Phorbas Posidoni N.Sp. (Porifera: Poecilosclerida) from the Aegean Sea, with a Discussion of the Family Anchinoidae

Total Page:16

File Type:pdf, Size:1020Kb

Phorbas Posidoni N.Sp. (Porifera: Poecilosclerida) from the Aegean Sea, with a Discussion of the Family Anchinoidae JOURNAL OF NATURAL HISTORY, 1991, 25, 827-836 Phorbas posidoni n.sp. (Porifera: Poecilosclerida) from the Aegean Sea, with a discussion of the family Anchinoidae ELENI VOULTSIADOU-KOUKOLIRAt and ROB W. M. VAN SOEST~ t Department of Zoology, University of Thessaloniki, 54006 Thessaloniki, Greece Institute of Taxonomic Zoology (Zoologisch Museum), University of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam, The Netherlands (Accepted 8 March 1991) A new species of the genus Phorbas Duchassaing and Michelotti, 1864 (junior synonym Anchinoe Gray, 1867) is described from the Aegean Sea (Eastern Mediterranean). Phorbas posidoni n.sp. is distinguished from the known Mediter- ranean species of Phorbas primarily by its elaborate repent-ramose habit combined with an extensive, spongin-enforced, plumoreticulate skeleton of oxea-like tornotes and the usual echinating acanthostyles and arcuate chelae. As in most Phorbas the surface bears characteristic areolae. The only other elaborate Mediterranean species, P. mercator (Schmidt) differs clearly in spiculation, since it lacks micro- scleres and has strongyles as the main skeletal elements. Phorbas fictitius (Bowerbank), P. tenacior (Topsent) and P. paupertas sensu Topsent are all incrusting and differ in spicule form and sizes as well. Likewise, Phorbas species from the neighbouring East Atlantic each show clear differences with P. posidoni n.sp. The genus Phorbas is discussed and compared with Pronax Gray (1867) sensu Lrvi (1973). It is proposed to unite the two genera because the alleged sharp difference between the two (only oxea-like tornotes in the skeletal tracts in Phorbas, only acanthostyles in the skeletal tracts in Pronax) is compromised by a range of intermediate conditions in various species. KEYWORDS: Systematics, Porifera, Poecilosclerida, Aegean Sea, Eastern Mediterranean. Introduction During recent collecting in the Aegean Sea (Eastern Mediterranean) a species of the family Anchinoidae Topsent, 1928 (=Phorbasidae De Laubenfels, 1936) was dis- covered, which could not be assigned to any of the known species from the Mediterranean (for a list of these, see Pulitzer-Finali, 1983). The new species is unusual because of its elaborate repent-ramose habit, whereas most other Anchinoidae are massively incrusting. In addition to providing a description of the new species, this paper also intends to address the confused taxonomy of the family Anchinoidae, based on a critical survey of the literature and examination of the characters of the type species of most genera assigned to the family, notably Phorbas amaranthus, type of Phorbas Duchassaing and Michelotti (1864), and Pronax plumosa, type of Pronax Gray (1867). Based on the usually employed criteria in Mediterranean sponge taxonomy, see, for example, Topsent (1928), Vacelet (1969), Pulitzer-Finali (1983), the new species 002~2933/91 $5.00 © 1991 Taylor & Francis Ltd. 828 E. Voultsiadou-Koukoura and R. W. M. van Soest conforms to the genus Anchinoe Gray (1867). This genus is defined by most authors as having smooth diactinal spicules in the skeletal tracts and in the ectosomal region, with echinating acanthostyles only in echinating condition. Recently, Anchinoe was synonymized with Phorbas Duchassaing and Michelotti (1864) (Van Soest, 1984, and Discussion below) and this slowly gained wide acceptance (Bergquist, 1961; Van Soest, 1984; Bergquist and Fromont, 1988; Uriz and Rosell, 1989). Accordingly, the new Eastern Mediterranean species is assigned to Phorbas. Systematic description Order POECILOSCLERIDA Topsent, 1928 Family ANCHINOIDAE (Topsent, 1928 (emended) (junior synonym: Phorbasidae De Laubenfels, 1936; Van Soest, 1984; even though the genus Anchinoe is now