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Diogenes Pugilator INPN 1 Le diogène des sables Diogenes pugilator (Roux, 1829) Citation de cette fiche : Noël P., 2016. Le diogène des sables Diogenes pugilator (Roux, 1829). in Muséum national d'Histoire naturelle [Ed.], 4 août 2016. Inventaire national du Patrimoine naturel, pp. 1-18, site web http://inpn.mnhn.fr Contact de l'auteur : Pierre Noël, SPN et DMPA, Muséum national d'Histoire naturelle, 43 rue Buffon (CP 48), 75005 Paris ; e-mail [email protected] Résumé. Chez le diogène des sables la carapace est dentée sur les côtés ; sa longueur peut atteindre 11 mm. La couleur générale est jaunâtre ou verdâtre, avec des lignes rouges sur les pattes. Les pédoncules oculaires sont forts et cylindriques. Le chélipède gauche est nettement plus fort que le droit et courbé sur le bord inférieur. Les pattes dépassent à peine la grande pince. L'espèce présente une forte variabilité morphologique intraspécifique. Les femelles sont ovigères en été. Il y a cinq stades larvaires zoé et une mégalope. L'espèce est dépositivore et a un comportement fouisseur. Divers poissons sont ses prédateurs. De nombreux parasites appartenant à des groupes variés (crustacés, vers...) ont été observés chez le diogène des sables. Pour s'abriter, il utilise des coquilles de nombreuses espèces de gastéropodes. Il se trouve de la côte à -40 m de profondeur sur des fonds sableux ou sablo-vaseux, et parfois dans des herbiers de phanérogames marines (zostères, posidonies, cymodocées) ou des zones avec des algues. Ce pagure est présent sur les côtes de l'Atlantique européen et africain de la Norvège à l'Angola, y compris la Méditerranée et la Mer Noire. Figure 1. Aspect général in situ, dans une coquille de Cerithium vulgatum. Photo © Stéphane Jamme. Classification : Phylum Arthropoda Latreille, 1829 > Sub-phylum Crustacea Brünnich, 1772 > Super-classe Multicrustacea Regier, Shultz, Zwick, Hussey, Ball, Wetzer, Martin & Cunningham, 2010 > Classe Malacostraca Latreille, 1802 > Sous-classe Eumalacostraca Grobben, 1892 > Super- ordre Eucarida Calman, 1904 > Ordre Decapoda Latreille, 1802 > Sous-ordre Pleocyemata Burkenroad, 1963 > Infra-ordre Anomura H. Milne Edwards, 1832 > Super-famille Paguroidea Latreille, 1802 > Famille Diogenidae Ortmann, 1892 > Genre Diogenes Dana, 1851. Synonymes (Zariquiey Álvarez 1968 ; McLaughlin & al. 2010 ; Noms vernaculaires: GBIF 2016 ; INPN 2016 ; WoRMS 2016): Diogène (Loyer 1995 ; Livory 2001) ; pagure des sables Diogenes pugilator var. intermedius Bouvier, 1891 (Göthel 1996 ; Vacquier 2007 ; WoRMS 2016) ; pagure du Diogenes pugilator orientalis Codreanu & Balcesco 1968 sable (Luther & Fiedler 1965) ; pagure athlète (Roux 1829) ; Diogenes pugilator pugilator Roux, 1829 diogène boxeur ; pagure boxeur. Pagurus algarbiensis de Brito Capello, 1875 Pagurus arenarius Lucas, 1846 Principaux noms étrangers. Pagurus bocagii de Brito Capello, 1875 Anglais : small hermit crab, sand hermit crab, South-claw Pagurus curvimanus Clément, 1874 hermit crab. Pagurus dillvyni A. Milne-Edwards & Bouvier, 1900 Allemand : Diogeneskrebs ; Sand-Einsiedler. Pagurus dillwynii Bate, 1851 Espagnol : ermitaño guerrero. Pagurus dilwynnii Alcock, 1905 Catalan : Bernat ermità de sorra. Pagurus Lafonti Fischer, 1872 Italien : paguro pugile. Pagurus ponticus Kessler, 1860 Néerlandais : Kleine heremietkreeft, kleine heremiet, klein Pagurus pugilator Roux, 1829 heremietkreeftje. Pagurus varians Costa, 1838 Ukrainien : Рак-самітник діоген. Diogenes varians Heller, 1863. N° des bases de données : GBIF ID : 5716791 ; INPN Diogenes pugillator [erreur occasionnelle]. Cd_Nom : 18465 ; WoRMS ; AphiaID : 107199. 