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1 L'hydrodictye Réticulée Hydrodictyon Reticulatum (Linnaeus)

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1 L'hydrodictye Réticulée Hydrodictyon Reticulatum (Linnaeus) 1 L'hydrodictye réticulée Hydrodictyon reticulatum (Linnaeus) Bory de Saint-Vincent 1824 Citation de cette fiche : Couté A., Noël P., 2016. L'hydrodictye réticulée Hydrodictyon reticulatum (Linnaeus) Bory de Saint-Vincent 1824. in Muséum national d'Histoire naturelle [Ed.], 6 septembre 2016. Inventaire national du Patrimoine naturel, pp. 1-9, site web http://inpn.mnhn.fr Contact des auteurs : Alain Couté, Muséum national d'Histoire naturelle, Département RDDM, UMR 7245 CNRS/MNHN, éq. Cyanobactéries, Cyanotoxines et Environnement, 57 rue Cuvier (CP 39), 75231 Paris cedex 05 ; e-mail [email protected] ; Pierre Noël, Muséum, SPN et DMPA, 43 rue Buffon (CP 48), 75231 Paris cedex 05 ; e-mail [email protected] Résumé Hydrodictyon reticulatum est une algue verte organisée en réseau dont les cellules, allongées et cylindriques, ont un chloroplaste pariétal réticulé avec de nombreux pyrénoïdes au stade âgé. Jeunes, elles ne possèdent qu’un noyau tandis qu’elles en ont un nombre important en vieillissant. Les cellules, qui mesurent jusqu’à 1,5cm de longueur, sont disposées en un réseau à mailles polyédriques en forme de sac. La taille du maillage augmente avec l’âge et les individus les plus anciens peuvent mesurer jusqu’à 20cm de longueur. Cette micro-algue est libre sur le sédiment ou flotte à la surface de l’eau. Elle prolifère en période estivale et peut subsister assez tard dans l’année grâce à une multiplication et une reproduction complexes faisant intervenir des cellules mobiles à deux flagelles égaux. Cette espèce ubiquiste se rencontre en faciès lentique dans divers types de plans d’eau naturels et artificialisés (mares, rizières…). H. reticulatum est présent un peu partout dans le monde dans les régions tempérées et tropicales, présentant une alternance de périodes d’inondations et de sécheresses. Figure 1. Aspect général et détail, d'après Komarek & Fott (1983) et Bourrelly (1990). Classification : Règne Plantae Haeckel 1866 > Phylum Chlorophyta Pascher 1914 > Classe Chlorophyceae Wille 1884 > Ordre Sphaeropleales Luerssen 1877 > Famille Hydrodictyaceae (Gray) Dumortier 1829 > Genre Hydrodictyon Roth 1797. Synonymes (AlgaeBase, 2016 ; INPN, 2016 ; WoRMS, 2016): Noms vernaculaires: Conferva reticulata Dillenius 1742 Historia Filet à réseaux (Bory de Saint-Vincent, 1824). muscorum., t. 4, f. 14. [lectotype fide Irvine in Hydrodyctie (Bory de Saint-Vincent, 1824). Spencer, Irvine & Jarvis 2009: 244]. Filet d'eau (Saint-Maxent, 2001 ; DORIS, 2016). Conferva reticulata Linnaeus 1753 : 1165 Réseau d’eau (Collectif, 1997 ; Saint-Maxent, 2001). Conferva reticulatum Linnaeus 1753 Principaux noms étrangers. Byssus reticulata (Linnaeus) F. H. Wiggers 1780 Anglais : water net (Wells & al., 1999). Byssus cancellata Linnaeus 1767 Allemand : Wassernetz. Espagnol : red de agua. Hydrodictyon utriculatum Roth 1800 Néerlandais : waternetje. Hydrodictyon pentagonum Vaucher 1803 Danois : Vandnet. Japonais : amimidoro. Hydrodyction reticulatum Bory de Saint-Vincent 1824 Russe : водяная сеточка ; гидродикцион сетчатый. N° des bases de données : AlgaeBase : 27496 ; GBIF ID : 7890524 ; INPN Cd_Nom : 707372 ; ITIS : 6080 ; WoRMS AphiaID : 381000. Etymologie : Hydrodictyon : du grec [hydro] = eau, mouiller, onde et du grec [dictuon] = filet de pêche, soit <<filet aquatique>>. reticulatum : du latin [reticulum] = petit filet, réseau, résille, treillis. NB. Entre le genre et l'espèce, il s'agit donc plus ou moins d'une tautologie ou d'un pléonasme insistant bien sur l'aspect très particulier de cette algue ! 2 Description. Hydrodictyon reticulatum est une algue verte filamenteuse réticulée dont les cellules, allongées, cylindriques, avec un chloroplaste en réseau, mesurent jusqu’à 1,5cm de longueur. À l’état jeune, elles ne possèdent qu’un noyau et un pyrénoïde mais le nombre de ces derniers s’accroît rapidement avec l’âge. Les cellules sont organisées en un réseau à mailles polyédriques le plus souvent hexagonales ou pentagonales (en réalité on observe de quatre à sept côtés). L’ensemble du thalle est d’abord en forme de sac cylindrique arrondi aux deux extrémités et de couleur verte. Les individus les plus âgés sont de couleur jaune, de grande taille, visibles à l’œil nu, mesurant plusieurs centimètres de longueur et de largeur. La taille du maillage augmente avec l’âge et le coenobium ou cénobe (= réseau à mailles) dans sa totalité peut atteindre 20cm et même 1m (en Nouvelle Zélande, selon Wells et al., 1999). Le thalle est souvent déchiré et l’ensemble se présente sous l’aspect d’un tulle vert à mailles hexagonales. Chaque cellule produit, par divisions mitotiques, un grand nombre de zoospores biflagellées qui nagent un certain temps sans sortir de la