considered a junior synonym of Phorbas the family group name Anchinoidae is valid according to the ICZN rules). Poecilosclerida with plumose or plumoreticulate skeletal tracts consisting of choanosomal styles and/or smooth diactinal spicules, echinated by styles; ectosomal skeleton consisting of grouped smooth diactinal spicules lying perpendicularly, paratangentially or tangentially to the surface. Microscleres: chelae (if present) are arcuate; sigmata may frequently be absent; toxa and rhaphides are absent. Genus Phorbas Duchassaing and Michelotti, 1864 Synonymy: Pronax Gray, 1867: 536; Anchinoe Gray, 1867; Suberotelites Schmidt, 1868; Plumohalichondria Carter, 1876; ?Clathrissa Von Lendenfeld, 1888; Stylostichon Topsent, 1892; Lissopocillon Ferrer Hernandez, 1916; Merriamium De Laubenfels, 1936. Definition (emended): Anchinoidae in which the styles composing the choanosomal tracts, if present, are heavily spined acanthostyles; echinating styles are likewise heavily spined. Occasionally, the coring acanthostyles are completely replaced by the diactinal smooth spicules. For justification of the synonymy see below. FIG. 1. Holotype of Phorbas posidoni n.sp. Phorbas posidoni n.sp. (Porifera: Poecilosclerida) 829 Phorbas posidoni n.sp. HOLOTYPE,Zoological Museum of Amsterdam ZMA POR 8969, Gulf of Kavala, North Aegean Sea, 17 May 1977. Description. Form: the sponge is a ramose, bushy aggregate, more or less erect to sprawling, about 11 cm long, 9 cm high and 3 cm wide. The branches often anastomose and have a diameter of 0.5 to 1.5 cm and a height up to 5 cm; they are in most cases depressed and terminally thinned (Fig. 1). Colour: brown in formalin. Consistency: soft, elastic and compressible. Surface: it is smooth, with many tangential areolate pore fields. Oscules are numerous, irregularly distributed, up to 1 mm in diameter. The ectosome is fleshy and easily detachable. Skeleton: the ectosomal skeleton is made of abundant isochelae and hastate oxeas. Oxeas are arranged, at right angles to the surface, around areolate pore fields (Fig. 2). This vertical arrangement is not so strict in areas among the pore fields, where oxea-like tornotes can be found even tangentially laid in the ectosome. Isochelae are scattered without any pattern on the dermal membrane. The choanosomal skeleton is composed of well developed spongin fibres forming a more or less well organized network (Fig. 3). Oxea-like tornotes, identical to those found in the ectosome, are embedded in the spongin fibers, which are provided with echinating acanthostyles. No acanthostyles are found inside the fibres, thus having an accessory role in the skeleton. The spongin fibres have a diameter of l0 to 80 #m and they are in most areas totally cored with spicules, although in many of them only two-thirds of their width is cored. The spongin network is equally developed both in the base and near the surface of the sponge. Isochelae are present in the choanosome of the sponge but in lower abundance than in the ectosome. Spicules: megascleres. Diactinal spicules forming the main skeleton, are smooth oxea-like tornotes, measuring 265-310 x 3.5-6 #m. Acanthostyles, usually straight but sometimes slightly curved, covered with spines all along their length (Fig. 4). The spines, most of which are hooked become gradually smaller forward to the apex of the spicules. Acanthostyles measure 110-145 x 7#m. Microscleres: arcuate isochelae measuring 21.5~8.8 #m. Habitat. The sponge was dredged from a depth of about 30m. The substratum was sandy silt mixed with biogenic detritus. FIG. 2. Ectosomal skeleton of Phorbas posidoni n.sp. 830 E. Voultsiadou-Koukoura and R. W. M. van Soest FIG. 3. Choanosomal skeleton of Phorbas posidoni n.sp. F~G. 4. Spicules of Phorbas posidoni n.sp. a, hastate oxea; b, acanthostyle; c, isochelae. Etymology. The species is named after the Greek mythological God of the sea Posidon (Neptune). Discussion Comparison with related species from the Mediterranean The new species differs from