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Leaf evolution in the subclass Ophioglossidae -Rhyniales: 1. Aglaophyton major, sporophyte (a); male gametophyte Lyonophyton rhyniensis (b); 2. Stockmansella remyi; 3. Teruelia diezii; 4. Huvenia kleui; 5. Rhynia gwynne-vaughanii, sporophyte (a), gametophyte Remyophyton delicatum (b); 6. Horneophyton lignieri, sporophyte (a), female gametophyte Langiophyton mackiei (b), sporangium/synangium with columella (c). Modified Kidston, Lang (1921) (1a, 5a); Remy et al. (1993) (1b, 6b); Taylor et al. (2009) (1b); Schultka, Hass (1997) (2); Cascales-Miñana, Gerrienne (2017) (3); Hass, Remy (1991) (4); Kerp et al. (2010) (5b); Eggert (1974) (6a, c). Author: M. Hrabovský.

Leaf evolution in the subclass Ophioglossidae -Rhyniales: 1. Aglaophyton major, sporophyte (a); male gametophyte Lyonophyton rhyniensis (b); 2. Stockmansella remyi; 3. Teruelia diezii; 4. Huvenia kleui; 5. Rhynia gwynne-vaughanii, sporophyte (a), gametophyte Remyophyton delicatum (b); 6. Horneophyton lignieri, sporophyte (a), female gametophyte Langiophyton mackiei (b), sporangium/synangium with columella (c). Modified Kidston, Lang (1921) (1a, 5a); Remy et al. (1993) (1b, 6b); Taylor et al. (2009) (1b); Schultka, Hass (1997) (2); Cascales-Miñana, Gerrienne (2017) (3); Hass, Remy (1991) (4); Kerp et al. (2010) (5b); Eggert (1974) (6a, c). Author: M. Hrabovský.

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The possible relationships between microphyllous and megaphyllous leaves were presented in the previous part of this series. Leaves could be comprehended as single or branched microphylls or as compound systems of microphylls. The primitive forms of such compound systems, called appendages, led to the proposal of various phylogenetic trees based on...

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Context 1
... (Rhyniales) share several morphological characteristics with Renaliales and Cooksoniales. The sporangia on 'lateral branches' occurring in Huvenia (Hass, Remy 1991; Fig. 5.4), in fact dichotomized microphylls, are reduced to single sporangia ( Fig. 5.2) in Stockmansella (Schultka, Hass 1997), 'adventitious branches' in Rhynia (Kidston, Lang 1917;Lemoigne 1970; Fig. 5.5) or protuberances present almost in all genera ( Fig. 5.1-2,4-5). The genera with absent lateral sporangia (Rhynia or Teruelia) have one ...
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... several morphological characteristics with Renaliales and Cooksoniales. The sporangia on 'lateral branches' occurring in Huvenia (Hass, Remy 1991; Fig. 5.4), in fact dichotomized microphylls, are reduced to single sporangia ( Fig. 5.2) in Stockmansella (Schultka, Hass 1997), 'adventitious branches' in Rhynia (Kidston, Lang 1917;Lemoigne 1970; Fig. 5.5) or protuberances present almost in all genera ( Fig. 5.1-2,4-5). The genera with absent lateral sporangia (Rhynia or Teruelia) have one terminal sporangium (Kidston, Lang 1917;Lemoigne 1970;Cascales-Miñana, Gerrienne 2017;Fig. ...
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... and Cooksoniales. The sporangia on 'lateral branches' occurring in Huvenia (Hass, Remy 1991; Fig. 5.4), in fact dichotomized microphylls, are reduced to single sporangia ( Fig. 5.2) in Stockmansella (Schultka, Hass 1997), 'adventitious branches' in Rhynia (Kidston, Lang 1917;Lemoigne 1970; Fig. 5.5) or protuberances present almost in all genera ( Fig. 5.1-2,4-5). The genera with absent lateral sporangia (Rhynia or Teruelia) have one terminal sporangium (Kidston, Lang 1917;Lemoigne 1970;Cascales-Miñana, Gerrienne 2017;Fig. ...
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... was also questioned by Edwards (2003) or Kadereit, Bresinsky (2013). There are more serious morphological features than water-conducting cells, above all the gametophyte known as Lyonophyton rhyniensis (Fig. 5.1b) that is related to other rhynialean gametophytes (Remy, Remy 1980; Kerp et al. 2004;Taylor et al. 2005). Another genus Horneophyton (Fig. 5.6a) was also excluded from Rhyniales by Kenrick, Crane (1997) as an oldest lineage of polysporangiophytes due to unique sporangia (Eggert 1974). Doweld (2001) classified this genus even among bryophytes due to presence of columella in branched sporangia (Fig. 5.6c). However, its vascular gametophyte Langiophyton mackiei (Fig. 5.6b) ...
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... (Remy, Remy 1980; Kerp et al. 2004;Taylor et al. 2005). Another genus Horneophyton (Fig. 5.6a) was also excluded from Rhyniales by Kenrick, Crane (1997) as an oldest lineage of polysporangiophytes due to unique sporangia (Eggert 1974). Doweld (2001) classified this genus even among bryophytes due to presence of columella in branched sporangia (Fig. 5.6c). However, its vascular gametophyte Langiophyton mackiei (Fig. 5.6b) resembles the gametophytes of other Rhyniales (Remy, Hass 1991). The reclassification of Horneophyton into Rhyniales and Ophioglossidae allows to describe the homology of the unique sporangia with columella from a new perspective. The formation of synangia is known ...
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... genus Horneophyton (Fig. 5.6a) was also excluded from Rhyniales by Kenrick, Crane (1997) as an oldest lineage of polysporangiophytes due to unique sporangia (Eggert 1974). Doweld (2001) classified this genus even among bryophytes due to presence of columella in branched sporangia (Fig. 5.6c). However, its vascular gametophyte Langiophyton mackiei (Fig. 5.6b) resembles the gametophytes of other Rhyniales (Remy, Hass 1991). The reclassification of Horneophyton into Rhyniales and Ophioglossidae allows to describe the homology of the unique sporangia with columella from a new perspective. The formation of synangia is known from both Psilotales and Ophioglossales. The compound synangia of ...
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... not associated with sporophylls and microphyll reduction (Kidston, Lang 1920;Gensel et al. 1969 as Lycopodiidae. More advanced Asteroxylon (Fig. 6.3), Hueberia, Kaulangiophyton (Fig. 6.4) lack venation in microphylls (Kidston, Lang 1920;1921;Gensel et al. 1969;Kerp et al. 2003;Yang et al. 2009) hence are misinterpreted as enations. Nothia aphylla (Fig. 6.5a) covered by many emergences could be also related to Asteroxylon (Daviero- Gomez et al. 2005). Its male gametophyte described as Kidstonophyton discoides (Fig. 6.5b) resembles the gametophytes of Rhyniales ( Remy et al. ...
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... (Fig. 6.4) lack venation in microphylls (Kidston, Lang 1920;1921;Gensel et al. 1969;Kerp et al. 2003;Yang et al. 2009) hence are misinterpreted as enations. Nothia aphylla (Fig. 6.5a) covered by many emergences could be also related to Asteroxylon (Daviero- Gomez et al. 2005). Its male gametophyte described as Kidstonophyton discoides (Fig. 6.5b) resembles the gametophytes of Rhyniales ( Remy et al. ...

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