Difference between revisions of "Notholaena"

R. Brown

Prodr. 145. 1810.

Common names: Cloak fern
Etymology: Greek notho, false, and chlaena, coat, in reference to the reflexed leaf segment margins that form false indusia
Treatment appears in FNA Volume 2.
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--><span class="statement" id="st-d0_s0" data-properties=""><b>Plants </b>usually on rock.</span> <span class="statement" id="st-d0_s1" data-properties="stem growth form or orientation;stem architecture or arrangement;stem orientation;stem orientation;stem orientation;stem architecture"><b>Stems </b>short-creeping to compact, ascending to horizontal, usually branched;</span> <span class="statement" id="st-d0_s2" data-properties="scale coloration;scale coloration;scale shape;scale shape;scale shape;central stripe coloration;margin coloration">scales black or often bicolored with dark central stripe and lighter margins, linear-subulate to lanceolate;</span> <span class="statement" id="st-d0_s3" data-properties="margin shape;margin shape;margin shape;margin shape;margin shape">margins ciliate, denticulate, or entire.</span> <span class="statement" id="st-d0_s4" data-properties="leaf architecture;leaf arrangement or growth form;leaf some measurement"><b>Leaves </b>monomorphic, clustered, 4–35 cm.</span> <span class="statement" id="st-d0_s5" data-properties="petiole coloration;petiole coloration;petiole shape;petiole shape;groove count;groove dehiscence or orientation;gland pubescence"><b>Petiole </b>brown or black, usually rounded, flattened, or with single longitudinal groove adaxially, often bearing scales, hairs, or farinose glands, with single vascular-bundle.</span> <span class="statement" id="st-d0_s6" data-properties="blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade shape;blade texture;blade architecture or function or pubescence;blade reflectance;blade coloration or pubescence or relief;farina coloration;farina coloration"><b>Blade </b>linear-lanceolate, ovate, deltate, or pentagonal, pinnate-pinnatifid to 4-pinnate, leathery, abaxially covered by yellowish or whitish farina (completely obscured by stellate scales in Notholaena aschenborniana), adaxially often sparsely glandular, dull, not striate;</span> <span class="statement" id="st-d0_s7" data-properties="rachis course">rachis straight.</span> <span class="statement" id="st-d0_s8" data-properties="ultimate segment architecture;ultimate segment architecture;ultimate segment architecture;ultimate segment fusion;ultimate segment shape;ultimate segment shape;ultimate segment shape;ultimate segment shape;ultimate segment width"><b>Ultimate </b>segments of blade sessile to subsessile, often adnate to costae, narrowly elliptic to oblong-ovate or deltate, usually less than 4 mm wide;</span> <span class="statement" id="st-d0_s9" data-properties="base shape;base shape;base shape;base shape">base rounded to truncate or cuneate;</span> <span class="statement" id="st-d0_s10" data-properties="stalk reflectance;stalk coloration">stalks (when present) usually lustrous and dark-colored;</span> <span class="statement" id="st-d0_s11" data-properties="segment margin orientation;segment margin arrangement;false indusium prominence">segment margins recurved to form confluent, poorly defined false indusia extending entire length of segment.</span> <span class="statement" id="st-d0_s12" data-properties="vein fusion;vein prominence;vein architecture;vein arrangement"><b>Veins </b>of ultimate segments free, obscure, pinnately branched and divergent distally.</span> <span class="statement" id="st-d0_s13" data-properties="false indusium coloration;false indusium size or width;false indusium position;sporangium position"><b>False </b>indusia greenish, narrow, clearly marginal, occasionally concealing the sporangia.</span> <span class="statement" id="st-d0_s14" data-properties="spore count;spore count;spore count"><b>Sporangia </b>confined to submarginal vein tips, containing 64, 32, or 16 spores, intermixed with farina-producing glands.</span> <span class="statement" id="st-d0_s15" data-properties="spore coloration;spore coloration;spore coloration;spore shape;spore shape;spore texture;equatorial-ridge count;equatorial-ridge prominence"><b>Spores </b>black to dark-brown, globose or tetrahedral-globose, granulate, lacking prominent equatorial-ridge.</span> <span class="statement" id="st-d0_s16" data-properties=""><b>Gametophytes </b>glandular-farinose (gametophytes of all other Pteridaceae genera lack farina).</span> <span class="statement" id="st-d0_s17" data-properties="gametophyte pubescence;x chromosome count">x = 30.</span><!--
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--><span class="statement" id="st-undefined" data-properties=""><b>Plants </b>usually on rock. <b>Stems</b> short-creeping to compact, ascending to horizontal, usually branched; scales black or often bicolored with dark central stripe and lighter margins, linear-subulate to lanceolate; margins ciliate, denticulate, or entire. <b>Leaves</b> monomorphic, clustered, 4–35 cm. <b>Petiole</b> brown or black, usually rounded, flattened, or with single longitudinal groove adaxially, often bearing scales, hairs, or farinose glands, with single vascular bundle. <b>Blade</b> linear-lanceolate, ovate, deltate, or pentagonal, pinnate-pinnatifid to 4-pinnate, leathery, abaxially covered by yellowish or whitish farina (completely obscured by stellate scales in <i>Notholaena aschenborniana</i>), adaxially often sparsely glandular, dull, not striate; rachis straight. <b>Ultimate</b> segments of blade sessile to subsessile, often adnate to costae, narrowly elliptic to oblong-ovate or deltate, usually less than 4 mm wide; base rounded to truncate or cuneate; stalks (when present) usually lustrous and dark-colored; segment margins recurved to form confluent, poorly defined false indusia extending entire length of segment. <b>Veins</b> of ultimate segments free, obscure, pinnately branched and divergent distally. <b>False</b> indusia greenish, narrow, clearly marginal, occasionally concealing the sporangia. <b>Sporangia</b> confined to submarginal vein tips, containing 64, 32, or 16 spores, intermixed with farina-producing glands. <b>Spores</b> black to dark brown, globose or tetrahedral-globose, granulate, lacking prominent equatorial ridge. <b>Gametophytes</b> glandular-farinose (gametophytes of all other Pteridaceae genera lack farina). <b>x</b> = 30.</span><!--
  