the known Mediterranean species (cf. appendix in Pulitzer-Finali, 1983) assignable to Phorbas especially in the elaborate growth form and the development of discrete spongin-enforced spicule tracts forming a reticulation. In the elaborate growth form, it is approached perhaps only by Phorbas mercator Phorbas posidoni n.sp. (Porifera: Poecilosclerida) 831 (Schmidt, 1868 as Suberotelites, see also redescription of Topsent (1938) and Pulitzer- Finali, 1983), and P. dendyi (Ferrer Hernandez, 1916 as Lissopocillon). From the former species the new Phorbas differs clearly in its possession of chelae (absent in P. mercator) and oxea-like tornotes (tylostrongylotes in P. mercator). The latter species is described as a massive sponge, but the photo (Ferrer Hernandez, 1916, Fig. 9) shows short projections and elevations. It is described in a new genus on account of the occurrence of 'pocillos', modifications of the arcuate chelae not worthy of generic separation. A further difference with our new species is the possession of two distinct sizes of acanthostyles in P. dendyi sensu Ferrer Hernandez. Note that if the synonymization of Phorbas and Pronax/Stylostichon is accepted, the combination of Phorbas dendyi would be preoccupied by that of Topsent (1892). The other species of 'Anchinoe', namely A. tenacior Topsent 1925, A. paupertas sensu Topsent (1925) and A. fictitius (Bowerbank 1866), differ in being thinly or massively incrusting. Moreover, of these only A.fictitius is similar in spiculation to our new species, differing, however, in possessing the usual two size categories of acanthostyles. Both A. tenacior and A. paupertas have smooth strongyles instead of oxea-like tornotes. The Mediterranean species assigned to 'Pronax', namely P. dives Topsent (1891), P. flbulatum Topsent (1893) and P. lieberkuehni Burton (1930), all differ
Recommended publications
  • Proposal for a Revised Classification of the Demospongiae (Porifera) Christine Morrow1 and Paco Cárdenas2,3*
    Morrow and Cárdenas Frontiers in Zoology (2015) 12:7 DOI 10.1186/s12983-015-0099-8 DEBATE Open Access Proposal for a revised classification of the Demospongiae (Porifera) Christine Morrow1 and Paco Cárdenas2,3* Abstract Background: Demospongiae is the largest sponge class including 81% of all living sponges with nearly 7,000 species worldwide. Systema Porifera (2002) was the result of a large international collaboration to update the Demospongiae higher taxa classification, essentially based on morphological data. Since then, an increasing number of molecular phylogenetic studies have considerably shaken this taxonomic framework, with numerous polyphyletic groups revealed or confirmed and new clades discovered. And yet, despite a few taxonomical changes, the overall framework of the Systema Porifera classification still stands and is used as it is by the scientific community. This has led to a widening phylogeny/classification gap which creates biases and inconsistencies for the many end-users of this classification and ultimately impedes our understanding of today’s marine ecosystems and evolutionary processes. In an attempt to bridge this phylogeny/classification gap, we propose to officially revise the higher taxa Demospongiae classification. Discussion: We propose a revision of the Demospongiae higher taxa classification, essentially based on molecular data of the last ten years. We recommend the use of three subclasses: Verongimorpha, Keratosa and Heteroscleromorpha. We retain seven (Agelasida, Chondrosiida, Dendroceratida, Dictyoceratida, Haplosclerida, Poecilosclerida, Verongiida) of the 13 orders from Systema Porifera. We recommend the abandonment of five order names (Hadromerida, Halichondrida, Halisarcida, lithistids, Verticillitida) and resurrect or upgrade six order names (Axinellida, Merliida, Spongillida, Sphaerocladina, Suberitida, Tetractinellida). Finally, we create seven new orders (Bubarida, Desmacellida, Polymastiida, Scopalinida, Clionaida, Tethyida, Trachycladida).
