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Étude Morphologique Et Histologique Des Scutes Des Macroramphosidae (Teleostei, Gastérostéiformes)

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Étude Morphologique Et Histologique Des Scutes Des Macroramphosidae (Teleostei, Gastérostéiformes) Cah. Biol. Mar. (1996) 37 : 189-203 Étude morphologique et histologique des scutes des Macroramphosidae (Teleostei, Gastérostéiformes) Olivier FEDRIGO *, François J. MEUNIER *, **, Guy DUHAMEL * ET Jean-Yves SIRE ** * Laboratoire d'Ichtyologie générale et appliquée, Muséum National d'Histoire Naturelle, Paris (France) ** Equipe de Recherche "Formations squelettiques", URA CNRS 1137, Laboratoire d'Anatomie comparée, Université D. Diderot, Paris (France) Correspondance : F.J. Meunier, Laboratoire d'Ichtyologie générale et appliquée, Muséum National d'Histoire Naturelle, 43, rue Cuvier, 75231 Paris Cedex 05 (France) Fax : (33) 1 40 79 37 71 - E-mail : [email protected] Résumé : Les scutes des Macroramphosidae (Macroramphosus, Notopogon et Centriscops) sont de petite taille (0,1 à 1,5 mm) et composées d'une base osseuse aplatie à partir de laquelle s'élève une hampe surmontée, pour le premier genre, d'une couronne carénée dont la marge postérieure est ornementée d'une ou plusieurs pointes. La morphologie des scutes (étudiée au MEB) varie selon leur position sur le corps, surtout chez Notopogon, et leurs caractéristiques, notamment l'ornementation, sont propres à chaque genre, avec des variations interspécifiques au sein du genre, d'où une utilisation pos- sible en systématique. L'orientation antéro-postérieure des ornementations (carènes) suggère qu'elles pourraient avoir un rôle hydrodynamique comparable à ce qui a été décrit pour les écailles placoïdes des Chondrichthyens. La structure et la minéralisation des scutes, étudiées plus en détail chez M. scolopax et TV. xenosoma, sont identiques ; elles sont entièrement constituées d'os à fibres parallèles (pseudo-lamellaire) et la minéralisation y est homogène. L'os est avasculaire et acellu- laire, les ostéoblastes n'étant pas incorporés dans la matrice qui, par ailleurs, présente des lignes d'arrêt de croissance très nettes. Cette organisation structurale est totalement différente de celle des écailles élasmoïdes. Abstract : Morphological and histological study of scutes in Macroramphosidae (Teleostei, Gasterosteiformes). In the three genera of Macroramphosidae, Macroramphosus, Notopogon and Centriscops, the scutes are small (0.1 to 1.5 mm) and composed of a flat basal plate from the middle of which a shaft arises. In Macroramphosus this shaft is covered by a ridged crown, the posterior margin of which is ornamented with one or several spines. The morphology of the scutes (SEM study) varies according to their location on the body, particularly in Notopogon, and the characteristics of the ornamentations are different from one genus to another. Moreover, as these ornamentations also show interspecific variations, they could be used for systematic purposes. The antero-posterior orientation of the superficial ridges led us to suggest that they have a hydrodynamical function similar to that described for the placoid scales in Chondrichthyans. A detailed, structural and ultrastructural (TEM) study of the scutes in M. scolopax and N. xenosoma has shown a similar tissue composition. They are constituted entirely of parallel-fibered bone which is homogeneously mineralized. This bone is avascular and acellular, and its matrix shows clear lines of arrested growth. This structural organization is completely different from that of elasmoid scales. Keywords : Macroramphosidae, scute, morphology, histology, systematics, phylogeny. Introduction Reçu le 17 avril 1996 ; accepté après révision le 11 octobre 1996. Les Ostéichthyens possèdent une très grande diversité de Received 17 April 1996; accepted in revised form 11 October 1996. structures dermiques tégumentaires (divers types d'écailles, 190 LES SCUTES DES MACRORAMPHOSIDAE scutes, plaques dermiques...) dont les relations phylétiques N. lilliei Regan, 1914 et N. macrosolen Barnard, 1925) sont mal connues. Cette diversité se traduit à la fois par une d'une épine sur la troisième plaque épaxiale inférieure. multitude Je formes et d'organisations et par une grande Divers aspects de la systématique, de la biogéographie et de variété te tissus. C'est afin d'établir un catalogue le plus la biologie des Macroramphosidae ont été développés complet possible de cette diversité tissulaire dans le der- récemment par l'un d'entre nous (Duhamel, 1995). moquelette des Ostéichthyens actuels, et ainsi pouvoir Le présent travail comprend une étude morphologique metre en évidence les homologies de leurs constituants, que (microscopie électronique à balayage = MEB) et une étude nous avons réalisé depuis quelques années une série de tra- histologique fondée sur des observations obtenues à partir vaux comparatifs sur divers