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Fendler's sandwort

eremogone, sandwort

Habit Plants ± cespitose, bluish green, not glaucous, with woody base. Plants perennial, often densely matted, usually with branched, woody base.
Taproots

slender or usually stout;

rhizomes absent.

Stems

erect, (2–)10–30(–40) cm, stipitate-glandular.

prostrate (nonflowering stems) or ascending to erect (flowering stems), simple or branched, terete.

Leaves

basal leaves persistent;

cauline leaves in (4–)5+ pairs, reduced or not;

basal blades ascending or recurved, filiform, 1–10(–11) cm × 0.2–0.4 mm, flexuous, herbaceous, apex apiculate to spinose, glabrous to puberulent, not glaucous.

usually congested at or near base of flowering stems, usually connate, mostly sessile;

blade 1-veined, needlelike or filiform to subulate or narrowly linear, succulent or not, apex blunt to usually apiculate or spinose.

Inflorescences

(1–)3–35-flowered, ± open cymes.

terminal, open to congested or umbellate cymes or sometimes flowers solitary;

bracts paired (or clustered at summit of peduncle in some E. congesta varieties), reduced, foliaceous or scarious.

Pedicels

3–25 mm, stipitate-glandular.

erect or flowers sessile.

Flowers

sepals weakly to prominently 1–3-veined, linear-lanceolate, 4–7.5 mm, not enlarging in fruit, margins broad, apex acuminate, moderately to densely stipitate-glandular on herbaceous portion;

petals white, oblong-elliptic to spatulate, 4–8 mm, 0.9–1.3 times as long as sepals, apex entire to somewhat erose;

nectaries as lateral and abaxial rounding of base of filaments opposite sepals, 0.2 × 0.4 mm.

perianth and androecium weakly perigynous;

hypanthium shallowly cup-shaped;

sepals 5, distinct or barely connate proximally, green (sometimes purplish in E. capillaris and E. eastwoodiae), linear-lanceolate to ovate, 1.8–12 mm, margins white, scarious, apex obtuse or rounded to acute, acuminate, or spinose, not hooded;

petals 5, white, yellowish white, or occasionally pink or brownish, clawed or not, blade apex entire, erose, emarginate, or rarely 2-fid;

nectaries usually 5, prominent at (or adjacent to in E. eastwoodiae) base of filaments opposite sepals, rarely absent;

stamens 10, arising from hypanthium;

filaments distinct;

staminodes absent;

styles 3, filiform, 2.5–3 mm, glabrous proximally;

stigmas 3, subcapitate, smooth to papillate (50x).

Capsules

5–7 mm, glabrous.

ovoid to urceolate, opening by 6 ascending to recurved teeth;

carpophore present or usually absent.

Seeds

black, ovoid to pyriform with hilar notch, 1.5–1.9 mm, tuberculate;

tubercles rounded, elongate to rounded-conic.

1–10, dark reddish or greenish brown, tan, blackish purple, black, or gray, ovoid to pyriform or suborbicular, laterally compressed, smooth, rugulose, or tuberculate, marginal wing absent, appendage absent.

x

= 11.

2n

= 44.

Eremogone fendleri

Eremogone

Phenology Flowering spring–late summer.
Habitat Sagebrush plains, pine forests, and mountain slopes to alpine zones
Elevation 1200-4300 m (3900-14100 ft)
Distribution
from FNA
AZ; CO; NM; TX; UT; WY
[WildflowerSearch map]
[BONAP county map]
from USDA
North-temperate regions; esp w North America; Eurasian mountains; Asia Minor
[BONAP county map]
Discussion

We agree with M. F. Baad (1969) in not formally recognizing varieties within Eremogone fendleri. B. Maguire (1947, 1951) recognized five varieties, defined chiefly on leaf and sepal characteristics. While some specimens can be “matched” to varieties, many appear intermediate between them, forming a continuum of variation.

B. Maguire (1947) noted that Eremogone fendleri is “probably to be found in the states of Sonora and Chihuahua, Mexico”; we have not seen any collections from that area.

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

Species ca. 89 (14 in the flora).