2 Etymologie : Le genre Diogenes est dédié au philosophe Diogène de Sinope (-412 -323 AC) qui vécut dans un tonneau ; la coquille où vit le pagure est assimilé à un tonneau (Perrier 1929) ; le nom d'espèce pugilator désigne un athlète au pugilat ou pugiliste, c'est à dire un boxeur. << D'un caractère peut-être plus irascible que ses congénères, il semble s'agiter vivement lorsqu'on le saisit, et loin de se hâter de se soustraire au danger en se blotissant au fond de son domicile, on dirait qu'il cherche à frapper de son poing celui qui l'inquiète >> (Roux 1828). Fig. 2. Vue dorsale. Iconographie illustrant la description originale de l'espèce © Roux 1829. Description (Bouvier 1940 ; Zariquiey Álvarez 1968). La carapace est dentée sur les côtés en avant du sillon subcervical. Les dents latérales du front sont assez nettes. Les pédoncules oculaires sont forts et cylindriques plus courts que les pédoncules antennulaires et antennaires. La saillie médiane de l'arceau oculaire est en pointe étroite. L'écaille ophthalmique est denticulée sur son bord oblique antérieur. L'écaille antennaire est en triangle aigu ; elle est denticulée en dedans. Les fouets antennaires portent des soies. Le chélipède gauche est fortement courbé sur son bord inférieur. Il est nu, avec une rangée de dents au bord inférieur du mérus et au bord supérieur du carpe et du propode. Ces deux derniers articles ont de fins granules aigus sur leur face externe. Les doigts sont calcifiés jusqu'à la pointe et sont en contact ; ils ont quelques grosses dents en regard sur leurs bords respectifs. Le chélipède droit est bien plus réduit que la gauche. Sa pince est beaucoup moins courbe au bord inférieur et porte des petits tubercules aigus et de longs poils sur sa face externe. Il y a une rangée de dents au bord supérieur du carpe. Le chélipède gauche présente un dimorphisme sexuel important d'une part et une variabilité individuelle d'autre part. Chez les femelles, il est plus large et arrondi et le carpe et la "main" sont courts et couverts de dents (Forest & Guinot 1956 ; Zariquiey Álvarez 1968) (voir figure 3). Les pattes locomotrices sont comprimées latéralement. Elles dépassent à peine en longueur la grande pince et sont légèrement dentées au bord antérieur du carpe et du propode. Leur dactyle a une forme de sabre, est plus long que le propode et possède une petite griffe terminale et un sillon longitudinal. Ces pattes portent souvent des soies longues et nombreuses. Le telson est un peu denté sur sa marge postérieure. La longueur de la carapace est de 2,5 à 11 mm chez les mâles, 3 à 9 mm chez les femelles et 3,5 mm à 11 mm chez les femelles ovigères (Forest 1956 ; Forest & Guinot 1956). 