paroi de la cellule maternelle. Les zoospores finissent par se regrouper en un jeune cénobe réticulé qui sera libéré par dégradation de la paroi maternelle (Lukov, 1964 ; Bourrelly, 1990 ; Saint-Maxent, 2001 ; Chou et al., 2006). H. reticulatum vit libre dans la colonne d’eau, posé sur le fond ou encore flottant en surface (Champion & Clayton, 2000 ; Chou et al., 2006). Le démarrage de la période de développement de cette micro-algue est généralement assez tardif, à la fin du printemps. La prolifération est le plus souvent estivale et peut persister assez tard en saison (Saint Maxent, 2001). Figure 2. Hydrodictyon reticulatum in situ ; photo prise en plongée à faible profondeur. L'aselle (Asellus aquaticus) donne l'échelle © Claude Zimmermann, Golf National, Saint-Quentin en Yvelines, 19 juin 2016. Risques de confusion, espèces voisines, variations infra-spécifiques. On connait quatre espèces pour le genre Hydrodictyon (AlgaeBase, 2016 ; WoRMS, 2016) : H. africanum S. Yamanouchi 1913 et H. patenaeforme Pocock 1937 forment des filets aplatis en une seule nappe de quelques centaines de cellules seulement tandis que H. reticulatum (Linnaeus) Bory de Saint Vincent 1824 et H. indicum Iyengar 1925, forment classiquement des sacs faits de milliers de cellules qui ne se séparent à maturité que chez H. indicum (Chou et al., 2006). L’espèce H. reticulatum se subdivise en trois variétés (WoRMS, 2016) : H. reticulatum var. bernardii Playfair 1914, H. reticulatum var. minimum Playfair 1914 et H. reticulatum var. nodosum Playfair 1914. Biologie. Multiplication : H. reticulatum est composé de milliers de cellules cénocytiques ne se séparant pas à maturité (Coffey & Miller, 1988 ; Chou et al., 2006). La multiplication (qui n’intervient qu’en conditions favorables) se réalise par production de « zoospores » biflagellées qui au final forment un filet ou cénobe fils, d’abord retenu à 3 l’intérieur de la cellule mère (Pocock, 1960 ; Marchant & Pickett-Heaps, 1972a). À maturité, les cellules mères du cénobe relâchent de nouveaux petits filets microscopiques. Reproduction : (intervient en conditions défavorables) elle s’effectue par le biais de « zoïdes » ou gamètes biflagellés tous de même dimensions, qui, en fusionnant, engendrent des zygospores (= zygote ou œuf). Celles-ci subissent la méïose et donnent naissance à 4 ou 8 zoospores, puis des stades polyèdriques et enfin un filet plat de 100 à 300 cellules (Marchant & Pickett-Heaps, 1972d). Chaque cellule de ce dernier peut produire de petits filets fils cylindriques végétatifs (Coffey & Miller, 1988). Les gamètes non fécondés (azygospores) donnent des spores uniques qui se transforment aussi en stades polyèdriques. H. reticulatum a donc un cycle biologique complexe (Marchant & Pickett-Heaps, 1971, 1972a, 1972b, 1972c, 1972d ; Marchant, 1972 ; Pirson, 1972) présentant une alternance de stades mobiles et de stades fixes. Trois de ces stades, les zygospores, les azygospores et les polyèdres (= cellules de résitance), ont une paroi épaisse renfermant des substances de réserve. Ils peuvent constituer des stades de résistance (= dormants) du cycle vital (Coffey & Miller, 1988). On observe des variations annuelles ou pluriannuelles avec apparition et disparition de nombreux individus sans qu’on en connaisse la cause (Wells et al., 1999 ; Pierre, 2001). Le taux de croissance est optimal pour des températures comprises entre 26 et 30°C (Mathey, 1993). Parasites : H. reticulatum peut être parasité par un champignon microscopique, Plyctochytrium hydrodictyi (Braun A. 1855) Schrôter J. 1882 (Duché 1945 [comme H. utriculatum]), Chitridineae, Rhyzidiaceae. Ecologie. H. reticulatum présente un caractère ubiquiste marqué. Cette micro-algue se rencontre communément en faciès lentique dans divers types de plans d’eau naturels : cours d’eau avec des cuvettes, fossés, marais, mares et prairies marécageuses (Van Meel, 1944 ; Prescott, 1970 ; Coffey & Miller, 1988 ; Rodriguez & Vergon, 1996 ; Lambert-Servien et al., 2008) ou en zones artificielles comme des installations liées à la pisciculture, des rizières (Pocock, 1960 ; Chou et al., 2006), des pièces d’eau d’agrément (Coffey & Miller, 1988), des terrains de golf ou des lavoirs (DORIS, 2016), ou encore des réservoirs de stockage d’eau destinée, après traitement, à l’alimentation. Des cénobes végétatifs ou des spores de résistance peuvent être transportés par des oiseaux aquatiques, des insectes volants ou encore, de façon accessoire, par les transferts de poissons ou de plantes aquatiques (Coffey & Miller, 1988). L’espèce peut ainsi coloniser de nouveaux secteurs. L’habitat typique serait les régions tempérées avec alternance de périodes d’inondation et de sécheresse comme la vallée du Nil. Cette micro-algue peut proliférer à faible pH destiné à traiter d’autres proliférations algales avec un algicide comme le sulfate de cuivre (Fitzgerald,
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