| SYSTEMATIC BIOLOGY |
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| THE VASCULAR CRYPTOGAMS | |||||
Vascular Cryptogam is an old botanical phrase, and it refers to those vascular plants that do not make seeds. Thus, cryptogam (literally hidden gametophyte) refers to the production of a separate, usually very small, archegoniate gametophyte. These are well represented in the fossil record. Kenrick and Crane (1997) report that individual spores of land plants (these could be mosses, hornworts or vascular plants) have been found in the Lower Silurian (see the Geological Time Scale), a phase in the evolution of land plants that they refer to as "Eotracheophytic". Unequivocal evidence of vascular land plants has been found in the mid Silurian, when the first radiation of tracheophytes occurred. The early Devonian Period, however, saw an explosion in the diversity of early land plants which continued through the mid Permian, a time called the Eutracheophytic by Kenrick and Crane (1997). In fact, we know so much about the diversity and structural details of the extinct taxa that taxonomic systems of the vascular cryptogams usually include them (e.g., Bold et al. 1987). The wealth of fossil evidence from the Devonian to the present also provides the basis and support for the Telome Theory, a theory about the origins of all vascular plant organs from a simple dichotomizing axis (see Gorentz et al., 2002 for a good review of that theory). Pearson (1995) explored the relationships between the green plants primarily by summarizing structural and developmental evidence. His phylogenies relevant to the vascular cryptogams include a diagram of the origin and early divergence and the fern taxa. Rothwell (1999) examined 101 morphological characters of extinct and extant taxa from all major groups of vascular cryptogams. His analysis supports the phylogenetic relationships that are assumed in Pearson (1995) and Bold et al. (1987). Interpretations of relationships based on molecular evidence can be seen in a diagram from Pryer et al. (2001). Combined structure, fossil, and molecular evidence can be seen in Tudge (2000) and the Tree of Life Project. Clearly, as indicated by Bold et al. (1987) in the case of the vascular cryptogams "Nature mocks at human categories". With that advice, I present a taxonomic system of the vascular cryptogam phyla that roughly conforms to that of Bold et al. (1987), as modified by Kenrick and Crane (1997) and Smith et al. (2006). Those taxa that are marked with a cross (+) and in maroon letters are extinct (as are all lower taxa within that taxon). Orders are followed by representative genera.
Concepts and Terms for the Vascular Cryptogams CLASS RHYNIOPSIDAORDER RHYNIALES
Rhynia, Horneophyton, Cooksonia, Hicklingia, Eogaspesiea, Taeniocrada, Hediea, Yarravia, Hostomella, Nothia?
CLASS ZOSTEROPHYLLOPSIDA
ORDER ZOSTEROPHYLLALES.
Baragwanathia, Zosterophyllum, Sawdonia, Kaulangiophyton, Crenaticaulis, Serrulacaulis.
CLASS TRIMEROPHYTOPSIDA
ORDER TRIMEROPHYTALES.
Dawsonites, Pertica, Psilophyton, Trimerophyton.
CLASS AGLOSSOPSIDA
ORDER ASTEROXYLALES+
Asteroxylon
ORDER LYCOPODIALES
Lycopodium, Huperzia, Lycopodites+.
ORDER PROTOLEPIDODENDRALES+
Protolepidodendron, Baragwanthathia, Leclercqia.
CLASS GLOSSOPSIDA
CLASS CLADOXYLIDOPSIDA+ORDER LEPIDODENDRALES+
Sigillaria, Lepidodendron, Lepidocarpon, Lepidophloios, Phytokneme.
ORDER SELAGINELLALES
Selaginella, Selaginellites+
ORDER PLEURONEMIALES+
Pleuronemia.
ORDER ISOETALES
Isoetes, Nathorstiana+, Isoeites+, Stylites+.
ORDER CLADOXYLALES
Cladoxylon
ORDER PSEUDOSPOROCHNALES
Pseudosporochnus
ORDER CALAMOPHYTALES
Calamophyton
CLASS COENOPTERIDOPSIDA+
ORDER STAUROPTERIDALES
Stauropteris
ORDER BOTRYOPTERIDALES
Botryopteris
ORDER RHACOPHYTONALES
Rhacophyton, Protocephalopteris, Protopteridophyton
ORDER ZYGOPTERIDALES
Zygopteris, Clepsydropsis, Etapteris, Metaclepsydropsis, Ankyropteris
CLASS PSILOTOPSIDA
ORDER OPHIOGLOSSALES
Family Ophioglossaceae: Botrychium, Helminthostachys, Mankyua, Ophioglossum, Cheiroglossa
ORDER PSILOTALES
Family Psilotaceae: Psilotum, Tmesipteris
CLASS EQUISETOPSIDA [SPHENOPSIDA]
ORDER HYENIALES+
Hyenia
ORDER EQUISETALES
Family Equisetaceae: Equisetum, Equisetites+, Phyllotheca+
ORDER CALAMITALES+
Calamites, Calamostachys, Calamocarpon, Annularia, Asterophyllites.
