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doradille des murailles, wall rue, wall-rue spleenwort

spleenwort

Roots

not proliferous.

fibrous, not proliferous or proliferous and producing tiny plantlets.

Stems

short-creeping to erect, often branched;

scales very dark brown throughout, narrowly deltate, 1–3 × 0.1–0.25 mm, margins with widely spaced teeth.

erect, rarely long-creeping;

scales basally attached, clathrate.

Leaves

monomorphic.

Petiole(s)

reddish brown proximally, green distally, dull, 1–9 cm, (1/2–)1–2 times length of blade;

indument of dark brown, narrowly deltate scales proximally grading into multicellular hairs.

not articulate.

Blade(s)

deltate-ovate to obovate or oblanceolate, 1–2(–3)-pinnate to 2-pinnate-pinnatifid, 2–6 × 1–4 cm, somewhat thick, glabrous;

base obtuse;

apex acute to rounded, not rooting.

1–4-pinnate, of diverse size and shape.

Pinnae

in 2–4 pairs, deltate-ovate to obdeltate;

proximal (largest) pinnae 7–30 × 5–20 mm;

base truncate to acute;

margins finely (sometimes coarsely) incised;

apex rounded to acute.

Veins

free, evident.

Indusia

present.

Sori

as many as 30 or more per pinna, usually 1–5 per segment, on both basiscopic and acroscopic sides.

Spores

64 per sporangium.

Rachis

green, dull, glabrous except for very sparse, minute hairs.

x

= 36.

2n

= 144.

Asplenium ruta-muraria

Asplenium

Habitat Limestone (or calcareous shale) cliffs and boulders, rarely invading masonry
Elevation 0–1000 m (0–3300 ft)
Distribution
from FNA
AL; AR; CT; IN; KY; MA; MD; MI; MO; NC; NH; NJ; NY; OH; PA; RI; TN; VA; VT; WV; ON; QC; Europe; e Asia
[WildflowerSearch map]
[BONAP county map]
from USDA
Worldwide
[BONAP county map]
Discussion

The relationship of North American Asplenium ruta-muraria to its European counterparts is incompletely understood and bears further investigation. Based on features of the stems, M. L. Fernald (1928) segregated the North American taxon as A. cryptolepis, but most current authors agree that morphologic differentiation of North American and European material is too slight and inconsistent for recognition at the specific level. In Europe, two ploidy levels are treated as subspecies, diploid A. ruta-muraria subsp. dolomiticum Lovis & Reichstein and tetraploid A. ruta-muraria subsp. ruta-muraria, the latter representing the most compelling case for true autopolyploidy (i.e., based on chromosomal homology) known in ferns (G. Vida 1970). Chromosome counts of North American plants are consistently tetraploid; whether or not these plants are referable to subsp. ruta-muraria will remain unclear until additional evidence (e.g., isozymes) is obtained. Meanwhile, North American material should be designated simply as A. ruta-muraria, the convention used in most current manuals.

Although M. L. Fernald (1928) recognized Asplenium cryptolepis var. ohionis (= A. ruta-muraria var. subtenuifolium Christ), based on its acute rather than rounded segment apices, leaves assignable to this variety may occur on plants also bearing leaves more similar to those of the type variety (R. Cranfill 1980). The former is not recognized taxonomically here.

Numerous hybrids of Asplenium ruta-muraria with various taxa are known from Europe (T. Reichstein 1981), but only three are known from North America, all exceedingly rare. These are the hybrids with A. rhizophyllum (A. × inexpectatum), with A. trichomanes (A. × clermontae Syme), and with A. platyneuron (A. × morganii).

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

Species ca. 700 (28 species, 3 nothospecies in the flora).