    [Show full text]
  • Is There a Mediterranean Cold-Water Coral Sponge Fauna?
    fmars-08-662899 June 9, 2021 Time: 17:47 # 1 ORIGINAL RESEARCH published: 15 June 2021 doi: 10.3389/fmars.2021.662899 Mediterranean Coral Provinces as a Sponge Diversity Reservoir: Is There a Mediterranean Cold-Water Coral Sponge Fauna? Andreu Santín1*, Jordi Grinyó1,2, Maria Jesús Uriz3, Claudio Lo Iacono1, Josep Maria Gili1 and Pere Puig1 1 Institut de Ciències del Mar (ICM-CSIC), Barcelona, Spain, 2 Department of Ocean Systems Sciences, NIOZ Royal Netherlands Institute for Sea Research and Utrecht University, Den Burg, Netherlands, 3 Centre d’Estudis Avançats de Blanes (CEAB-CSIC), Blanes, Spain Cold-water coral reefs (CWC) are known to be biodiversity hotspots, however, the sponge assemblages found to dwell within these habitats haven not been studied in depth to date in the Mediterranean Sea. The present article provides the first insight Edited by: on the associated sponge fauna of the recently discovered CWC communities on the Shirley A. Pomponi, Florida Atlantic University, Catalan Margin and, to a lesser extent, the Cabliers Coral Mound Province, while also United States reviewing the current knowledge of the sponge fauna dwelling in all the Mediterranean Reviewed by: CWC provinces. In regards to the studied areas, some rare species are cited for Sergi Taboada, Autonomous University of Madrid, the first time in the Mediterranean or redescribed, while two of them, Hamacantha Spain (Hamacantha) hortae sp. nov. and Spongosorites cabliersi sp. nov. are new to science. Giorgio Bavestrello, At a basin scale, Mediterranean CWC appear as poriferan biodiversity hotspots, yet University of Genoa, Italy current diversity values on each site rather represent a small fraction of its actual *Correspondence: Andreu Santín fauna.
    [Show full text]
  • Svenzea Zeai, a Caribbean Reef Sponge with a Giant Larva, And
    Svenzea zeai, a Caribbean Reef Sponge with a Giant Larva, and Scopalina ruetzleri: A Comparative Fine-Structural Approach to Classification (Demospongiae, Halichondrida, Dictyonellidae) Author(s): Klaus Rützler, Rob W. M. van Soest, Belinda Alvarez Source: Invertebrate Biology, Vol. 122, No. 3 (Summer, 2003), pp. 203-222 Published by: Blackwell Publishing on behalf of American Microscopical Society Stable URL: http://www.jstor.org/stable/3227199 Accessed: 27/10/2010 19:43 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=black. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Microscopical Society and Blackwell Publishing are collaborating with JSTOR to digitize, preserve and extend access to Invertebrate Biology.
    [Show full text]
  • Demospongiae, Poecilosclerida), New Names for the Mediterranean ‘Phorbas Paupertas’ and ‘Phorbas Coriaceus’
    Zootaxa 1873: 26–38 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition) Phorbas topsenti and Phorbas tailliezi (Demospongiae, Poecilosclerida), new names for the Mediterranean ‘Phorbas paupertas’ and ‘Phorbas coriaceus’ JEAN VACELET & THIERRY PEREZ Centre d’Océanologie de Marseille, Aix-Marseille Université, CNRS UMR 6540 DIMAR, Station Marine d’Endoume, rue Batterie des Lions, 13007 Marseille, France. E-mail: [email protected] Abstract A common Mediterranean species of Phorbas has been frequently identified under the specific name paupertas, refer- ring to the sponge described as Hymeniacidon paupertas by Bowerbank 1866 from the north coasts of Europe. It is actu- ally different from this species, and a new name, Phorbas topsenti sp.nov., is proposed. A description of the morphology, spicule complement and cytology is given. Another Mediterranean species of Phorbas, Phorbas tailliezi sp. nov. is described. Key words: Taxonomy, Porifera, Hymedesmiidae, Mediterranean, Phorbas topsenti, new species Introduction A common, remarkable Mediterranean sponge has been frequently identified under the specific name pauper- tas in the genera Anchinoe, Hymedesmia or Phorbas after Topsent (1934, 1936). As pointed out by van Soest et al. (2000), this thickly encrusting sponge, of a vivid red color, is actually different from the species described as Hymeniacidon paupertas by Bowerbank (1866: 223), at present classified in Hymedesmia. Here we re-describe the morphology of the Mediterranean specimens and erect a separate new species to harbor them. We also illustrate the ultrastructure of its cell types and of its bacterial symbionts, which could be useful for species distinction, and the differences with Phorbas fictitius (Bowerbank, 1866) and Hamigera hamigera (Schmidt, 1862), which are very similar by field characters.
