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
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