Des Miliolacea aux Alveolinacea (grands foraminifères): la faune du Crétacé supérieur et du Tertiaire inférieur du Chiapas (S.E. Mexique)

Found in Chiapas (South-East of Mexico), Praealveolina michaudi nov. sp. and Chubbina jamaicensis (of Campanian-Maestrichtian age) are described, both evolved from Nummoloculina sp. while Raadshoovenia guatemalensis (upper Paleocene-lower Eocene) arises from Quinqueloculina. We precisely analyze and modelize the nepionic coiling during the evolutionary change from Miliolacea to Alveolinacea. During ontogenesis as well as during phylogenesis, quinqueloculine coiling progresses to streptospiral and then to planispiral chamber arrangement with the increase of volume of embryo and of chambers. Appearance of these large foraminifers are understood as a consequence of the rise of K-strategies in their mode of life, following the development of photosynthetic symbioses.     

Anglesites gen. nov. (Ammonoidea, Ancyloceratina), un nouveau genre d’ammonites hétéromorphes du Barrémien supérieur du Sud-Est de la France

The revision of the Crioceras puzosianum d’Orbigny, 1842 made during the revision of the Paléontologie Française of d’Orbigny, shows that this taxon belongs to a new genus: Anglesites gen. nov. This new genus, from upper Barremian age, is monospecific for the moment and is homeomorphic to the Leptoceratoides from the Lower Barremian. It is temporarily included in the subfamily of the Leptoceratoidinae. A neotype for the “Crioceras” puzosianum d’Orbigny, 1842 is herein designated.     

Les premiers Anguilliformes : I. Révision des genres cénomaniens Anguillavus HAY, 1903 et Luenchelys nov. gen.

New specimens from Cenomanian outcrops of Lebanon permit redescribtion of the first eel-like fossil fishes. Anguillavus bathshebaeHay, 1903 becomes synonymous with the type species A. quadripinnisHay, 1903. A new fossil eel with pelvic fins, Anguillavus mazeni nov. sp., is furthermore proposed. In the same way, the revision of Luenchelys minimus nov. gen., nov. sp., reveals some morphological variations at the specific level and a widespread biogeographical repartition which potentially can be important. This whole anatomical revision is a necessary basis for integrating these fossils into the Anguilliformes (Teleostei, Elopomorpha) and gives an idea of the plesiomorphic structural pattern of the order, in which the modern forms are particularly specialised.     

Phylloceratoidea du Jurassique moyen et supérieur du Nord-Est de l’Iran (Monts Binalud)

Phylloceratidae are often the most frequent taxa within the Middle and Late Jurassic ammonite faunas of North Iran but so far they have not been noticed much within the existing literature and have been described only sporadically. In this study, the Phylloceratidae of north Iran are treated systematically and described comprehensively. These include 16 taxa, most of them being previously unpublished for Iran.     

Pseudocitrobacter gen. nov., a novel genus of the Enterobacteriaceae with two new species Pseudocitrobacter faecalis sp. nov., and Pseudocitrobacter anthropi sp. nov,isolated from fecal samples from hospitalized patients in Pakistan

Four isolates of Gram-negative facultatively anaerobic bacteria, three of them producing NDM-1 carbapenemase, were isolated from hospitalized patients and outpatients attending two military hospitals in Rawalpindi, Pakistan, and studied for their taxonomic position. Initially the strains were phenotypically identified as Citrobacter species. Comparative analysis of 16S rRNA gene sequences then showed that the four strains shared >97%, but in no case >98.3%, 16S rRNA gene sequence similarities to members of the genera Citrobacter, Kluyvera, Pantoea, Enterobacter and Raoultella, but always formed a separate cluster in respective phylogenetic trees. Based on multilocus sequence analysis (MLSA) including partial recN, rpoA, thdF and rpoB gene sequence and respective amino acid sequence analysis it turned out that the strains also here always formed separate clusters. Based on further comparative analyses including DNA–DNA hybridizations, genomic fingerprint analysis using rep- and RAPD-PCRs and physiological tests, it is proposed to classify these four strains into the novel genus Pseudocitrobacter gen. nov. with a new species Pseudocitrobacter faecalis sp. nov. with strain 25 CIT^T (= CCM 8479^T = LMG 27751^T) and Pseudocitrobacter anthropi sp. nov. with strain C138^T (= CCM 8478^T = LMG 27750^T), as the type strains, respectively.     