 
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|distribution=North America;Mexico;West Indies;Central America;South America.
 
|distribution=North America;Mexico;West Indies;Central America;South America.
|discussion=<p>As pointed out by R. M. Tryon (1956), J. T. Mickel (1979), and many others, Notholaena in the broad sense is poorly defined and difficult to distinguish from either Cheilanthes or Pellaea. North American taxa traditionally assigned to Notholaena represent at least four distinct evolutionary lineages (M. D. Windham 1986). In order to clarify species relationships and generic boundaries among cheilanthoid ferns, Notholaena is defined here in a very restricted sense. The pubescent, nonfarinose species (such as N. newberryi) have been placed in Cheilanthes following R. M. Tryon and A. F. Tryon (1982). The scaly, nonfarinose taxa of the N. sinuata complex have been transferred to Astrolepis for reasons discussed by D. M. Benham and M. D. Windham (1992). The glabrous and farinose species related to Pellaea (e.g., N. jonesii and N. dealbata) have been placed in Argyrochosma (M. D. Windham 1987). The species retained in Notholaena following this reorganization (i.e., members of the farinose N. grayi -- N. standleyi alliance) form a coherent, monophyletic group found only in the Western Hemisphere. The correct generic name for this group is in dispute because Notholaena has been lectotypified by several authors citing three different type species. The rules of priority favor the first typification (by J. Smith in 1875) based on N. trichomanoides (Linnaeus) Desvaux, which is definitely a member of the group here called Notholaena. R. E. G. Pichi-Sermolli (1989), however, urged acceptance of the typification by C. Christensen ([1905–]1906). Both the second and third lectotypifications of Notholaena were based on species unrelated to the North American taxa discussed here. If Smith's typification is overturned, the correct generic name for our species will be Chrysochosma (J. Smith) Kümmerle.</p><!--
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|discussion=<p>As pointed out by R. M. Tryon (1956), J. T. Mickel (1979), and many others, <i>Notholaena</i> in the broad sense is poorly defined and difficult to distinguish from either <i>Cheilanthes</i> or <i>Pellaea</i>. North American taxa traditionally assigned to <i>Notholaena</i> represent at least four distinct evolutionary lineages (M. D. Windham 1986). In order to clarify species relationships and generic boundaries among cheilanthoid ferns, <i>Notholaena</i> is defined here in a very restricted sense. The pubescent, nonfarinose species (such as N. newberryi) have been placed in <i>Cheilanthes</i> following R. M. Tryon and A. F. Tryon (1982). The scaly, nonfarinose taxa of the N. sinuata complex have been transferred to <i>Astrolepis</i> for reasons discussed by D. M. Benham and M. D. Windham (1992). The glabrous and farinose species related to <i>Pellaea</i> (e.g., N. jonesii and N. dealbata) have been placed in <i>Argyrochosma</i> (M. D. Windham 1987). The species retained in <i>Notholaena</i> following this reorganization (i.e., members of the farinose <i>N. grayi</i> -- <i>N. standleyi</i> alliance) form a coherent, monophyletic group found only in the Western Hemisphere. The correct generic name for this group is in dispute because <i>Notholaena</i> has been lectotypified by several authors citing three different type species. The rules of priority favor the first typification (by J. Smith in 1875) based on N. trichomanoides (Linnaeus) Desvaux, which is definitely a member of the group here called <i>Notholaena</i>. R. E. G. Pichi-Sermolli (1989), however, urged acceptance of the typification by C. Christensen ([1905–]1906). Both the second and third lectotypifications of <i>Notholaena</i> were based on species unrelated to the North American taxa discussed here. If Smith's typification is overturned, the correct generic name for our species will be Chrysochosma (J. Smith) Kümmerle.</p><!--
 
