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cystoptère ténue, Mackay's brittle fern, Mackay's fragile fern, upland brittle bladderfern

bulb bladderfern, bulblet bladder fern, bulblet fragile fern, cystoptère bulbifère

Stems

creeping, not cordlike, internodes short, beset with old petiole bases, hairs absent;

scales tan to light brown, lanceolate, radial walls thin, luminae tan.

creeping, not cordlike, internodes very short, less than 0.5 cm, heavily beset with old petiole bases, hairs absent;

scales uniformly brown to somewhat clathrate, lanceolate, radial walls brown, luminae clear.

Leaves

monomorphic, clustered at stem apex, to 40 cm, nearly all bearing sori.

monomorphic, clustered at stem apex, to 75 cm, seasonally bearing sori (earliest leaves lack sori, subsequent leaves with sori).

Petiole

dark at base, mostly green to straw-colored distally, shorter than or nearly equaling blade, base sparsely scaly.

reddish when young, usually green or straw-colored throughout (occasionally darker) in mature specimens, shorter than blade, base sparsely scaly.

Blade

lanceolate to narrowly elliptic, 1(–2)-pinnate-pinnatifid, widest at or just below middle, apex short-attenuate;

rachis and costae lacking gland-tipped hairs or bulblets;

axils of pinnae lacking multicellular, gland-tipped hairs.

broadly to narrowly deltate, 2-pinnate to 2-pinnate-pinnatifid, almost always widest at base, apex long-attenuate;

rachis and costae usually densely covered by unicellular, gland-tipped hairs, often with bulblets;

axils of pinnae occasionally with multicellular, gland-tipped hairs.

Pinnae

typically at acute angle to rachis, often curving toward blade apex, margins crenulate;

proximal pinnae pinnatifid to pinnate-pinnatifid, ± equilateral, basiscopic pinnules not enlarged;

basal basiscopic pinnules sessile, base cuneate to obtuse, distal pinnae ovate to narrowly elliptic.

mostly perpendicular to rachis, not curving toward blade apex, margins serrate;

proximal pinnae pinnate-pinnatifid to pinnatifid, ± equilateral, basiscopic pinnules not enlarged, basal basiscopic pinnules sessile to short-stalked, bases truncate to obtuse;

distal pinnae ovate to oblong.

Veins

directed into teeth and notches.

directed into notches.

Indusia

ovate to cup-shaped, without gland-tipped hairs.

cup-shaped, apex truncate, typically invested with unicellular, gland-tipped hairs.

Spores

spiny, usually 39–50 µm. 2n = 168.

spiny, usually 33–38 µm. 2n = 84.

Cystopteris tenuis

Cystopteris bulbifera

Phenology Sporulating summer–fall. Sporulating summer–fall.
Habitat Mostly on shaded rock and cliff faces but also occasionally on forest floors Cracks and ledges on cliffs, rarely terrestrial, usually on calcareous substrates
Elevation 0–2800 m (0–9200 ft) 0–2500 m (0–8200 ft)
Distribution
from FNA
AR; AZ; CT; DE; IA; IL; IN; KS; KY; MA; MD; ME; MI; MN; MO; NC; NE; NH; NJ; NV; NY; OH; OK; PA; RI; TN; UT; VA; VT; WI; WV; NB; NS; ON; QC
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from FNA
AL; AR; AZ; CT; DE; GA; IA; IL; IN; KY; MA; MD; ME; MI; MN; MO; NC; NH; NJ; NM; NY; OH; PA; RI; SC; TN; TX; UT; VA; VT; WI; WV; NB; NF; NS; ON; QC
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Discussion

Long recognized as Cystopteris fragilis var. mackayi, C. tenuis was returned to species status by R. C. Moran (1983b). It is probably an allotetraploid originating from C. protrusa and an extinct diploid related to C. fragilis (C. H. Haufler 1985; C. H. Haufler and M. D. Windham 1991).

Cystopteris tenuis is common in eastern North America and less frequent at the northern and western perimeter of its range. In the center of its distribution (Minnesota, Iowa, Illinois, Wisconsin, Indiana, Ohio, Pennsylvania), the narrow, elliptic pinnae angled toward the blade apex and the rounded teeth make C. tenuis relatively distinct from C. fragilis and C. protrusa (although the early season, sterile leaves of C. protrusa often resemble those of C. tenuis). In the west and especially in the northeast, C. tenuis and C. fragilis are difficult to distinguish. For the most part, C. fragilis is confined to higher latitudes and elevations than C. tenuis, but the two species can be sympatric and occasionally form sterile tetraploid hybrids. Cystopteris protrusa and C. tenuis are infrequently sympatric, but where they are, sterile triploid hybrids can occur. Hybrids between C. tenuis and C. tennesseensis are recognized as C. × wagneri (R. C. Moran 1983). Hybridization between C. tenuis and C. bulbifera has also been reported (R. C. Moran 1982b). This hybrid, C. × illinoensis R. C. Moran, is known only from the type and needs to be studied further.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Cystopteris bulbifera usually occurs on moist calcareous cliffs, but it also grows on rock in dense woods and occasionally occurs terrestrially in northern swamps. Blades on most individuals are narrowly deltate and distinctively long-attenuate. The rachises, costae, and indusia are densely beset with gland-tipped, unicellular hairs. Mature specimens often have deciduous bulblets. These characteristics readily distinguish C. bulbifera from other diploid species.

Hybridization and allopolyploidy involving Cystopteris bulbifera and other North American Cystopteris species have generated several species. In the eastern portion of its range, C. bulbifera and C. protrusa are the diploid progenitors of the tetraploid C. tennesseensis (C. H. Haufler et al. 1990). In northeastern North America, C. bulbifera has hybridized with tetraploid C. fragilis, ultimately resulting in the hexaploid C. laurentiana (R. F. Blasdell 1963). In the southwest, the diploid C. reevesiana and disjunct representatives of C. bulbifera are the progenitors of the tetraploid C. utahensis (C. H. Haufler and M. D. Windham 1991). In addition to these fertile allopolyploids, sterile hybrids are also possible when C. bulbifera is sympatric with its polyploid derivatives. Sterile hybrids between C. bulbifera and C. tennesseensis have been identified from several localities. Cystopteris bulbifera may hybridize with C. tenuis to form C. × illinoensis (C. H. Haufler et al. 1990; R. C. Moran 1982b). Diploid sexual C. bulbifera may be distinguished from these allopolyploid species and sterile hybrids because the hybrid-derived taxa (1) will normally have less prominent glandular hairs, (2) will have misshapen bulblets, (3) will more likely have blades that are widest above the base, and (4) will have large spores (in sexual allopolyploids) or malformed spores (in sterile hybrids).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Source FNA vol. 2. FNA vol. 2.
Parent taxa Dryopteridaceae > Cystopteris Dryopteridaceae > Cystopteris
Sibling taxa
C. bulbifera, C. fragilis, C. laurentiana, C. montana, C. protrusa, C. reevesiana, C. tennesseensis, C. utahensis
C. fragilis, C. laurentiana, C. montana, C. protrusa, C. reevesiana, C. tennesseensis, C. tenuis, C. utahensis
Synonyms Nephrodium tenue, C. fragilis var. mackayi Polypodium bulbiferum
Name authority (Michaux) Desvaux (Linnaeus) Bernhardi: Neues J. Bot. 1(2): 10. (1805)
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