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Trichomanes (1996) • Kathleen Pryer & J.-Y British Pteridological Society Meeting, Reading, 31 Oct. 2009 Genetic relationships within the Hymenophyllaceae Sabine Hennequin Université Pierre et Marie Curie, Paris Centre de recherche sur la Paléobiodiversité et les Paléoenvironnements Systematics of the Hymenophyllaceae Previous works on the family: main contributors • C. Presl (1843, 1849) • E. B. Copeland (1937, 1938, 1942) • C. V. Morton (1968) • R. Pichi Sermolli (1977) • K. Iwatsuki (1970es and 1980es) Since the development of molecular phylogeny: • Jean-Yves Dubuisson: PhD on Trichomanes (1996) • Kathleen Pryer & J.-Y. Dubuisson (2001): first phylogeny of the family • S. Hennequin: PhD on Hymenophyllum (2004) • Atsushi Ebihara: PhD on Hymenophyllaceae, with a focus on Asiatic region (2005) several publications since 1996 Revision of the Hymenophyllaceae (Ebihara and coll. 2006, in Blumea) Phylogenetic reconstruction using DNA sequences In the lab: DNA extraction, amplification, sequencing On the computer: (Natural Histrory Museum, Darwin Center labs) DNA sequences Phylogeny = tree of relationships Alignment of the sequences Various phylogenetic methods Position of Hymenophyllaceae in the fern tree basal lineage of leptosporangiate ferns origin: Permian (270 Ma; Pryer et al. 2004) Pryer et al. 2004 Characteristics of the family • very thin lamina (usually 1 cell thick) • marginal sorus involucre receptacle • ~ 600 sp. • hygrophilous strategy • distributed in tropical rain forests and humid temperate forests; a few representatives in Europe Habitat diversity in filmy ferns climbing and growth forms hemi-epiphyte true liana epiphytic terrestrial Mare Longue forest, 300m, La Réunion Bélouve forest, 1600m, La Réunion The Hymenophyllaceae were traditionally divided in 2 genera Trichomanes (~300 species) tubular sorus Hymenophyllum (~300 species) bivalved sorus Problematic taxa of the family Cardiomanes reniforme Hymenoglossum cruentum (New Zealand) (Chile) Rosenstockia rolandi- Serpyllopsis caespitosa principis (New Caledonia) (Chile + Argentina) Microtrichomanes (9 to 14 species) Phylogeny of the Hymenophyllaceae rbcL gene 96 Trichomanes s.l. and 50 Hymenophyllum s.l. "Trichomanes s.l." Serpyllopsis caespitosa Rosenstockia rolandi-principis "Hymenophyllum s.l." Microtrichomanes Trichomanes pallidum Hymenoglossum cruentum Cardiomanes reniforme Hennequin et al. 2008 Phylogeny of the Hymenophyllaceae trichomanoids (~300 species) Terrestrial, climbing, or epiphytic Various growth forms Tubular sori hymenophylloids (~300 species) Epiphytic long-creeping rhizomes Mostly bivalved sori incl. problematic taxa Phylogenetic relationships in the trichomanoids Ebihara et al. 2006 Phylogenetic relationships in the trichomanoids Cephalomanes 4 sp Paleotropics Erect to short creeping rhizome Venation anadromous Blades once pinnate to bipinnatifid terrestrial Ce Cephalomanes atrovirens Cephalomanes obscurum Phylogenetic relationships in the trichomanoids Callistopteris 5 sp Paleotropics Erect to short creeping rhizome Ca Venation anadromous Bristle-like reddish hairs on stipes and rachises terrestrial C. apiifolium Phylogenetic relationships in the trichomanoids Abrodictyum 25 sp Pantropical Laminar cells reduced terrestrial Ab A. meifolium (La Réunion) A. caudatum Phylogenetic relationships in the trichomanoids Trichomanes > 60 sp Neotropical (+ a few in Africa - Indian Ocean) Venation anadromous or Morphological variation catadromous Tr Ecological variation Subg. Trichomanes > 30 sp T. crispum T. pinnatum (Guadeloupe) Phylogenetic relationships in the trichomanoids (Trichomanes) Subg. Feea > 5 sp T. osmundoides (Guadeloupe) Subg. Davalliopsis > 1 sp T. elegans (Guadeloupe) Subg. Lacostea > 4 sp T. tuerckheimii T. pedicellatum (Costa Rica) (French Guyana) Phylogenetic relationships in the trichomanoids Polyphlebium 15 sp Temperate regions of S. hemisphere, mountain forest in low latitudes Po epiphytic P. venosum Phylogenetic relationships in the trichomanoids Didymoglossum > 30 sp Di Pantropical Dwarf epiphytes Roots absent (but root-like shoots) D. kapplerianum (Guyane française) D. punctatum (Guadeloupe) D. membranaceum (Guadeloupe) Phylogenetic relationships in the trichomanoids Vandenboschia > 15 sp Va Pantropical, extending to temperate regions of N. hemisphere Terrestrial or climbing Example: V. speciosum V. giganteum (La Réunion) Phylogenetic relationships in the trichomanoids Cr Crepidomanes > 30 sp Paleotropics, extending to N temperate regions False veinlets parallel to true veins • Subg. Crepidomanes epiphytic Roots absent (but root-like shoots) Cr. bipunctatum (La Réunion) Cr. brevipes (Philippines) • Subg. Nesopteris 4 sp. Paleotropics terrestrial Phylogenetic relationships in the hymenophylloids rbcL + rps4-trnS + 1 genus: Hymenophyllum; 9 subgenera rbcL-accD (3641 bp) Hymenophyllum ~100 sp (incl. Rosenstockia, Serpyllopsis, Hemicyatheon, Craspedophyllum...) Mecodium ~70 sp Sphaerocionium ~70 sp (incl. Microtrichomanes) Globosa ~25 sp Myrmecostylum > 8 sp Pleuromanes 5 sp Hymenoglossum 3 sp ? Diploophyllum 1 sp Fuciforme 2 sp Cardiomanes 1 sp Phylogenetic relationships in the hymenophylloids Subg.Cardiomanes 1 sp Subg Diploophyllum 1 sp D C H. nephrophyllum (New Zealand) H. dilatatum (New Zealand) Phylogenetic relationships in the hymenophylloids Subg. Fuciforme 2 sp Southern South America Short creeping rhizome F H. fuciforme (Chile) H. pulcherrimum (Chile) Phylogenetic relationships in the hymenophylloids Subg. Hymenoglossum 2 or 3 sp ? Southern Chile and Madagascar Hg + Pl H. cruentum (Chile) Phylogenetic relationships in the hymenophylloids Subg. Pleuromanes 5 sp Asia-Pacific Hg + Pl H. pallidum H. flabellatum Phylogenetic relationships in the hymenophylloids Subg. Myrmecostylum > 8 sp My Southern Chile, New Zealand, New Caledonia H. krauseanum (Chile) paniense H. plicatum (Chile) H. scabrum (New Zealand) Phylogenetic relationships in the hymenophylloids Subg. Globosa ~25 sp Gl Tropics to temperate regions of Asia-Pacific; +1 in southern South America glabrous H. flexuosum (Pacific) H. caudiculatum (Chile) Phylogenetic relationships in the hymenophylloids Characteristics of the "basal" subgenera • 2-4 cell thick (at least partly) • Rhizome generally thicker (diameter > 1mm) • Fronds generally longer (> 20 cm) • Rhizome anatom : stele more developped (protostele with xylemian parenchyma, or dorsi-ventral) Sphaerocionium, Basal subgenera Mecodium, Globosa Hymenophyllum Pleuromanes Myrmecostylum Fuciforme Diploophyllum • All types of sori, but receptacles often broad and Cardiomanes bearing sporangiophores H. badium H. dilatatum • Chromosome number n = 36 (also in Sphaerocionium) Phylogenetic relationships in the hymenophylloids Subg. Sphaerocionium ~70 sp Sph (incl. Microtrichomanes); pantropical Hairs on the axes and margin, sometimes lamina 2 cm H. digitatum Up to sori 70 6 cm cm H. capillare H. hygrometricum H. hirsutum H. digitatum Phylogenetic relationships in the hymenophylloids Mec Subg. Mecodium ~70 sp (incl. Microtrichomanes); pantropical n = 28 Glabrous, entire margins H. polyanthos (La Réunion – Guadeloupe) Note: Species of Diploophyllym, Globosa, Fuciforme, and some of Myrmecostylum and Pleuromanes formerly placed in Mecodium Phylogenetic relationships in the hymenophylloids Hy Subg. Hymenophyllum ~100 sp n = 11, 12, 13, 14, 18, 21, 22, 26, 28 Dentate margin Sori with a broaden base, position close to rachis Hairs on axes H. tunbrigense Phylogenetic relationships in the hymenophylloids Temperate or high elevation clade Some minute species (fronds < 3 cm) H. armstrongii (New Zealand) H. peltatum (Chile) H. caespitosum (Chile) H. pectinatum (Chile) H. sibthorpioides (La Réunion) Phylogenetic relationships in the hymenophylloids Asia-Pacific clade n =21 or 22 H. revolutum (New Zealand) H. bivalve (New Zealand) H. barbatum (Japan) Phylogenetic relationships in the hymenophylloids Relationships within subg. Hymenophyllum Temperate or high elevation Asia-Pacific Southern South America New Zealand taxa (Hennequin et al., submitted) Habitat diversity in filmy climbing ferns Hemi-epiphyte True liana epiphytic terrestrial Divergence times and evolution of habitats • Estimation of divergence times • Habitat coded using 4 states: terrestrial hemiepiphytic T climbing epiphytic • Ancestral state reconstruction Diversification of hymenophylloids more recent than trichomanoids H Occurrence of epiphytism in the Cretaceous, with major radiation in the Tertiary (Hennequin et al., 2008) Conclusions & prospects New classification: trichomanoids lineage: 8 genera hymenophylloid lineage: 1 genus Not all nodes of the phylogeny are strongly supported may require some modifications Regressive evolution: adaptation to very humid conditions, moss strategy 250 species sequenced out of the 600 of the family (160 /300 in trichomanoids, 90/300 in Hymenophyllum) still plenty of work to do! Still to investigate: biogeography, evolution of the sorus, chromosome numbers Aknowledgments: UMR 7207 at University Pierre et Marie Curie & MNHN Paris: Jean-Yves Dubuisson Natural History Museum Tokyo: Atsushi Ebihara Department of Biology, Duke University: Kathleen Pryer, Eric Schuettpelz NHM London: H. Schneider Sponsors: NSF CAREER, NESCent, Synthesys All photographers sharing pictures on the web! .
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