Much of this treatment follows those by B. Maguire (1947, 1951) and M. F. Baad (1969) of Arenaria sect. Eremogone. The Eremogone “group” is morphologically distinctive (cespitose or matted, woody perennials with filiform to subulate leaves, stiffly erect to ascending flowering stems, and open to congested or umbellate cymes) and nearly all of the published chromosome counts are based on x = 11 (see also Baad), an uncommon base number in Arenaria in the narrow sense. The decision to recognize Eremogone, as used in W. A. Weber and R. C. Wittmann (1992) and R. D. Dorn (2001), is based largely on results of the molecular study of subfamily Alsinoideae by M. Nepokroeff et al. (2001). In that study, Arenaria (excluding Minuartia and Moehringia) was shown to be polyphyletic, with two distinct clades. One of the clades consists of seven species: five from Arenaria subg. Eremogone and two from the closely related subg. Eremogoneastrum; therefore the Eremogone group, as delimited by Maguire, is monophyletic.

While Eremogone is a distinct genus, there is considerable morphological variability among the species, with many, often geographically isolated, morphological types; see J. C. Hickman (1971) for a commentary on the situation, numerous examples of intermediate forms, and notes on their possible evolution. With this in mind, we have chosen not to adopt an infrageneric classification; J. McNeill’s (1962) outline of Arenaria suggests that three species groups could be recognized within Eremogone.

Because of the aforementioned variability and intermediacy within and between taxa, the production of a satisfactory key has been a challenge and, as with those by B. Maguire (1947, 1951), likely will prove inadequate for at least a low percentage of specimens.

Reports of winged seeds in Eremogone possibly are derived from examination of immature material. The tubercles along the abaxial edge that have not yet expanded can appear as a lighter-colored band around a portion of the seed.

The morphology of the nectaries in Eremogone is very diverse, both among species and, in one case, among infraspecific taxa (E. macradenia). We feel that a morphogenetic study using scanning electron microscopy similar to the investigation of nectaries in Schiedea by W. L. Wagner and E. M. Harris (2000) may shed light on relationships within Eremogone.

Some suggestions for the identification of taxa of Eremogone follow.

In determining the shape of the sepals, several flowers should be evaluated and the membranous margins must be included, not just the central herbaceous portion. As this genus is in many ways a “problematic group,” some effort beyond what may be considered “normal” may be needed for accurate identification. This is because of the frequent need to observe floral details at high magnification (20 or 30×), preferably with a dissecting microscope. Furthermore, on dried specimens, a flower or two may need to be “boiled up” or otherwise rehydrated so that good, clear observations can be made. This is a practice that too often is avoided, to the detriment of accurate observation and identification, and should be standard in the study of pressed material.