Halorubellus salinus gen. nov., sp. nov. and Halorubellus litoreus sp. nov., novel halophilic archaea isolated from a marine solar saltern

Two extremely halophilic archaeal strains GX3(T) and GX26(T) were isolated from the Gangxi marine solar saltern near the Weihai city of Shandong Province, China. Cells from the two strains were pleomorphic and stained Gram-negative, colonies were red-pigmented. Strains GX3(T) and GX26(T) were able to grow at 25-50 °C (optimum 37 °C), at 1.4-5.1M NaCl (optimum 3.1M), at pH 5.5-9.5 (optimum pH 7.0) and neither strain required Mg(2+) for growth. Cells lyse in distilled water and the minimal NaCl concentration to prevent cell-lysis was 8% (w/v). The major polar lipids of the two strains were PA (phosphatidic acid), PG (phosphatidylglycerol), PGP-Me (phosphatidylglycerol phosphate methyl ester) and three major glycolipids (GL1, GL2 & GL3) chromatographically identical to S-TGD-1 (sulfated galactosyl mannosy glucosyl diether), S-DGD-1 (sulfated mannosyl glucosyl diether), and DGD-1 (mannosyl glucosyl diether) respectively, an unidentified lipid (GL4) was also detected in strain GX26(T). Phylogenetic analysis based on 16S rRNA gene revealed that strain GX3(T) and strain GX26(T) formed a distinct clade with the closest relative, Haladaptatus paucihalophilus (89.9-92.4% and 90.4-92.7, respectively). The rpoB' gene similarities between strains GX3(T) and GX26(T), and between the two strains and the closest relative, Halorussus rarus TBN4(T) are 96.5%, 84.3% and 83.9%, respectively. The DNA G+C contents of strain GX3(T) and strain GX26(T) are 67.3 mol% and 67.2 mol%, respectively. The DNA-DNA hybridization value between strain GX3(T) and strain GX26(T) was 44%. The phenotypic, chemotaxonomic and phylogenetic properties suggest that strain GX3(T) and strain GX26(T) represent two novel species in a new genus within the family Halobacteriaceae, Halorubellus salinus gen. nov., sp. nov. (type strain GX3(T)=CGMCC 1.10384(T)=JCM 17115(T)) and Halorubellus litoreus sp. nov. (type strain GX26(T)=CGMCC 1.10386(T)=JCM 17117(T)).     

Description de Artareites landii nov. (Ammonoidea) du Barrémien supérieur de Majastre (Sud-Est de la France) et discussion sur les Helicancylidae Hyatt, 1894

In the Majastre area (Alpes-de-Haute-Provence, South-Eastern France), the hemipelagic domain of the Lower Cretaceous is exposed to the favour of the “Décrochement du Poil”. This palaeogeographical and paleotectonic position allowed the harvest of a diversified ammonite-fauna in the level with “small Barremites”. It is precisely dated here for the first time, and takes place between the base of the Barremense subzone and the base the Breistrofferi biohorizon of the Vandenheckei biozone (base of the Upper Barremian). Within this fauna, Artareites landii nov. sp. is considered here as a possible descendant of the genus Dissimilites Sarkar due to its characteristics and stratigraphic position. The genus Artareites nov. gen. is certainly an important milestone of the Helicancylidae towards the Toxoceratoides genus morphology. In this case, in agreement with the morphological, ontogenical, stratigraphical and paleobiogeographical parameters, some North American species could be considered as potential intermediate. Due to their characteristics and their joint origin, genus of small size included in the Helicancylidae represent a coherent set with the rank of a family, and is independent of the Ancyloceratidae. Its content is revised and limited to the genus phyletically associated around the main axis of Acrioceras-Dissimilites-Artareites nov. gen. and Toxoceratoides-Helicancylus, whose elements are connected by a number of forms. In this case, Monodites nov. gen. is created for the Hauterivian and Lower Barremian very specific forms Monodites crevolai (Vermeulen et al.) and Monodites trumpyi (Kakabadze et Thieuloy) which does not seem to be integrated in the taxa already recognized in the literature. The family Monoditidae nov. fam. is proposed to mark this uniqueness.     

Studies of the Liagoraceae (rhodophyta) of Western Australia: Gloiotrichus fractalis gen. et sp. nov. and Ganonema helminthaxis sp. nov.