éléments du dermosquelette de de lames minces (microscopie photonique et microradiogra- la lignée actinoptérygienne : écailles ganoïdes de polyptères phie) et de coupes semi-fines et ultrafines (microscope élec- (Meunier, 1980 ; Sire et ai, 1987) ou de lépisostées (Sire, tronique à transmission = MET) des scutes de quelques 1994), scutes de Siluriformes (Sire et Meunier, 1993) et espèces de cette famille. écailles élasmoïdes dans divers groupes de Téléostéens : Ostéoglossiformes (Meunier, 1984a), Cyprinidae Matériel et méthodes (Zylberberg et Meunier, 1981), Characiformes (Meunier, Matériel 1996), Cichlidae (Sire et Meunier, 1981). Nous nous L'étude des scutes a été réalisée à partir d'un ensemble de sommes intéressés également à la structure des plaques der- spécimens appartenant aux trois genres de miques de Téléostéens plus évolués, comme les Scombridae Macroramphosidae : Macroramphosus scolopax, Centris- Thunninae (Meunier et Sire, 1981) et les Ostraciidae cops humerosus (Richardson, 1848) pour les deux genres (Meunier et Francillon-Vieillot, 1995) et, très récemment, à monospécifiques et Notopogon xenosoma, N. lilliei, celle des plaques dermiques de quelques Gastérostéi- N. armatus, pour le troisième. formes : Gasterosteidae, Syngnathidae et Macrorham- La morphologie et l'histologie des scutes ont été plus phosidae. Ce sont les scutes de cette dernière famille qui particulièrement étudiées chez Macroramphosus scolopax font l'objet du présent travail. En effet, les connaissances (Campagne Thalassa, station U20, 1994) (Ls = 100, 101, sur leur morphologie et leur histologie sont à la fois frag- 123 et 141 mm) et Notopogon xenosoma (MNHN 1994-23) mentaires et anciennes (Hertwig, 1872) et elles sont nette- (Ls = 93, 110 et 117 mm). Les prélèvements ont été faits ment insuffisantes pour établir des relations phylogéniques dans différentes régions du corps (Fig. la, b) sur ces spéci- entre les divers tissus qui les constituent. Nous avons aussi mens qui avaient été préalablement fixés au formol et voulu vérifier s'il était possible d'utiliser d'éventuelles conservés dans de l'alcool. caractéristiques morphologiques et/ou histologiques de ces formations tégumentaires comme critères systématiques D'autres prélèvements ont été réalisés dans la zone 8 dans la famille des Macroramphosidae. (Fig. 1b) sur un spécimen de chacune des espèces sui- vantes : Notopogon lilliei (MNHN 1995-11) (Ls = 165 mm), Les Macroramphosidae sont grégaires et benthiques et N. armatus (MNHN 1991-448) (Ls = 107 mm) et vivent jusqu'à plus de 500 mètres de profondeur. Ils sont Centriscops humerosus (MNHN 1989-301) (Ls = 157 mm) caractérisés par un corps comprimé latéralement, 3 ou 4 pour les observations au MEB. plaques osseuses dermiques latéro-dorsales (ou plaques épaxiales), un revêtement dermique composé de scutes, un Méthodes rayon épineux dorsal très développé et un museau tubulaire Etude morphologique terminé par des mâchoires réduites à de minuscules osselets Pour étudier la distribution des différents types de scutes formant un tout petit ‘‘bec’’ terminal (Le Danois, 1959 ; sur le corps, deux spécimens de Macroramphosus scolopax Duhamel, 1995). Cette famille comporte 7 espèces distri- et de Notopogon xenosoma ont été colorés à 1'alizarine par buées dans trois genres. Le premier genre, avec Macroram- la technique de Simons et Van Horn (1971) sans l'étape du phosus scolopax (L. 1758), possède deux nageoires dorsales bleu alcian ; des volets de peau (environ 25 mm2) ont été bien séparées ce qui le distingue des deux autres. Il possède prélevés et montés entre lame et lamelle pour des observa- une distribution géographique assez vaste. Les deux autres tions au microscope photonique. genres se différencient par le nombre de plaques épaxiales Pour tous les autres spécimens, des morceaux de tégu- (trois pour Notopogon et quatre pour Centriscops) ; ils ne ment (environ 50 mm2) ont été débarrassés de l'épiderme sont présents que dans la ceinture subtropicale et tempérée par un traitement à la potasse (1 %) pendant environ 15 de l'hémisphère sud. Le genre Centriscops est monospéci- minutes, puis rincés à l'eau distillée ; après déshydratation fique tandis que le genre Notopogon comporte cinq espèces dans une série croissante d'alcools les échantillons ont été réparties en deux groupes sur la base de la présence séchés, collés sur un support en nickel, métallisés et obser- (N. xenosoma Regan, 1914 et N. femandezianus Delfin, vés dans un microscope électronique à balayage, MEB 1899) ou de l'absence (N. armatus Sauvage, 1879, (JEOL-SEM-35). 0. FEDRIGO, F. J. MEUNIER, G. DUHAMEL, J.-Y. SIRE 191 Des tranches (sections transversales et longitudinales) d'environ 200 |im d'épaisseur ont été obtenues avec une tronçonneúse Isomet, amincies progressivement avec des abrasifs de grain décroissant et polies sur velours jusqu'à une épaisseur finale de 80 à 100 µm. Ces lames minces ont été
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