    [Show full text]
  • New and Little-Known Poecilosclerid Sponges from the Mexican
    Zoological Studies 51(7): 1139-1153 (2012) New and Little-Known Poecilosclerid Sponges from the Mexican Pacific Ocean Jose Maria Aguilar-Camacho* and Jose Luis Carballo Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México (Estación Mazatlán), Avenida Joel Montes Camarena s/n, PO Box 811 Mazatlan C.P., 82000, Mexico (Accepted August 3, 2012) Jose Maria Aguilar-Camacho and Jose Luis Carballo (2012) New and little-known poecilosclerid sponges from the Mexican Pacific Ocean. Zoological Studies 51(7): 1139-1153. Five species belonging to the order Poecilosclerida are described based on material from the Mexican Pacific Ocean. Two of them are little known from this region: Biemna rhadia de Laubenfels 1930 and Discorhabdella urizae Maldonado, Carmona, van Soest and Pomponi 2001. Three species are new to science: Phorbas reginae sp. nov., a black massive sponge with 2 categories of acanthostyles, tornotes, and arcuate isochelae; Ectyonopsis sigmata sp. nov., a red encrusting sponge with acanthostyles, acanthostrongyles, ectosomal tornotes, anchorate isochelae, and sigmas; and Myxillodoryx nicolae gen. nov., sp. nov., a pinkish-red, cushion-shaped sponge with ectosomal tylotes, choano- somal acanthostyles, arcuate tridentate isochelae, unguiferate multidentate isochelae, and sigmas. The new genus Myxillodoryx gen. nov. is characterized by having 2 types of isochelae: arcuate and unguiferate multidentate. Based on the former classification of the order Poecilosclerida, we allocated this genus to the family Coelosphaeridae even if this species has 1 diagnostic feature of the family Myxillidae (unguiferate multidentate chelae). We emended the genus Ectyonopsis Carter 1883a to include species bearing anchorate or unguiferate isochelae and sigmas as microscleres.
    [Show full text]
  • Which Signaling Mechanisms Sponge Larvae Use to Become Adult Sponges? Ilya Borisenko, Olga Podgornaya, Alexander Ereskovsky
    From traveler to homebody: Which signaling mechanisms sponge larvae use to become adult sponges? Ilya Borisenko, Olga Podgornaya, Alexander Ereskovsky To cite this version: Ilya Borisenko, Olga Podgornaya, Alexander Ereskovsky. From traveler to homebody: Which sig- naling mechanisms sponge larvae use to become adult sponges?. Advances in Protein Chemistry and Structural Biology, 116, pp.421-449, 2019, 10.1016/bs.apcsb.2019.02.002. hal-02072762 HAL Id: hal-02072762 https://hal.archives-ouvertes.fr/hal-02072762 Submitted on 19 Mar 2019 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. From traveler to homebody: Which signaling mechanisms sponge larvae use to become adult sponges? Ilya Borisenkoa, ⁎, Olga I. Podgornayaa, b, c, Alexander V. Ereskovskya, d a St. Petersburg State University, St. Petersburg, Russia b Far Eastern Federal University, Vladivostok, Russia c Institute of Cytology RAS, St. Petersburg, Russia PROOF d Mediterranean Institute of Marine and Terrestrial Biodiversity and Ecology (IMBE), Aix Marseille University, CNRS, IRD, Avignon Université, Marseille, France ⁎ Corresponding author: Email address: [email protected] (I. Borisenko) Contents 1. Introduction 2 2. Embryonic development 3 3. Morphology of the metamorphosis 6 4. The fate of ciliated cells during metamorphosis of H.