Two new taxa of Liagoraceae (Nemaliales) are described from Western Australia. Gloiotrichus fractalis gen. et sp. nov. has been collected from 3–20 m depths at the Houtman Abrolhos, Western Australia. Plants are calcified, extremely lubricous, and grow to 17 cm in length. Carpogonial branches are straight, 6 or 7 cells in length, arise from the basal or lower cells of cortical fascicles, and are occasionally compound. Branched sterile filaments of narrow elongate cells arise on the lower cells of the carpogonial branch prior to gonimoblast initiation, at first on the basal cells, then on progressively more distal cells. Following presumed fertilisation the carpogonium divides transversely, with both cells giving rise to gonimoblast filaments. The distal cells of the carpogonial branch then begin to fuse, with fusion progressing proximally until most of the cells of the carpogonial branch are included. As fusion extends, the filaments on the carpogonial branch are reduced to the basal 2 or 3 cells. The gonimoblast is compact and bears terminal carposporangia. Spermatangial clusters arise on subterminal cells of the cortex, eventually displacing the terminal cells. The sequence of pre- and post-fertilisation events occurring in the new genus separates it from all others included in the Liagoraceae, although it appears to have close affinities with the uncalcified genus Nemalion. Ganonema helminthaxis sp. nov. was collected from 12 m depths at Rottnest Island, Western Australia. Plants are uncalcified and mucilaginous, the axes consisting of a few (< 10) primary medullary filaments, each cell of which gives rise to a cortical fascicle at alternate forks of the pseudodichotomies borne on successive medullary cells. Subsidiary (adventitious) filaments and rhizoids comprise the bulk of the thallus. Carpogonial branches are straight, (3-)4(-6) cells in length, arise on the basal 1–4 cells of the cortical fascicles, and are frequently compound. Carposporophytes develop from the upper of two daughter cells formed by a transverse division of the fertilised carpogonium. Ascending and descending sterile filaments girdle the carpogonial branch cells and arise mostly on the supporting cell prior to fertilisation. Ganonema helminthaxis is the first completely non-calcified member of the genus, and its reproductive and vegetative morphology supports the recognition of Ganonema as a genus independent from Liagora. Liagora codii Womersley is a southern Australian species displaying features of Ganonema, to which it is transferred.     

Proposal of Ensifer psoraleae sp. nov., Ensifer sesbaniae sp. nov., Ensifer morelense comb. nov. and Ensifer americanum comb. nov

In a survey of rhizobia associated with the native legumes in Yunnan Province, China, seven and nine strains isolated from the root nodules of Psoralea corylifolia, Sesbania cannabina and Medicago lupulina were respectively classified into the novel genomic species groups I and II in the genus Ensifer (former Sinorhizobium) based on the sequence analyses of the 16S rRNA gene. Analyses of concatenated housekeeping genes (atpD, recA and glnII) further revealed that they were distinct lineages in the genus, and group I was most similar to Ensifer terangae and Ensifer garamanticus (both with 94.2% similarity), while group II was most similar to Ensifer adhaerens (94.0%). These groups could be distinguished from closely related species by DNA–DNA relatedness, MALID-TOF MS, cellular fatty acid profiles and a series of phenotypic characters. Therefore, two novel species were proposed: Ensifer psoraleae sp. nov. (seven strains, type strain CCBAU 65732^T = LMG 26835^T = HAMBI 3286^T) and Ensifer sesbaniae sp. nov. (nine strains, type strain CCBAU 65729^T = LMG 26833^T = HAMBI 3287^T). They had a DNA G + C mol% (T_m) of 58.9 and 60.4, respectively. Both of the type strains formed effective nodules on common bean (Phaseolus vulgaris) and their hosts of origin. In addition, the previously described species Sinorhizobium morelense and Sinorhizobium americanum were renamed as Ensifer morelense comb. nov. and Ensifer americanum comb. nov. according to the accumulated data from different studies.     