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

Key
1. Inflorescences usually dense or ± open, capitate, umbellate cymes
→ 2
1. Inflorescences ± open, rarely compact cymes
→ 4
2. Petals 1.5-2 times as long as sepals; sepals 3-6.5 mm, ovate to lanceolate
E. congesta
2. Petals 0.8-1.1 times as long as sepals; sepals (5-)6-12 mm, lanceolate or usually linear-lanceolate
→ 3
3. Stems glabrous, with 6-10 pairs of well-developed cauline leaves, leaves markedly overlapping
E. franklinii
3. Stems scabrid-puberulent, with 1-4 pairs of well-developed leaves or reduced distally, usually little overlapping
E. hookeri
4. Sepal apices obtuse to rounded or barely acute, sometimes abruptly mucronate
→ 5
4. Sepal apices acute to acuminate, sometimes spinose
→ 10
5. Basal leaves abundant, blade (2-)4-8 cm, straight to curved in one direction, flexuous, apex acute or acuminate to weakly spinose; valves of capsule at least sparsely stipitate-glandular
E. capillaris
5. Basal leaves abundant or sparse, blade 0.3-2(-4) cm, recurved in various directions, apex often stiff and spinose; capsules glabrous
→ 6
6. Basal leaf blades ascending or often arcuate-spreading, rigid, herbaceous; plants glaucous or not
→ 7
6. Basal leaf blades erect to spreading, flexuous or rigid, herbaceous or ± succulent; plants not glaucous
→ 8
7. Sepals ± stipitate-glandular; plants glaucous; cauline leaves in 1-3 pairs; petals 1.5-3 times as long as sepals
E. aculeata
7. Sepals glabrous or nearly so; plants not glaucous; cauline leaves in 5-7 pairs; petals 1.3-1.5 times as long as sepals
E. aberrans
8. Sepals stipitate-glandular; basal leaf blades ± succulent, ± flexuous; Crater Lake and vicinity, sw Oregon
E. pumicola
8. Sepals glabrous or nearly so; basal leaf blades herbaceous, rigid
→ 9
9. Sepals 1.8-3 mm in flower; nectaries neither grooved nor cupped, 0.3-0.4 mm; s California
E. ursina
9. Sepals 3-5(-6) mm in flower; nectaries with transverse groove or elongate cup, 0.6 mm; se Idaho, n Utah, Wyoming
E. kingii
10. Sepal apices broadly acute, sometimes spinose
→ 11
10. Sepal apices narrowly acute to acuminate or spinose
→ 14
11. Stems usually 20-40(-100) cm; leaf blades usually 2-6 mm; basal leaves sparse or absent; nectaries rectangular, 2-lobed and 0.7-1.5 mm, or narrowly longitudinally rectangular, truncate, densely minutely pubescent with erect to spreading hairs, 0.7-0.8 mm, or laterally or abaxially rounded, 0.3-0.4 mm
→ 12
11. Stems usually 3-20 cm; leaf blades usually 0.3-3 cm; basal leaves abundant; nectaries rounded, 0.2-0.6 mm
→ 13
12. Sepals 3-4.3 mm, in fruit to 5.5 cm; cymes open, branches spreading; nectaries as lateral and abaxial rounding of filament bases, 0.3-0.4 mm
E. ferrisiae
12. Sepals 4.5-7.2 mm, in fruit to 8 mm; cymes ± compact, branches ascending or erect; nectaries rectangular, 2-lobed or truncate, 0.7-1.5 mm
E. macradenia
13. Cauline leaves in 5-7 pairs, reduced distally; petals yellowish white, 5.8-10 mm; capsules 7-10 mm; n Arizona
E. aberrans
13. Cauline leaves in (1-)2-4+ pairs, reduced distally or not; petals white or rarely pink, (3-)4-7 mm; capsules 4.5-7 mm; Great Basin and to n and w
E. kingii
14. Sepals glabrous throughout or essentially so
→ 15
14. Sepals moderately to densely stipitate-glandular
→ 16
15. Petals 0.9-1.1 times as long as sepals; nectaries longitudinally rectangular, apically cleft or emarginate, 1-2 mm; Arizona, Colorado, New Mexico,Utah, Wyoming
E. eastwoodiae
15. Petals 1.5-1.7 times as long as sepals; nectaries rounded at base of filaments, 0.3-0.4 mm; se Nevada
E. stenomeres
16. Cauline leaves usually in 2-4 pairs, blade usually 0.3-3 cm; stems usually 2-20 cm; nectaries rounded lobe with transverse groove or elongate cup; Colorado Plateau and Great Basin
E. kingii
16. Cauline leaves in (4-)5+ pairs, blade usually 3-11 cm; stems usually 7-40 cm; nectaries rounded, not transversely grooved or cupped
→ 17
17. Leaves flexuous; petals 0.9-13 times as long as sepals; Rocky Mountains proper and outlying mountains
E. fendleri
17. Leaves rigid; petals 1.5-1.7 times as long as sepals; se Nevada
E. kingii
Source FNA vol. 5, p. 64. FNA vol. 5, p. 56. Authors: Ronald L. Hartman, Richard K. Rabeler, Frederick H. Utech.
Parent taxa Caryophyllaceae > subfam. Alsinoideae > Eremogone Caryophyllaceae > subfam. Alsinoideae
Sibling taxa
E. aberrans, E. aculeata, E. capillaris, E. congesta, E. eastwoodiae, E. ferrisiae, E. franklinii, E. hookeri, E. kingii, E. macradenia, E. pumicola, E. stenomeres, E. ursina
Subordinate taxa
E. aberrans, E. aculeata, E. capillaris, E. congesta, E. eastwoodiae, E. fendleri, E. ferrisiae, E. franklinii, E. hookeri, E. kingii, E. macradenia, E. pumicola, E. stenomeres, E. ursina
Synonyms Arenaria fendleri, Arenaria fendleri subsp. brevifolia, Arenaria fendleri var. brevifolia, Arenaria fendleri var. diffusa, Arenaria fendleri var. porteri, Arenaria fendleri var. tweedyi, Arenaria tweedyi
Name authority (A. Gray) Ikonnikov: Novosti Syst. Vyssh. Rast. 10: 139. (1973) Fenzl: Vers. Darstell. Alsin., 13, unnumbered plate. (1833)
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