--><p>Species ca. 25 (10 in the flora).</p>
 
--><p>Species ca. 25 (10 in the flora).</p>
 
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|publication year=1810
 
|publication year=1810
 
|special status=
 
|special status=
|source xml=https://jpend@bitbucket.org/aafc-mbb/fna-fine-grained-xml.git/src/287ef3db526bd807d435a3c7423ef2df1e951227/V2/V2_771.xml
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|source xml=https://bitbucket.org/aafc-mbb/fna-data-curation/src/2e0870ddd59836b60bcf96646a41e87ea5a5943a/coarse_grained_fna_xml/V2/V2_771.xml
 
|genus=Notholaena
 
|genus=Notholaena
|base shape=rounded;truncate or cuneate
 
|blade architecture or function or pubescence=glandular
 
|blade coloration or pubescence or relief=not striate
 
|blade reflectance=dull
 
|blade shape=pentagonal pinnate-pinnatifid;4-pinnate
 
|blade texture=leathery
 
|central stripe coloration=dark
 
|equatorial-ridge count=lacking
 
|equatorial-ridge prominence=prominent
 
|false indusium coloration=greenish
 
|false indusium position=marginal
 
|false indusium prominence=defined
 
|false indusium size or width=narrow
 
|farina coloration=whitish;yellowish
 
|gametophyte pubescence=glandular-farinose
 
|gland pubescence=farinose
 
|groove count=single
 
|groove dehiscence or orientation=longitudinal
 
|leaf architecture=monomorphic
 
|leaf arrangement or growth form=clustered
 
|leaf some measurement=4cm;35cm
 
|margin coloration=lighter
 
|margin shape=entire;denticulate;entire;denticulate;ciliate
 
|petiole coloration=black;brown
 
|petiole shape=flattened;rounded
 
|rachis course=straight
 
|scale coloration=bicolored;black
 
|scale shape=linear-subulate;lanceolate
 
|segment margin arrangement=confluent
 
|segment margin orientation=recurved
 
|sporangium position=concealing
 
|spore coloration=black;dark-brown
 
|spore count=16;32;64
 
|spore shape=tetrahedral-globose;globose
 
|spore texture=granulate
 
|stalk coloration=dark-colored
 
|stalk reflectance=lustrous
 
|stem architecture=branched
 
|stem architecture or arrangement=compact
 
|stem growth form or orientation=short-creeping
 
|stem orientation=ascending;horizontal
 
|ultimate segment architecture=sessile;subsessile
 
|ultimate segment fusion=adnate
 
|ultimate segment shape=narrowly elliptic;oblong-ovate or deltate
 
|ultimate segment width=0mm;4mm
 
|vein architecture=branched
 
|vein arrangement=divergent
 
|vein fusion=free
 
|vein prominence=obscure
 
|x chromosome count=30
 
 
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Latest revision as of 20:25, 5 November 2020

Plants usually on rock. Stems short-creeping to compact, ascending to horizontal, usually branched; scales black or often bicolored with dark central stripe and lighter margins, linear-subulate to lanceolate; margins ciliate, denticulate, or entire. Leaves monomorphic, clustered, 4–35 cm. Petiole brown or black, usually rounded, flattened, or with single longitudinal groove adaxially, often bearing scales, hairs, or farinose glands, with single vascular bundle. Blade linear-lanceolate, ovate, deltate, or pentagonal, pinnate-pinnatifid to 4-pinnate, leathery, abaxially covered by yellowish or whitish farina (completely obscured by stellate scales in Notholaena aschenborniana), adaxially often sparsely glandular, dull, not striate; rachis straight. Ultimate segments of blade sessile to subsessile, often adnate to costae, narrowly elliptic to oblong-ovate or deltate, usually less than 4 mm wide; base rounded to truncate or cuneate; stalks (when present) usually lustrous and dark-colored; segment margins recurved to form confluent, poorly defined false indusia extending entire length of segment. Veins of ultimate segments free, obscure, pinnately branched and divergent distally. False indusia greenish, narrow, clearly marginal, occasionally concealing the sporangia. Sporangia confined to submarginal vein tips, containing 64, 32, or 16 spores, intermixed with farina-producing glands. Spores black to dark brown, globose or tetrahedral-globose, granulate, lacking prominent equatorial ridge. Gametophytes glandular-farinose (gametophytes of all other Pteridaceae genera lack farina). x = 30.