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

Key
1. Blades simple, pinnatifid, or forked (sometimes with free pinnae at base), not pinnate throughout.
→ 2
1. Blades pinnate throughout or pinnatifid only in distal 1/3, pinnae undivided to 1–3-divided.
→ 8
2. Blades less than 2–3 mm wide, linear, frequently forking or with 1–3 small, narrow projections.
A. septentrionale
2. Blades more than 10 mm wide, linear-lanceolate to lanceolate, if lobed, segments resembling adnate pinnae.
→ 3
3. Veins anastomosing to form areoles; blades rooting at tip.
A. rhizophyllum
3. Veins free; blades with apical buds or not but not rooting at tip.
→ 4
4. Blades simple, linear to oblanceolate.
→ 5
4. Blades pinnatifid, elliptic, lanceolate, to deltate.
→ 6
5. Blades cordate at base; roots not proliferous.
scolopendrium var. americanum
5. Blades tapered at base; roots proliferous.
A. serratum
6. Petioles 1 cm or less, dull; stem scales 0.6–1 mm wide, sparsely denticulate.
A. dalhousiae
6. Petioles 1–10 cm, lustrous; stem scales less than 0.5 mm wide, entire.
→ 7
7. Petioles dark-pigmented only at base; blade narrowly deltate.
A. pinnatifidum
7. Petioles and proximal midrib darkly pigmented abaxially; blade oblong- lanceolate.
A. ebenoides
8. Blades 2–4-pinnate, with only largest pinnae divided to all pinnae divided.
→ 9
8. Blades 1-pinnate, pinnae undivided (except in some individuals of A. pumilum).
→ 18
9. Blades deltate-ovate, with 2–5 pairs of pinnae; pinnae generally deltate-ovate to obdeltate, rarely lanceolate to narrowly elliptic; color in living plants dull bluish green.
A. ruta-muraria
9. Blades ovate-lanceolate to linear, with 5–20 pairs of pinnae; pinnae linear to oblong; color in living plants shiny green.
→ 10
10. Delicate root proliferations usually present (commonly buried in mossy soil in crevices, but enabling clone formation); damp, shaded limestone; Florida.
→ 11
10. Delicate root proliferations absent or rare; mainly rock crevices; plants not in Florida.
→ 15
11. Spores uniform and normal.
→ 12
11. Spores of different sizes and shapes, malformed (irregular in size and shape).
→ 13
12. Blades oblong-lanceolate, mostly 2- pinnate; pinnae lanceolate to oblong- lanceolate.
A. cristatum
12. Blades narrowly lanceolate, 2–3- pinnate; pinnae ovate to ovate- deltate.
A. verecundum
13. Blades linear-lanceolate, s Florida.
A. ×biscayneanum
13. Blades oblong-lanceolate, c Florida.
→ 14
14. Medial pinnae pinnate throughout; pinnules mostly deeply notched apically.
A. ×curtissii
14. Medial pinnae pinnate only in proximal 1/2; pinnules mostly shallowly notched apically.
A. plenum
15. Buds at blade, pinna, and pinnule apices minute, scaly, dormant; Arizona.
A. exiguum
15. Buds absent; foothills and mountains in e United States and s Rocky Mountains.
→ 16
16. Petioles all pale green except at base.
A. montanum
16. Petioles dark reddish brown.
→ 17
17. Blades lanceolate to linear- lanceolate, 1–2-pinnate; proximal pinnae often somewhat reduced; e United States foothills and mountains.
A. bradleyi
17. Blades deltate-ovate to deltate- lanceolate, 2–3-pinnate; proximal pinnae usually largest; s Rocky Mountains.
A. adiantum-nigrum
18. Pinna pairs 1–2(–4); blades ovate-deltate, with fine nonglandular hairs on veins.
A. pumilum
18. Pinna pairs 5–35; blades mainly oblong to lanceolate, usually with glandular hairs or narrow hairlike scales on veins or blades glabrous.
→ 19
19. Pinnae with conspicuous basal auricles overlapping rachis; sori inframedial; leaves ± dimorphic, upright and tall, or spreading and short; plants often terrestrial.
A. platyneuron
19. Pinnae bases not overlapping rachis; sori medial to supramedial; leaves monomorphic; plants mainly on rock (epiphytic in A. auritum).
→ 20
20. Blades mostly 4–9 cm wide, linear- lanceolate to oblong-lanceolate.
→ 21
20. Blades mostly less than 3 cm wide, mainly linear.
→ 22
21. Blades linear-lanceolate, to 35 cm; pinna pairs 10–22, base with conspicuous acroscopic auricle; epiphytic on tree trunks.
A. auritum
21. Blades oblong to oblong-lanceolate, to 20 cm; pinna pairs 4–8, base with or without small acroscopic auricle; on mossy limestone.
A. abscissum
22. Sori 1(–3), mainly confined to basiscopic side of strongly asymmetric pinnae.
A. monanthes
22. Sori 4–10, on both sides of costa of nearly symmetric pinnae.
→ 23
23. Blade apex prolonged into whiplike extension of rachis terminating in bud.
A. palmeri
23. Blade apex not prolonged, terminal bud absent.
→ 24
24. Pinnae ovate, oblong-ovate, to rhombic, mainly 4–7 mm, length 1–2 times width.
→ 25
24. Pinnae oblong-lanceolate to oblong, mainly 6–10 mm, length 3–5 times width.
→ 28
25. Rachises dark reddish brown throughout.
A. trichomanes
25. Rachises dark in proximal 1/3–2/3 or only at base.
→ 26
26. Rachises dark in proximal 1/3–2/3; pinna margins entire to shallowly crenate.
A. adulterinum
26. Rachises dark only at base; pinna margins dentate-crenate.
→ 27
27. Pinnae with basiscopic 1/2 much reduced; blades mainly less than 1 cm wide; n North America.
A. trichomanes-ramosum
27. Pinnae nearly symmetric; blades mainly more than 1 cm wide; s Florida.
A. trichomanes-dentatum
28. Pinna margins deeply lobed, especially acroscopic and distal margins, cut 1/3 or more distance from margin to costa; s California.
A. vespertinum
28. Pinna margins nearly entire or shallowly crenate- dentate; se United States to e Arizona.
→ 29
29. Pinna margins nearly entire; sori supramedial; spores 32; widespread in s United States.
A. resiliens
29. Pinna margins ± shallowly crenate- dentate; sori medial to inframedial; spores 32 or 64; local in se United States.
→ 30
30. Pinna margins shallowly crenulate; auricle present at pinna base; veins obscure; spores 32 per sporangium.
A. ×heteroresiliens
30. Pinna margins serrate to crenate; auricle rudimentary or absent at pinna base; veins evident; spores 64 per sporangium.
A. heterochroum
Source FNA vol. 2. FNA vol. 2. Authors: Warren H. Wagner Jr., Robbin C. Moran, Charles R. Werth.
Parent taxa Aspleniaceae > Asplenium Aspleniaceae
Sibling taxa
A. abscissum, A. adiantum-nigrum, 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. 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
A. abscissum, A. adiantum-nigrum, 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 var. americanum, A. septentrionale, A. serratum, A. trichomanes, A. trichomanes-dentatum, A. trichomanes-ramosum, A. verecundum, A. vespertinum, A. ×biscayneanum, A. ×curtissii, A. ×heteroresiliens
Synonyms A. cryptolepis, A. cryptolepis var. ohionis, A. ruta-muraria var. cryptolepis
Name authority Linnaeus: Sp. Pl. 2: 1081. (1753) Linnaeus: Sp. Pl. 2: 1078. 1753; Gen. Pl. ed. 5, 485, (1754)
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