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black spleenwort

spleenwort family

Habit Plants terrestrial, on rock, or rarely epiphytic.
Roots

not proliferous.

Stems

ascending or short-creeping, infrequently branched;

scales dark brown to blackish throughout, narrowly deltate, 2–4(–5) × 0.2–0.5 mm, margins entire or shallowly denticulate to serrulate.

erect or nearly erect, rarely long-creeping, scaly.

Leaves

monomorphic.

monomorphic, rarely almost dimorphic with fertile leaves taller and more erect than sterile ones.

Petiole(s)

dark reddish brown proximally, often fading to green distally, lustrous, 2–20 cm, 2/3–2 times length of blade;

indument of black filiform scales and minute hairs.

with 1 vascular bundle X-shaped in cross section or with 2 vascular bundles back to back and C-shaped.

Blade(s)

deltate, 2–3-pinnate, 2.5–10 × 2–6.5 cm, thick, hairs dark, scattered, minute;

base truncate;

apex acute to acuminate, not rooting.

extremely diverse, simple to 4-pinnate, commonly with tiny glandular hairs and a few linear scales, rarely with spreading hairs.

Pinnae

in 4–10 pairs, deltate to lanceolate;

most proximal (largest) pinnae 1.5–4 × 1–2.5 cm;

base obliquely obtuse;

segment margins coarsely incised;

apex acute.

Veins

free, evident.

free to anastomosing.

Indusia

usually present, shape conforming to sorus and originating along 1 side of sorus.

Sori

1–numerous pairs per pinna [1–6 pairs per segment], on both basiscopic and acroscopic sides.

borne on veins, ± lunate to linear.

Sporangia

with stalk of 1 row of cells, annulus vertical, interrupted by sporangial stalk.

Spores

64 per sporangium.

monolete;

perispore typically winged, spiny, reticulate, or perforate.

Gametophytes

surficial, green, cordate.

Rachis

greenish throughout or sometimes reddish brown proximally, lustrous, sparsely pubescent.

Steles

radially symmetric or dorsiventral (with structurally distinct abaxial and adaxial aspects) dictyosteles.

2n

= 144.

Asplenium adiantum-nigrum

Aspleniaceae

Habitat Cliffs
Elevation 1675–2300 m (5500–7500 ft)
Distribution
from FNA
AZ; CO; UT; Eurasia; Africa
[BONAP county map]
Worldwide
[BONAP county map]
Discussion

Asplenium adiantum-nigrum is principally a Eurasian species and occurs extremely rarely in North America (see M. G. Shivas 1969 and M. D. Windham 1983 for a discussion of the conspecificity of Western Hemisphere and Eastern Hemisphere material). It is an allotetraploid derived from hybridization of two European taxa, A. cuneifolium Viviani and A. onopteris Linnaeus (M. G. Shivas 1969). Hybrids involving A. adiantum-nigrum and other Asplenium species occur in Europe but are unknown in North America.

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

Members of this family can usually be identified by the combination of clathrate stem scales and indusiate linear sori. Supporting anatomic characteristics include the two vascular bundles in the petiole that unite distally in the petiole to form an X-shaped petiolar strand, and the single row of cells in the sporangial stalk. The scales consist of cells with dark, thick, radial walls and clear, thin, tangential walls, giving the scales a clathrate (latticelike) appearance reminiscent of lead moldings between plates of stained glass.

As construed here, Aspleniaceae comprise a single, huge, extremely diverse genus, Asplenium. A satisfactory taxonomic division into subgenera or satellite genera has not been possible because of the absence of any significant gaps. Various segregates have been proposed (e.g., Camptosorus, Phyllitis, Ceterach, Pleurosorus), but numerous "intergeneric" hybrids occur.

The members of Asplenium are popular with plant evolutionists, field naturalists, and fern gardeners, not only because of the interesting morphology of the plants but also because of their remarkable ability to form spectacular hybrids, often combining dramatically different leaf shapes. In North America, 23 diploid hybrids and allopolyploids have been recorded. At least two of these hybrid combinations occur as both sterile diploids and their fertile allotetraploid derivatives. Only those hybrids that are reproductively competent (through vigorous clone-forming by root proliferations or apogamy, or rarely through sexual reproduction) are treated in the key and fully described below.

Only about two-fifths of the reproductively competent species are believed to be cladistically divergent species; the other three-fifths are of hybrid origin (allopolyploids). For two of the allotetraploids, sterile diploids of the same parentage are also known. The most unusual allopolyploid phytogeographically is Asplenium adiantum-nigrum, the parents of which are known only in the Old World. These reticulate relationships are summarized in the reticulogram.

Polyploidy is widespread in Asplenium, and the chromosome numbers vary from 2 x to 6 x. Two species, Asplenium trichomanes and A. heterochroum, occur in different levels of polyploidy—2 x and 4 x, and 4 x and 6 x, respectively. The highest chromosome number known for Asplenium in North America is 2n = 216 (in A. trichomanes-dentatum and the hexaploid form of A. heterochroum). The only three apogamous taxa are A. monanthes (3 x), A. resiliens (3 x), and A. × heteroresiliens (5 x).

Genera 1, species ca. 700 (1 genus, 28 species, and 3 nothospecies in the flora).

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

Source FNA vol. 2. FNA vol. 2, p. 228. Authors: Warren H. Wagner Jr., Robbin C. Moran, Charles R. Werth.
Parent taxa Aspleniaceae > Asplenium
Sibling taxa
A. abscissum, A. adulterinum, A. auritum, A. bradleyi, A. cristatum, A. dalhousiae, A. ebenoides, A. exiguum, A. heterochroum, A. monanthes, A. montanum, A. palmeri, A. pinnatifidum, A. platyneuron, A. plenum, A. pumilum, A. resiliens, A. rhizophyllum, A. ruta-muraria, A. scolopendrium, A. septentrionale, A. serratum, A. trichomanes, A. trichomanes-dentatum, A. trichomanes-ramosum, A. verecundum, A. vespertinum, A. ×biscayneanum, A. ×curtissii, A. ×heteroresiliens
Subordinate taxa
Synonyms A. andrewsii, A. chihuahuense, A. dubiosum
Name authority Linnaeus: Sp. Pl. 2: 1081. (1753) Newman
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