ORDER PSEUDOBORNIALES+
Pseudobornia
ORDER SPHENOPHYLLALES+
Sphenophyllum, Bowmanites
CLASS MARATTOPSIDA
ORDER MARATTIALES
Family Marattiaceae: Angiopteris, Christensenia, Danaea, Marattia, Archangiopteris (?)
CLASS PTERIDOPSIDA [=FILICOPSIDA, POLYPODIOPSIDA]
ORDER OSMUNDALES
Family Osmundaceae: Osmunda, Leptopteris, Todea
ORDER HYMENOPHYLLALES
Family Hymenophyllaceae: Hymenophyllum, Cardiomanes, Hymenoglossum, Rosenstockia, Serpyllopsis, Sphaerocionium(?), Mecodium(?), Microtrichomanes, Abrodictyum, Callistopteris, Cephalomanes, Crepidomanes, Didymoglossum, Polyphlebium, Trichomanes, Vandenboschia
ORDER GLEICHENIALES
Family Gleicheniaceae (incl. Dicranopteridaceae, Stromatopteridaceae): Dicranopteris, Diplopterygium, , Gleichenella, Gleichenia, Sticherus, Stomatopteris.
Family Dipteridaceae (incl. Cheiropleuriaceae): Cheiropleuria, Dipteris
Family Matoniaceae: Matonia, Phanaerosorus
ORDER SCHIZAEALES
Family Lygodiaceae: Lygodium
Family Anemiaceae (incl. Mohriaceae): Anemia, Mohria
Family Schizaeaceae: Actinostachys, Schizaea, Anachorpteris*
ORDER SALVINIALES
Family Marsileaceae (incl. Pilulariaceae): Marsilea, Pilularia, Regnellidium.
Family Salviniaceae (incl. Azollaceae): Azolla, Salvinia.
ORDER CYATHIALES
Family Thyrsopteridaceae: Thyrsopteris
Family Loxomataceae: Loxomopsis, Loxoma
Family Culcitaceae: Culcita
Family Plagiogyriaceae: Plagiogyra
Family Cibotiaceae: Cibotium
Family Cyatheaceae (incl. Alsophilaceae, Hymenophyllopsidaceae): Alsophilia, Cyathea, Gymnosphaera, Hymenophyllopsis, Sphaeropteris
Family Dicksoniaceae (incl. Lophosoriaceae): Calochlaena, Dicksonia, Lophosoria
Family Metaxyaceae: Metaxya
Incertae Sedis: Amphidesmium, Cnemidaria, Lophosoria, Trichosorus
ORDER POLYPODIALES
Family Lindsaeaceae (incl. Cystodiaceae, Lonchitidaceae): Cystodium, Lindsaea, Lonchitis, Odontosoria, Ormoloma, Sphenomeris, Tapeinidium, Xyropteris
Family Saccolomataceae: Saccoloma
Family Dennstaedtiaceae (incl. Hypolepidaceae, Monachosoraceae, Pteridiaceae): Blotiella, Coptidipteris, Costaricia, Dennstaedtia, Histiopteris, Hypolepis, Leptolepia, Microlepia, Monachosorum, Oenotrichia, Paesia, Pteridium
Family Pteridaceae (incl. Acrostichaceae, Actiniopteridaceae, Adiantaceae, Anopteraceae, Antrophyaceae, Ceratopteridaceae, Cheilanthaceae, Cryptogrammaceae, Hemionitidaceae, Negripteridaceae, Parkeriaceae,Platyzomataceae, Sinopteridaceae, Taenitidaceae, Vittariaceae): Achrostichum, Actinopteris, Adiantopsis, Adiantum, Aleuropteris, Afropteris, Anathacorus, Anetium, Anogramma, Anopteris, Antrophyum, Argyrochosma, Aspidotis, Astrolepis, Austrogramme, Bommeria, Botryogramma, Cassebeera, Ceratopteris, Cerosora, Cheilanthes, Cheiloplecton, Chrysochosma, Chrysodium, Coniogramme, Cryptogramma, Dictyogramme, Doryopteris, Eriosorus, Haplopteris, Hecistopteris, Hemionitis, Hemionanthes, Holcochlaena, Holttumia, Holttumiella, Jamesonia, Llavea, Mildella, Monogramma, Negripteris, Nephropteris, Neurocallis, Neurosorus, Notholaena, Notogramme, Ochropteris, Onychium, Ormopteris, Parahemionitis, Pellea, Pentagramma, Pityrogramma, Pityromeria, Platytaenia, Platyzoma, Polytaenium, Pteris, Pterozonium, Radiovittaria, Rheopteris, Schizolepton, Scoliosorus, Syngramma, Taenitis, Trachypteris, Vaginularia, Vittaria.
