassic) group of scyphozoan cnidarians, are presented. The analysis sought to test whether the three conulariid subfamilies (Conulariinae Walcott, 1886 , Paraconulariinae Sinclair, 1952 and Ctenoconulariinae Sinclair, 1952 ) recognized in the Treatise on Invertebrate Paleontology (TIP) are monophyletic. A total of 17 morphological characters were scored for 16 ingroup taxa, namely the genera Archaeoconularia, Baccaconularia, Climacoconus, Conularia, Conulariella, Conularina, Ctenoconularia, Eoconularia, Glyptoconularia, Metaconularia, Notoconularia, Paraconularia, Pseudoconularia, Reticulaconularia, Teresconularia and Vendoconularia. The extant medusozoan taxa Cubozoa, Stauromedusae, Coronatae and Semaeostomeae served as outgroups. Unweighted analysisof the data matrix yielded 1057 trees, and successive weighting analysis resulted in one of the 1057 original trees. The ingroup is monophyletic with two autapomorphies: (1) the quadrate geometry of the oral region; and (2) the presence of a mineralized (phosphatic) periderm. Within the ingroup, the clade (Vendoconularia, Teresconularia, Conularina, Eoconularia) is supported by the sinusoidal longitudinal geometry of the transverse ridges, and the much larger clade (Baccaconularia, Glyptoconularia, Metaconularia, Pseudoconularia, Conularia, Ctenoconularia, Archaeoconularia, Notoconularia, Climacoconus, Paraconularia, Reticulaconularia) is supported by the presence of external tubercles, which, however, were lost in the clade (Notoconularia, Climacoconus, Paraconularia, Reticulaconularia). As proposed by Van Iten et al. (2000) , the clade (Notoconularia, Climacoconus, Paraconularia, Reticulaconularia) is supported by the termination and alternation of the transverse ribs in the corner sulcus. The previously recognized subfamilies Conulariinae, Paraconulariinae and Ctenoconulariinae were not recovered from this analysis. The diagnostic features of Conulariinae (continuation of the transverse ornament across the corner sulcus and lack of carinae) and Ctenoconulariinae (presence of carinae) are symplesiomorphic or homoplastic, and Paraconulariinae is polyphyletic. The families Conulariellidae Kiderlen, 1937 and Conulariopsidae Sugiyama, 1942 , also recognized in the TIP, are monogeneric, and since they provide no additional phylogenetic information, should be abandoned.     

The origin of tetraradial symmetry in cnidarians

Serially arranged sets of eight septa‐like structures occur in the basal part of phosphatic tubes of Sphenothallus from the early Ordovician (early Floian) Fenxiang Formation in Hubei Province of China. They are similar in shape, location and number, to cusps in chitinous tubes of extant coronate scyphozoan polyps, which supports the widely accepted cnidarian affinity of this problematic fossil. However, unlike the recent Medusozoa, the tubes of Sphenothallus are flattened at later stages of development, showing biradial symmetry. Moreover, the septa (cusps) in Sphenothallus are obliquely arranged, which introduces a bilateral component to the tube symmetry. This makes Sphenothallus similar to the Early Cambrian Paiutitubulites, having similar septa but with even more apparent bilateral disposition. Biradial symmetry also characterizes the Early Cambrian tubular fossil Hexaconularia, showing a similarity to the conulariids. However, instead of being strictly tetraradial like conulariids, Hexaconularia shows hexaradial symmetry superimposed on the biradial one. A conulariid with a smooth test showing signs of the ‘origami’ plicated closure of the aperture found in the Fenxiang Formation supports the idea that tetraradial symmetry of conulariids resulted from geometrical constrains connected with this kind of closure. Its minute basal attachment surface makes it likely that the holdfasts characterizing Sphenothallus and advanced conulariids are secondary features. This concurs with the lack of any such holdfast in the earliest Cambrian Torellella, as well as in the possibly related Olivooides and Quadrapyrgites. Bilaterally arranged internal structures in polyps representing probably the oldest medusozoans support the suggestions based on developmental evidence that the ancestor of cnidarians also was a bilaterally symmetrical animal. This is one more example of fossil data that strictly fit the molecular phylogenetic evidence but not necessarily morphology‐based zoological interpretations.     

