Eremogone

Plants perennial, often densely matted, usually with branched, woody base. Taproots slender or usually stout; rhizomes absent. Stems prostrate (nonflowering stems) or ascending to erect (flowering stems), simple or branched, terete. Leaves 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 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 erect or flowers sessile. Flowers: 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 (50×). Capsules ovoid to urceolate, opening by 6 ascending to recurved teeth; carpophore present or usually absent. Seeds 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.

Distribution

North-temperate regions, esp. w North America, Eurasian mountains, Asia Minor.

Discussion

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.

Selected References

Baad, M. F. 1969. Biosystematic Studies of the North American Species of Arenaria, Subgenus Eremogone (Caryophyllaceae). Ph.D. dissertation. University of Washington.

Hickman, J. C. 1971. Arenaria, section Eremogone (Caryophyllaceae) in the Pacific Northwest: A key and discussion. Madroño 21: 201–207.

Ikonnikov, S. S. 1973. Zametki o gvozdichnykh (Caryophyllaceae). I. O rode Eremogone Fenzl. Novosti Sist. Vyssh. Rast. 10: 136–142.

Maguire, B. 1947. Studies in the Caryophyllaceae. III. A synopsis of the North American species of Arenaria sect. Eremogone Fenzl. Bull. Torrey Bot. Club 74: 38–56.

Lower Taxa

Eremogone aberrans, Eremogone aculeata, Eremogone capillaris, Eremogone congesta, Eremogone eastwoodiae, Eremogone fendleri, Eremogone ferrisiae, Eremogone franklinii, Eremogone hookeri, Eremogone kingii, Eremogone macradenia, Eremogone pumicola, Eremogone stenomeres, Eremogone ursina

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	Eremogone 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	Eremogone franklinii
3	Stems scabrid-puberulent, with 1-4 pairs of well-developed leaves or reduced distally, usually little overlapping	Eremogone 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	Eremogone 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	Eremogone 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	Eremogone aberrans
8	Sepals stipitate-glandular; basal leaf blades ± succulent, ± flexuous; Crater Lake and vicinity, sw Oregon	Eremogone 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	Eremogone ursina
9	Sepals 3-5(-6) mm in flower; nectaries with transverse groove or elongate cup, 0.6 mm; se Idaho, n Utah, Wyoming	Eremogone 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	Eremogone 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	Eremogone macradenia
13	Cauline leaves in 5-7 pairs, reduced distally; petals yellowish white, 5.8-10 mm; capsules 7-10 mm; n Arizona	Eremogone 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	Eremogone 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	Eremogone 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	Eremogone 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	Eremogone 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	Eremogone fendleri
17	Leaves rigid; petals 1.5-1.7 times as long as sepals; se Nevada	Eremogone kingii