Family Aspleniaceae: Asplenium, Boniniella, Camptosorus, Loxoscaphe, Pleurosorus, Phyllitis, Ceterach, Thamnopteris, Antigramma, Holodictyum, Schaffneria, Sinephropteris, Hymenasplenium
Family Thelypteridaceae: Thelypteris, Cyclosorus, Ampeltopteris, Chingia, Amphineuron, Cyclogramma, Christella, Cyclosorus, Glaphyropteridopteris, Goniopteris, Meniscium, Menisorus, Mesophlebion, Pelazoneuron, Plesioneuron, , Pneumatopteris, Pronephrium, Pseudocyclosorus, Sphaerostephanos, Stegnogramma, Steiropteris, Trigonospora, Phegopteris, Pseudophegopteris, Macrothelypteris, Amauropelta, Coryphopteris, Metathelypteris, Oreopteris, Parathelypteris
Family Woodsiaceae (incl. Athyriaceae, Cystopteridaceae): Athyrium, Diplazium, Callipteris, Monomelangium, Ancystopteris, Cheilanthopsis, Cornopteris, Cystopteris, Deparia, Lunathyrium, Dryoathyrium, Athyriopsis, Dictyodroma, Diplaziopsis, Gymnocarpium, Currania, Hemidictyum, Homalosorus, Protowoodsia, Pseudocystopteris, Hymenocystis, Rhachidosorus, Woodsia
Family Blechnaceae (incl. Stenochlaenaceae): Blechnum, Brainea, Doodia, Pteridoblechnum, Sadleria, Salpichlena, Steenisioblechnum, Stenochlaena, Woodwardia, Anchystia, Cheiniopteris, Lorinseria
Family Onocleaceae: Matteuccia, Onoclea, Onocleopsis, Pentarhizidium
Family Dryopteridaceae (incl. Aspidiaceae, Bolbitidaceae, Elaphoglossaceae, Hypodematiaceae, Peranemataceae): Ctenitis, Dryopteris, Elaphoglossum, Polystichum, Acrophorus, Acrorumohra, Adenoderris, Arachnoides, Altaxipteris, Bolbitis, Egenolfia, Coveniella, Cyclodium, Cyrtogonellum, Cyrtomidictyum, Cyrtomium, Didymochlaena, Dryopolystichum, Dryopsis, Nothoperanema, Microstaphyla, Peltapteris, Hypodematium, Lastreopsis, Leucostegia, Lithostegia, Lomogramma, Maxonia, Megalastrum, Oenotrichia, Olfersia, Peranema, Phanerophlebia, Papuapteris, Plecosorus, Polybotrya, Polystichopteris, Revwattsia, Stenolepia, Stigmatopteris, Teratophyllum
Family Lomariopsidaceae (incl. Nephrolepidaceae): Cyclopeltis, Lomariopsis, Nephrolepis, Thysansoria
Family Tectariaceae: Aenigmopteris, Arthropteris, Heterogonium, Hypoderris, Pleocnemia, Psammiosorus, Psomiocarpa, Pteridrys, Tectaria, Triplophyllum
Family Oleandraceae: Oleandra
Family Davalliaceae: Araiostega, Davallia, Humata, Parasorus, Scyphularia, Davallodes, Pachypleuria
Family Polypodiaceae (incl. Drynariaceae, Grammitidaceae, Gymnogrammitidaceae, Loxogrammaceae, Platyceriaceae, Pleurisoriopsidaceae): Acrosorus, Adenophorus, Aenigmopteris, Aglaomorpha, Anathropteris, Arthromeris, Arthopteris, Atalopteris, Belvisia, Calymmodon, Campyloneurum, Ceradenia, Chlamydogramme, Christiopteris, Chrysogrammitis, Cochlidium, Colysis, Crypsinus, Ctenitis, Ctenitopsis, Ctenopteris, Cyclopeltis, Dicranoglossum, Dictymia, Dictyoxiphium, Drymotaenium, Drynaria, Dryopsis, Enterosora, Goniophlebium, Grammitis, Gymnogrammitis, Lastreopsis, Lecanopteris, Lellingeria, Lemmaphyllum, Lepidosorus, Leptochilus, Leucostegia, Loxogramme, Marginariopsis, Megalastrum, Melpomene, Microgramma, Microphlebodium, Micropolypodium, Microsorum, Nephrolepis, Neurodium, Niphidium, Paraselliguea, Pecluma, Phlebodium, Phymatosorus, Platycerium, Pleocnemia, Pleopeltis, Pleurosoriopsis, Podosorus, Polypodiodes, Polypodiopteris, Polypodium, Prosaptia, Psammiosorus, Pseudocolysis, Pteridrys, Pyrrosia, Rumohra, Scleroglossum, Selliguea, Solanopteris, Tectaria, Tectaridium, Terpsichore, Themelium, Triplophyllum, Trogostolon, Xiphopteris, Zygophlebia
Incertae Sedis: Acyatopteris, Anisocampium, Chrynephrium, Cornoathyrium, Craspedosorus, Diellia, Diplora, Kuniwatsukia, Rhachidosorus, Macrothelypteris, Nannothelypteris, Phanerophlebiopsis, Sorolepidium
CLASS PROGYMNOSPERMOPSIDA
ORDER ANEUROPHYTALES
Aneurophyton, Tetraxylopteris.
ORDER PROTOPITYALES
Protopitys
ORDER ARCHAEOPTERIDALES
Archaeopteris [Callixophyton, Svalbardia, Actinoxylon, Siderella, Actinopodium]
This page is maintained by Jack R. Holt & Carlos A. Iudica. Last modified: 03/25/08 .