    [Show full text]
  • Nudibranch Food
    List of the Worldwide Food Habits of Nudibranchs by Gary McDonald The "list of the worldwide food habits of nudibranchs" was originally available on the web as a supplement to two articles published in the Veliger: McDonald & Nybakken, 1997 & 1999. On 5 November 2014, it was brought to my attention that the list is no longer available on the web, so I have uploaded it to eScholarship in order to make it available once again. References MCDONALD, G. R., & J. W. NYBAKKEN. 1997. A worldwide review of the food of nudibranch mollusks. I. Introduction and the suborder Arminacea. Veliger 40(2):157-159. MCDONALD, G. R., & J. W. NYBAKKEN. 1999. A worldwide review of the food of nudibranch mollusks. II. The suborder Dendronotacea. Veliger 42(1):62-66. Nudibranch Food A List of the Worldwide Food Habits of Nudibranchs by Gary R. McDonald Long Marine Laboratory, 100 Shaffer Rd., Santa Cruz, Cal. 95060 [email protected] and James W. Nybakken Moss Landing Marine Laboratories, P.O. Box 450, Moss Landing, Cal. 95039 [email protected] This list is an attempt to compile all of the published food data for nudibranchs. More than 4,600 nudibranch papers were perused in an effort to find as much of the published food data as possible. More than 1,000 papers mentioned nudibranch food. Included are all of the articles that we were able to obtain prior to June 1996. The list is arranged phylogenetically so that trends in food preferences within various groups of nudibranchs may be seen more easily. The food items are listed alphabetically below each species of nudibranch.
    [Show full text]
  • New Data on Embryonic Development of Halisarca Dujardini Johnston, 1842 (Demospongiae, Halisarcida)
    New data on embryonic development of Halisarca dujardini Johnston, 1842 (Demospongiae, Halisarcida) Alexander V. ERESKOVSKY Elisaveta L. GONOBOBLEVA Department of Embryology, Biological Faculty, St. Petersburg State University Universitetskaja nab. 7/9, St. Petersburg 199034 (Russia) [email protected] Ereskovsky A. V. & Gonobobleva E. L. 2000. — New data on embryonic development of Halisarca dujardini Johnston, 1842 (Demospongiae, Halisarcida). Zoosystema 22 (2) : 355- 368. ABSTRACT Embryogenesis of Halisarca dujardini from the White Sea (Russia) was stu- died by light and transmission electron microscopy. The cleavage is equal and asynchronous. From the stage of 8-16 blastomeres, the cells divide perpendi- cularly to the surface of embryos and form a coeloblastula. The few internal KEY WORDS Embryonic development, cells of the prelarva are derived by multipolar migration of external cells. The Halisarca dujardini, larva (disphaerula) consists of two flagellated spheres: external and internal. ultrastructure, cleavage, The internal sphere is formed by invagination of lateral flagellated cells. The blastula, disphaerula is completely flagellated, but the flagella on posterior pole are morphogenesis, sparse. The peculiarities of Halisarca embryonic development indicates the larvae, comparative embryology, particular position of this genus in subclass Ceractinomorpha. The pattern of Porifera. development is most primitive in this subclass. ZOOSYSTEMA • 2000 • 22 (2) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.mnhn.fr/publication/ 355 Ereskovsky A. V. & Gonobobleva E. L. RÉSUMÉ Données nouvelles sur l’embryogenèse d’Halisarca dujardini Johnston, 1842 (Demospongiae, Halisarcida). L’embryogenèse d’Halisarca dujardini de la mer Blanche (Russie) a été étudiée en microscopie photonique et électronique. La segmentation est égale et asyn- chrone.
    [Show full text]