Sur l'origine du genre Microceratina Swanson, 1980 (Ostracoda, Eucytherurinae) et sa présence téthysienne dès le Jurassique supérieur

The discovery in the uppermost Jurassic of Lebanon of a species of the ostracod genus Microceratina Swanson, 1980 (Eucytherurinae), of which the oldest known species was until now of Maastrichtian age, confirms the Tethyan origin of this genus. Two new species are created: Microceratina bhannesensis nov. sp. from the Upper Jurassic (Kimmeridgian) of Lebanon, and Microceratina azazoulensis nov. sp. from the Cenomanian of Morocco. The genera Chapmanicytherura Weaver, 1982 and Erratacytheridea Herrig et al., 1997, are considered as probable junior synonyms of Microceratina. An exhaustive bibliographical analysis allowed to identifying several other Cretaceous species susceptible to be assigned to the genus Microceratina.     

<Emphasis Type="Italic">Aliihoeflea aestuarii</Emphasis> gen. nov., sp. nov., a novel bacterium isolated from tidal flat sediment

A novel Gram-negative and rod-shaped bacterium, designated N8^T, was isolated from tidal flat sediment. Phylogenetic analysis based on 16S rRNA gene sequences showed that N8^T strain is associated with the family Phyllobacteriaceae: two uncultured clones (98.4 and 99.8% 16S rRNA gene sequence similarity) and the genus Mesorhizobium (≤97.0%). The novel strain formed a separate clade with uncultured clones in the phylogenetic tree based on 16S rRNA gene sequences. Cellular fatty acid profiles predominately comprised C_18:1 ω7c and C_19:0 cyclo ω8c. The major isoprenoid quinone is ubiquinone-10 and genomic DNA G+C content is 53.4 mol%. The polyphasic taxonomic study indicates that the novel strain N8^T represents a novel species of the new genus in the family Phyllobacteriaceae, named Aliihoeflea aestuarii. The type strain is N8^T (= KCTC 22052^T= JCM 15118^T= DSM 19536^T).     

Hironoia fusiformis gen. et sp. nov.; a cornalean fruit from the Kamikitaba locality (Upper Cretaceous,Lower Coniacian) in northeastern Japan

The application of sieving techniques to bulk samples from the Ashizawa Formation, Futaba Group (Lower Coniacian) of northeastern Honshu, Japan, has yielded well-preserved mesofossil assemblages comparable with those recently described from eastern North America, Europe, and central Asia. Among the most abundant and distinctive components of these assemblages are fusiform fruits that are assigned here to a new genus and species, Hironoia fusiformis gen. et sp. nov. The fruits developed from an epigynous ovary with three to four locules. Each locule bears one seed and has a distinctive dorsal germination valve. These features of the fruit, along with the adnate calyx, indicate an affinity to extant Cornales and specifically the Cornaceae sensu lato. The recognition of an unequivocal cornalean fruit in the Early Coniacian–Early Santonian of Japan provides the earliest record of this group in the fossil record. It also establishes a minimum age for the early divergence of the asterid clade, a major group of living angiosperms comprising more than a third of all species of extant flowering plants. Electronic Publication     

Diaceratherium massiliae nov. sp. des argiles oligocènes de Saint-André et Saint-Henri à Marseille et de Les Milles près d’Aix-en-Provence (SE de la France), premier grand Rhinocerotidae brachypode européen

A revision of all previously collected mammalian fossils from the two Late Oligocene sites of Saint-André and Saint-Henri in Marseille (both from the MP 26 reference-level) allows us to identify three Rhinocerotoidea species: Protaceratherium albigense, Ronzotherium romani, Diaceratherium massiliae nov. sp., and maybe a fourth one, Eggysodon cf. gaudryi. Only the first two were previously known there. D. massiliae nov. sp. is found together with R. romani; it is the first case of sympatry ever known between the two genera. D. massiliae nov. sp. is then the most ancient Diaceratherium in Europe, where the genus was previously unknown before the MP 29 reference-level. It is a very large species whose limb bones proportions foreshadow these of the later species of the genus, especially D. lemanense from the Late Oligocene-Early Miocene. This suggests a possible phylogenetic link between D. massiliae nov. sp. and D. lemanense, and the coexistence of at least two different but partially contemporaneous lineages among the European Diaceratherium. In Les Milles near Aix-en-Provence, also from the MP 26 reference-level, the three species P. albigense, R. romani and D. massiliae nov. sp. were also found.     