Distribution

North America, Mexico, West Indies, Central America, South America.

Discussion

As pointed out by R. M. Tryon (1956), J. T. Mickel (1979), and many others, Notholaena in the broad sense is poorly defined and difficult to distinguish from either Cheilanthes or Pellaea. North American taxa traditionally assigned to Notholaena represent at least four distinct evolutionary lineages (M. D. Windham 1986). In order to clarify species relationships and generic boundaries among cheilanthoid ferns, Notholaena is defined here in a very restricted sense. The pubescent, nonfarinose species (such as N. newberryi) have been placed in Cheilanthes following R. M. Tryon and A. F. Tryon (1982). The scaly, nonfarinose taxa of the N. sinuata complex have been transferred to Astrolepis for reasons discussed by D. M. Benham and M. D. Windham (1992). The glabrous and farinose species related to Pellaea (e.g., N. jonesii and N. dealbata) have been placed in Argyrochosma (M. D. Windham 1987). The species retained in Notholaena following this reorganization (i.e., members of the farinose N. grayi -- N. standleyi alliance) form a coherent, monophyletic group found only in the Western Hemisphere. The correct generic name for this group is in dispute because Notholaena has been lectotypified by several authors citing three different type species. The rules of priority favor the first typification (by J. Smith in 1875) based on N. trichomanoides (Linnaeus) Desvaux, which is definitely a member of the group here called Notholaena. R. E. G. Pichi-Sermolli (1989), however, urged acceptance of the typification by C. Christensen ([1905–]1906). Both the second and third lectotypifications of Notholaena were based on species unrelated to the North American taxa discussed here. If Smith's typification is overturned, the correct generic name for our species will be Chrysochosma (J. Smith) Kümmerle.

Species ca. 25 (10 in the flora).

Key

1 Blades with scales or multicelled hairs in addition to whitish or yellowish farina. > 2
1 Blades with whitish or yellowish farina only, lacking scales or multicelled hairs. > 5
2 Nonfarinose indument on blades primarily of deeply dissected, stellate scales; farina on abaxial surface completely concealed by overlapping scales. Notholaena aschenborniana
2 Nonfarinose indument on blades of lanceolate scales and/or multicelled hairs; farina on abaxial surface readily apparent. > 3
3 Nonfarinose indument on blades of shiny, dark brown, needlelike hairs; sporangia containing 64 spores. Notholaena nealleyi
3 Nonfarinose indument on blades primarily of dull, light brown, narrowly lanceolate scales; sporangia containing 16 or 32 spores. > 4
4 Blades sparsely villous adaxially with long, multicelled, whitish hairs; abaxial scales ciliate. Notholaena aliena
4 Blades glabrous adaxially except for scattered glandular hairs; abaxial scales entire. Notholaena grayi
5 Petioles grooved or flattened adaxially; blades linear to narrowly deltate, 2–6 times longer than wide; basal pinnae not markedly larger than adjacent pair, ± equilateral, proximal basiscopic pinnules not greatly enlarged. > 6
5 Petioles rounded adaxially; blades pentagonal to ovate, 1–2 times longer than wide; basal pinnae much larger than adjacent pair, strongly inequilateral due to enlargement of proximal basiscopic pinnules. > 7
6 Blades glabrous adaxially at maturity; ultimate segments adnate to costae for entire width and clearly sessile; segment margins slightly recurved. Notholaena lemmonii
6 Blades glandular adaxially at maturity; at least some ultimate segments constricted proximally and slightly stalked; segment margins strongly revolute. Notholaena greggii
7 Blades clearly 3-pinnate proximally; at least some ultimate segments constricted proximally and slightly stalked. > 8
7 Blades pinnate-pinnatifid to 2-pinnate-pinnatifid; ultimate segments adnate to costae their entire width and clearly sessile. > 9
8 Stem scales weakly bicolored, brown margins very narrow; blades broadly pentagonal, adaxially glandular; segment margins recurved but not strongly revolute. Notholaena californica
8 Stem scales strongly bicolored, brown margins broad and conspicuous; blades narrowly deltate-pentagonal, adaxially glabrescent; segment margins strongly revolute. Notholaena neglecta
9 Blades ovate, fully pinnate beyond basal pinnae, distal pinnae not connected; abaxial farina white. Notholaena copelandii
9 Blades broadly pentagonal, pinnatifid beyond basal pinnae, distal pinnae connected by narrow wing of green tissue; abaxial farina yellow. Notholaena standleyi
... more about "Notholaena"
Michael D. Windham +
R. Brown +
Cloak fern +
North America +, Mexico +, West Indies +, Central America +  and South America. +
Greek notho, false, and chlaena, coat, in reference to the reflexed leaf segment margins that form false indusia +
tryon1956a +  and windham1986a +
Notholaena +
Pteridaceae +