3 Figure 3. Variabilité morphologique de la pince gauche. © Forest & Guinot 1956. La couleur d'ensemble est décrite comme variée selon les auteurs : brun livide (Roux 1828), jaunâtre ou verdâtre, souvent avec des lignes rouges sur les pattes (Bouvier 1940 ; Holthuis 1956), bleu verdâtre lavé de blanc et de brun (Bohn 1901), vert ou rouge clair taché de plus foncé (Perrier 1929), jaune finement tacheté de rouge (de Brito Capello 1875), blanc-gris taché de brun-rouge, abdomen bleu sur le sable et gris-bleu sur la vase (Noël 1992). Figure 4. Diogenes pugilator sorti de sa coquille. Figure 5. Vue dorsale, fide Holthuis & al. 1986. Banyuls-sur-mer, 10/05/1975. Photo ©Jean Lecomte. Risques de confusion, espèces voisines, variations infra-spécifiques. La grande variabilité morphologique de cette espèce a été signalée et étudiée par divers auteurs (Brito Capello 1875 ; Chevreux & Bouvier 1892 ; Bouvier 1901 ; Forest 1956 ; Rossignol 1962). Cette variabilité a conduit d'autres auteurs à décrire des espèces ou des variétés nouvelles (Miers 1881 ; Bouvier 1891 ; Balss 1921) qui ont été mises en synonymie par la suite. Ces variations concernent la pilosité qui peut être plus ou moins accentuée, les pinces dont la longueur peut être plus ou moins importante, et le nombre des denticulations sur le bord antéro-externe des écailles ophthalmiques qui sont de 4 ou 5 chez la forme pugilator typique contre 10 à 12 chez la forme denticulatus et jusqu'à 16 à 18 sur la côte catalane (Bouvier 1940 ; Zariquiey Álvarez 1968). Dans le genre Diogenes il existe une soixantaine d'espèces (essentiellement tropicales) au niveau mondial (McLaughlin & al. 2010 ; GBIF 2016 ; WoRMS 2016). D. pugilator est la seule espèce de ce genre en Europe 4 (d'Udekem d'Acoz 1999). Dans les zones sous juridiction française, il convient de signaler la présence de Diogenes pallescens Whitelegge, 1897 à La Réunion, en Nouvelle Calédonie et en Polynésie (INPN 2016). Biologie. Reproduction et développement. L'anatomie et l'ultrastructure de l'appareil génital mâle de D. pugilator ont été étudiées par Manjón-Cabeza & García Raso (2000) et les spermatozoïdes et les spermatophores l'ont été par Tirelli & al. (2008). L'espèce est gonochorique. Une stimulation sexuelle normalement liée à l'accouplement est nécessaire pour qu'il y ait mue puis ponte (Bloch 1933). Le mâle attend donc la mue de la femelle pour s'accoupler en une sorte de promenade pré-nuptiale au cours de laquelle le mâle s'agrippe à la coquille occupée par la femelle (Bloch 1935). Ceci est parfaitement illustré par une photo dans Anonyme (2016). Les femelles sont ovigères en été, de mai à septembre à Roscoff (Bourdon 1965) et de juin à août seulement en Irlande (McGrath & al. 2000). Les œufs ont un diamètre de 0,36 x 0,46 mm (Zariquiey Álvarez 1968). Les larves et le développement ont fait l'objet de différents signalements et études (Gurney 1927 ; Menon 1937 ; Gurney 1927, 1942 ; MacDonald & al. 1957 ; Pike & Williamson 1958, 1960 ; Bourdillon Casanova 1960 ; Thiriot 1974 ; Barnich 1996 ; González-Gordillo & al. 2001). Il y a cinq stades zoé et une mégalope (dos Santos 1999). En Bretagne à Roscoff, les larves sont rares dans le plancton de mai à novembre (Thiriot 1963) ; les glaucothoës et les premiers stades post-larvaires s'y observent en juin et en septembre (Pérez 1928 ; Bourdon 1965). Les larves sont présentes toute l'année sur les côtes du Portugal (Paula 1987 ; dos Santos 1999). Il y a plusieurs périodes de recrutement dans l'année (Dolbeth & al. 2006). La croissance de l'espèce est relativement rapide (Manjón- Cabeza & García Raso 1998a, 1998b) et la longévité chez cette espèce a été estimée à 21 mois dans une fourchette de 1 ou 2 ans (Dolbeth & al.
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