Heterorhabditidoides chongmingensis gen. nov., sp. nov. (Rhabditida: Rhabditidae), a novel member of the entomopathogenic nematodes

During a recent soil sample survey in Eastern China, a new entomopathogenic nematode species, collected from the Chongming Islands in the southern-eastern area of Shanghai, was discovered. Morphological characteristics of different developmental stages of the nematode combined with molecular data showed that this nematode is a new genus of Rhabditidae, and described as Heterorhabditidoides chongmingensis gen. nov., sp. nov., for that it shares more morphological characteristics with heterorhabditids than with steinernematids. For males, the papillae formula of bursa is 1, 2, 3, 3, with constant papillae number in the terminal group, stoma tubular-shaped and about 1.5 head width; cheilorhabdions cuticularized, esophageal collar present and long, median bulb present. For infective juveniles, EP = 90 (80-105) μm, ES = 104 (92-120) μm, tail length = 111 (89-159) μm, and a = 19.1 (15-21). The percentages of the nucleotides A, T, C and G in the ITS1 regions of the new species are significantly different from those of heterorhabditids and other rhabditids. Molecular phylogenetic trees based on 18S rDNA and the internal transcribed spacer (ITS) sequences data revealed that the new entomopathogenic nematode species forms a monophyletic group, which is a sister group of the clade comprised of some genera of Rhabditidae.     

Nouvelles données sur le genre Imerites Rouchadzé, 1933 (Ammonitida,Ancyloceratina) ; réponses et précisions apportées au travail de Bert,Delanoy et Canut, 2009

This work is mainly an answer to the work of Bert, Delanoy and Canut, on the genus Imerites Rouchadzé, and we add also some points to the knowledge of this genus. An answer is given for the implication of one of the authors (J.V.) of this present work, about the origin of the genus Imerites Rouchadzé, and some significant bibliographic references, not named in the 2009 work, are specified and placed in their context. The characteristics of the ornamental stage “Pseudoshasticrioceras” mainly defined by an oval and compressed section of the whorls and by regular ventrolateral clavi, well marked, which frame the venter in a typical way (Bert et al., p. 181), don’t allow to affirm the presence of this stage during the ontogeny of all species of the genus Imerites Rouchadzé, contrary to what Bert, Delanoy and Canut (p. 30, 31) wrote. The type-species of the genus Imerites Rouchadzé is made clear by the application of the rules of the I.C.Z.N. code. The article 23, with the paragraph 23.1 of the I.C.Z.N. code, imply the senior synonymy of Imerites cristatus (Orbigny) on Imerites giraudi (Kilian), even if the Kilian's species is still the type species of the genus Imerites Rouchadzé. The taxinomical validity of the species Imerites cristatus (Orbigny) is confirmed. Its morphological variability is clarified by the distinction of four referential morphotypes, Cristatus, Giraudi, Favrei and Raricostatum. Consequently, the assumption of a shape-dimensional dimorphism into the populations of the genus Imerites Rouchadzé is rejected, awaiting irrefutable proof. A new species, Imerites stephaniae sp. nov., is described. It was collected in the Gerhardtia sartousiana Zone, in the uppermost part of the Gerhardtia provincialis Subzone, and it is a probable ancestor of the cogeneric species of the Imerites giraudi Zone.     

La phalange distale du pouce d’Orrorin tugenensis (Miocène supérieur du Kenya)

The lion's share of articles dealing with the thumb anatomy of Plio-Pleistocene hominids has focussed on the capacity to manipulate and manufacture tools, and has largely neglected the locomotor aspects. However, in these hominids, the forelimb was still employed in locomotion. Certain of the anatomical characters classically associated with manipulation and/or fabrication of tools are already present in the Late Miocene species Orrorin tugenensis as shown by the terminal thumb phalanx BAR 1901'01. This specimen reveals crucial information suggesting that thumb morphology is not exclusively related to such tool using and manufacturing activities but reflects in a frequently bipedal creature a deeper adaptation concerning the precision grip essential for climbing and balancing, different from that of apes.     

