Hymenophyllum Seminar - the British Pteridological Society

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
Alignment of the sequences
Phylogeny = tree of
relationships
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
and growth forms
hemi-epiphyte
climbing
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
(New Zealand)
Serpyllopsis caespitosa
(Chile + Argentina)
Hymenoglossum cruentum
(Chile)
Rosenstockia rolandiprincipis (New Caledonia)
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
Ce
terrestrial
Cephalomanes atrovirens
Cephalomanes obscurum
Phylogenetic relationships in the trichomanoids
Callistopteris 5 sp
Paleotropics
Ca
Erect to short creeping rhizome
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)
Morphological variation
Venation anadromous or
catadromous
Ecological variation
Subg. Trichomanes > 30 sp
T. crispum
T. pinnatum (Guadeloupe)
Tr
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
(Costa Rica)
T. pedicellatum
(French Guyana)
Phylogenetic relationships in the trichomanoids
Polyphlebium
15 sp
Temperate regions of S. hemisphere, mountain forest in low latitudes
epiphytic
P. venosum
Po
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
Pantropical, extending to temperate regions of N. hemisphere
Terrestrial or climbing
Example: V. speciosum
V. giganteum (La Réunion)
Va
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)
• Subg. Nesopteris 4 sp.
Paleotropics
terrestrial
Cr. brevipes (Philippines)
Phylogenetic relationships in the hymenophylloids
rbcL + rps4-trnS +
rbcL-accD
(3641 bp)
1 genus: Hymenophyllum; 9 subgenera
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
H. nephrophyllum (New Zealand)
Subg Diploophyllum 1 sp
H. dilatatum (New Zealand)
D
C
Phylogenetic relationships in the hymenophylloids
Subg. Fuciforme 2 sp
Southern South America
F
Short creeping rhizome
H. fuciforme (Chile)
H. pulcherrimum (Chile)
Phylogenetic relationships in the hymenophylloids
Subg. Hymenoglossum 2 or 3 sp ?
Southern Chile and Madagascar
H. cruentum (Chile)
Hg + Pl
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)
Globosa
Basal subgenera
Sphaerocionium,
Mecodium,
Hymenophyllum
Pleuromanes
Myrmecostylum
Fuciforme
Diploophyllum
Cardiomanes
• All types of sori, but receptacles often broad and
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
70
cm
6
cm
sori
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
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
Hy
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
Hemi-epiphyte
ferns
climbing
True liana
epiphytic
terrestrial
Divergence times and evolution of habitats
• Estimation of divergence times
• Habitat coded using 4 states:
terrestrial
hemiepiphytic
climbing
epiphytic
T
• Ancestral state reconstruction
Diversification of
hymenophylloids more recent
than trichomanoids
Occurrence of epiphytism in the
Cretaceous, with major
radiation in the Tertiary
(Hennequin et al., 2008)
H
Conclusions & prospects
 New classification:
trichomanoids lineage: 8 genera
hymenophylloid lineage: 1 genus
Not all nodes of the phylogeny are strongly supported
some modifications
may require
 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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