Anatomy and phylogenetic significance of Eoconularia loculata, a conulariid from the Silurian of Gotland

The type material of Eoconularia loculata (Wiman, 1895), a conulariid with high, bifurcated septa, was originally found in an erratic boulder, hence the source-rock is as yet unidentified. Recently, a rich material of the species has been discovered at eleven localities in the Silurian Hemse Beds of Gotland Another two localities in the Slite Beds (Wenlockan, Sheinwoodian) revealed what is assumed to be the ancestor of E loculata. This ancestral form probably constitutes a separate taxon distinguished from E. loculata in having simple, unbranched septa. E. loculata is re-described, and four ontogenetic stages in septal development are recognized. During stage 1, the most juvenile stage, the septa are simple. The septa in stages 3 and 4, the adult stages, are coarse and bifurcated. The affinities of conulariids are discussed, with the conclusion that the group shares a number of similarities with modem scyphomans. The microstructures of the exoskeleton show several similarities with coronatids, and the septa are interpreted to be homologous with the internal structures of stauromedusids. The stratigraphical distribution of all currently known septate conulariids suggests that septa were a primitive morphologic feature ranging from early? Ordovician to late Silurian. The simplest type, however, persisted at least into the early Permian. Five of the eleven described septa types have been found only among the conulariids from Gotland. □Conulariida, EOCONULARIA LOCULATA, bifurcated septa, taxonomy, ecology, morphology, Scyphozoa, Sweden, Gotland, Silurian.     

Additional observations on the gross morphology and microstructure of Baccaconularia Hughes, Gunderson et Weedon, 2000, a Cambrian (Furongian) conulariid from the north-central USA

Baccaconularia Hughes, Gunderson et Weedon, 2000, from the Furongian Series (Cambrian System) of the north-central USA, has been interpreted as a conulariid cnidarian, based on a suite of gross morphological similarities shared only with other post-Cambrian genera currently assigned to this group. Closely spaced, squarish to subrectangular facial nodes of Baccaconularia are aligned in distinct longitudinal files. Nodes also display a subtler, more or less rectilinear transverse alignment, though this pattern commonly is disrupted by offset parallel to the longitudinal files. In their shape and pattern of arrangement, the nodes of Baccaconularia are most similar to the squarish to elongate nodes of Pseudoconularia Bouček, 1939. Longitudinal node files of Baccaconularia may also be compared with the longitudinal facial ridges of Conularia cambria Walcott, 1890 from the Furongian of Wisconsin. Apical angles of Baccaconularia range from approximately 13° to 14.5°. Scanning electron imaging of B. cf. robinsoni shows that its thin, phosphatic skeleton is finely lamellar, with the thickness of individual lamellae measuring approximately 1 μm. The skeleton also exhibits microscopic circular pores and crater-like pits that range from approximately 5 to 10 μm in diameter. These pores and pits are similar in size, geometry, areal density and pattern of arrangement to those of many post-Cambrian conulariids. Microscopic circular pores are documented here for the first time in the genus Archaeoconularia Bouček, 1939 from the Upper Ordovician of the Czech Republic. Although the origin of the pores and pits is open to alternative interpretations, the discovery of these features and fine lamination in Baccaconularia strengthens the argument that this genus is a Cambrian conulariid.     

Unstable B-chromosomes in a European population of Allium schoenoprasutn L. (Liliaceae)

In Allium schoenoprasum , small, dichromatic B-chromosomes occur in 74% of plants at Lac de Tracens, Pare National des Pyrénées. B-chromosomes from Lac de Tracens were compared with the previously well characterized Bs from British populations of A. schoenoprasum at the River Wye, Powys, and were found to differ in terms of both their morphology and the level of mitotic instability. At Lac de Tracens, two morphological types of B-chromosome were observed: a telocentric of about 1.15 μm in length and a metacentric of about 1.35 μm in length; the most frequent B-type in the River Wye populations (B^-1) was not found. B-chromosomes from the River Wye are essentially stable at root tip mitosis, whereas 25 out of 29 plants from Lac de Tracens (86%) showed between-cell variation in B-number, with non-modal numbers in 330 out of 794 cells (41.6%). Most of the variability occurred within individual roots, rather than between roots of single bulbs or of sister bulbs. No meiotic pairing was observed between the different Bs in hybrids from the geographically isolated populations, suggesting a lack of homology. An independent origin of B-chromosomes in the British and French populations is suggested.