Cristatopecten nov. gen. benoisti (Pectinidae, Bivalvia) du Chattien d'Aquitaine (France). Révision systématique et réflexions sur l'origine et l'évolution des Cristatopecten

The detailed study of a sample of “Pecten (Amussiopecten) benoisti” Cossmann and Peyrot (1914) from the Chattian outcrops of the Adour Basin, allowed to improve our knowledge of this species poorly known till now and to show its important ornamental variability. It follows that some nomenclatural points have been clarified at the generic level: we refute the attribution of this species to the genus Amussiopecten Sacco (1897) as used by authors—genus which was besides emended several times subsequently— and we propose its allocation to Cristatopecten nov. gen. Furthermore, this study allowed to remove the ambiguity existing originally as for the attribution of the species benoisti and burdigalensis to a same genus: our comparison of the latter two taxa shows that if benoisti has to be henceforth included in the new genus Cristatopecten (as its type species), burdigalensis (which is the type species of Amussiopecten) must be incorporated in the genus Flabellipecten Sacco (1897). The two genera Amussiopecten and Flabellipecten are therefore synonymous, as Depéret and Roman had already stated in 1910. The Cristatopecten from the Old World seem to be derived from a group of “Aequipecten” notably known in the Italian Rupelian, the group of “Aequipecten deletus” (Michelotti, 1861), of which we study here a sample from the Aquitaine Chattian. In the Aquitaine Basin, the Cristatopecten are represented, in the Chattian, by Cristatopecten benoisti (Cossmann and Peyrot, 1914); in the Lower Aquitanian, by C. praehaueri (Bongrain, Cahuzac and Freneix, 1994); in the Burdigalian-Langhian of the Saubrigues paleocanyon, by C. haueri (Michelotti, 1847) and C. miogallicus (Cossmann and Peyrot, 1914). During the whole Neogene, they spread and evolved in the Tethys and the Paratethys areas, and seem to have become extinct in the Pliocene with the disappearance of C. cristatus (Bronn, 1827), a species that we consider not as an Amusium s.s., but as the last representative of the Cristatopecten phylum.     

Évolution des environnements littoraux du bassin de Melilla-Nador (Nord-Est Maroc) au Messinien supérieur entre 6,0 et 5,77 Ma

This study of late Messinian deposits (E2 = TCC) from Melilla-Nador carbonate complex (Morocco) allowed us to characterize the environmental evolution between 6.0 and 5.77 Ma, which is a period considered as contemporaneous of the onset and development of the “messinian salinity crisis”. A statistical approach coupled together with classical petrographic study allows us to constrain the sedimentary dynamic of platforms and to propose deposition environment models. The Melilla-Nador platforms E2 deposits show a progressive continuous evolution of littoral environments in two major stages of available space creation and infill. The first stage corresponds to the build up of a reef complex between 6.0 and 5.85 Ma then the second stage, between 5.85 and 5.77 Ma, corresponds to the development of a mixed oolitic shallow ramp during the progressive infill of Melilla-Nador basin. The comparison with the Sorbas basin (Spain) TCC deposits exhibits a very close evolution of late Messinian littoral environments, which could be used as a correlation tool.     

Ormieresia carcini gen. n., sp. n., Microsporidie du Crabe Méditerranéan, Carcinus mediterraneus Czerniavsky, 1884: Cycle Évolutif et Étude Ultrastructurale

RÉSUMÉ. Une nouvelle microsporidie, Ormieresia carcini gen. n., sp. n., est trouvée dans Carcinus mediterraneus Czerniavsky, 1884. Son évolution et ses ultrastructures sont étudiees.
Le cycle débute par un méronte limité par une membrane unitaire et possédant un diplocaryon. Toute la sporogonie se déroule dans la musculature de l'höte. Les stades sporogoniaux sont enveloppés dans une membrane pansporoblastique persistante. Dans chaque pansporoblaste, la sporoblastogenése (donnant naissance à 8 sporoblastes) est particulière; un plasmode sporogonial octonucléé manque. Au cours des divisions schizogoniques et sporogoniques, chaque centre cinétique est formé de 2 plaques, une incluse dans l'enveloppe nucléaire, l'autre occupant une position extranucléaire. Les sporontes en division et les sporoblastes sécrètent des substances métaboliques (granules, tubes) qui sont déposées dans l'espace pansporoblastique.
La spore uninucléée est longue et cylindrique (19,1 × 2,4 μm). Un manubrium rectiligne traverse la spore. Son extrémité postérieure se rétrécit brusquement en un filament polaire décrlvant 4 ou 5 tours de spire. L'extrémité antérieure du manubrium est attachée au capuchon polaire lui-même comprimé par un double anneau. La partie antérieure du manubrium est entourée par un polaroplaste formé d'une zone spongieuse et d'une zone lamellaire.