Aliciella cliffordii J.M.Porter, 2011

Porter, J. Mark, 2011, Two new Aliciella species and a new subspecies in Ipomopsis (Polemoniaceae) from the western United States of America, Phytotaxa 15, pp. 15-25 : 19-20

Aliciella cliffordii J.M.Porter , sp. nov. ( Fig. 2 )

Species Aliciella cliffordii similis Aliciella haydenii (A. Gray) J.M. Porter , sed corolla caerulea vel alba differt.

Type:— J.M. Porter , K.D.Heil, & A.Clifford 13528; U.S.A. Arizona. Apache County: Navajo Indian Reservation, ca. 5 km S of Cove; 36° 32’ 01” N, 109° 13’ 04” W, 2225 m elev., 6 May 2003. (holotype: RSA; isotypes: ARIZ, SJNM) .

Taprooted, short-lived perennial, with a well-developed basal rosette of leaves, thus appearing biennial, 10–90 cm tall, stems sparsely and coarsely glandular pubescent with uniseriate glandular trichomes, erect but freely and divaricately branching, sometimes to the base, the lateral branches overtopping the primary axis. Basal leaves forming a rosette, entire, coarsely toothed to once-pinnatifid, 1.2–5 cm long, the rachis broad, 1– 4.5 mm wide, the segments 6–15, entire to rarely lobed, glandular and crisp puberulent with white, uniseriate eglandular trichomes, lobes cuspidate or mucronate. Lower cauline leaves pinnatifid to more commonly entire and linear, gradually to abruptly reduced in size upward, ultimately upper cauline leaves entire, 1–18 mm long, glandular puberulent. Inflorescence loosely open cymose-panicle, the flowers mostly crowded at the tips of the branches. Calyx cylindrical to campanulate, 3–4.7 mm long, tube 1.8–3.5 mm long at anthesis, glandular. Corolla 6–18 mm long, blue, paling to nearly white, corolla externally glabrous but bearing a patch of glandular trichomes centrally, at the proximal end (base) of the corolla lobe, narrowly funnelformsalverform; the tube, 5.5–12 mm long, lobes narrowly oblanceolate, 3–5.5(–6.0) mm long, 1.9–3.5 mm wide. Stamens equally inserted in the upper tube (at the sinuses of the corolla lobes), the free portion ca. 1 mm long, anthers 1.5–2.2 mm long, slightly exserted. Style well exserted and approaching herkogamous to included and reverse herkogamous. Fruit a capsule, 3-5 mm long, 2.3–2.9 mm wide, ovoid, dehiscing at the apex but valves remaining attached at the base, locules 3. Seed 1.8–2.2 mm long, 0.7–1 mm wide, narrowly ovoid to fusiform, or angled-truncate and lung-shaped, wingless or with a narrow partial wing at one end or along the margin, less than 0.2 mm wide, pale gray-brown.

Distribution:— Aliciella cliffordii is apparently restricted to Apache County, Arizona and adjacent San Juan County, New Mexico, occurring in sandy or clay badlands, associated with pinyon-juniper woodland and ponderosa pine forests. Soils are generally red. Occurrences range in elev. from 1525–1980 m (5000–6500 ft). A very localized species, it is endemic to the Navajo Nation on Beautiful Mountain and the adjacent eastern slopes of the Lukachukai and Chuska Mountains.

Phenology:—Flowering begins in May and continues through June or rarely to early July. Fruiting begins in mid-May and lasts through July.

Etymology:—The specific epithet, cliffordii , honors Arnold Clifford, enthusiastic field botanist and important collector in the Four Corners region and Navajo Nation.

Additional specimens examined (paratypes):— U.S.A. ARIZONA. APACHE COUNTY: 1 to 3 mi S of Cove, in the Chuska Mtns. (label states San Juan Co., NM, incorrectly), 8 June 1974, D.Atwood 6361 ( BRY, NY) ; Navajo Nation, W of Red Rock, on the eastern foot of Lukachukai Mtns , red silty loam, 14 May 1991, J.M. Porter 9559 ( RSA) ; Navajo Nation, drainage and road to Mexican Cry Mesa, on lower member of the Wingate Formation , 27 Sept. 2001, K.Heil & A.Clifford 18375 ( RSA, SJNM) . NEW MEXICO. SAN JUAN COUNTY: 12 mi west of Shiprock along Hwy 504, 15 May 1970, D.Atwood 2525 ( BRY, NY) .

Discussion:— Aliciella Brand sect. Aliciella subsect. Subnudae J.M.Porter ( Porter 1998: 31; see also Porter and Johnson 2000) is largely of the Colorado Plateau. It boasts some of the more colorful and attractive members of the Phlox family in the Four Corners region, including A. formosa (Greene ex Brand 1907: 119) J.M. Porter (1998: 33) , A. subnuda ( Gray 1870: 276) J.M. Porter (1998: 33) , and A. haydenii . The showy magenta flowers of A. formosa and A. haydenii are hawkmoth pollinated, with over 90% of insect visits due to hawkmoths (Porter 1993; Porter and Floyd-Hanna 1993). There are two subspecies of A. haydenii recognized, differing in corolla size, density of glands on the external corolla, and in part corolla color upon drying. Regardless of these differences, populations and the two races of A. haydenii share a common reproductive and pollination system (Porter 1993) and show some intergradation. This is consistent with the hypothesis that they represent a single species.

The series of isolated and disjunct populations near Beautiful Mountain, Arizona are very similar to Aliciella haydenii , but differ in their narrower corolla tube and lobes and pale floral color, having blue to nearly white corollas ( Fig. 2A, B ). The populations in the Beautiful Mountain area are pollinated primarily by beeflies (Bombilius lancifer Osten Sacken [1877: 251]; Fig. 2C ), and visited, to a lesser degree, by anthophorid bees, representing an ethological distinction from A. haydenii . Field studies over a four-year period, summing to nearly 200 hours of observation time, have verified that these populations are not visited by hawkmoths, rather B. lancifer (73% of visits) and several species of anthophorid bees (18% of visits). Nonpollinating visitors (e.g., syrphid flies) account for the remaining visits. I hypothesize that the populations near Beautiful Mountain correspond to a different reproductive and pollination system from A. haydenii , and therefore comprise a genetically independent species. In accordance, I treat it at the rank of species. While A. cliffordii seems technically weak if considering the morphological features alone, the different pollination system and geographic disjunction is consistent with a hypothesis of peripatric speciation, involving a change in pollination mechanisms. This provides an explicit hypothesis that can be experimentally tested. It is also important to note that, in the field, it is readily distinguishable from A. haydenii ( Fig. 3A, B ). The two species can be distinguished using the following key:

1 Corolla blue, paling to nearly white when fresh; corolla lobes narrowly lanceolate, 3–6 mm long, 1.9–3.5 mm wide; free portion of filaments 0.2–1.5 mm (mean 0.9 mm); restricted to the Beautiful Mountain area and near Red Rock, Arizona ......................................................................................................................................................... A. cliffordii

- Corolla magenta when fresh; corolla lobes oval to oblanceolate, slightly wider than above, 3.5–9 mm long, 2–4.2 mm wide; free portion of filaments 0.8–2.7 mm (mean 1.5 mm); scattered in the Four Corners region of Colorado, New Mexico and Utah ................................................................................................................................. A. haydenii

Figures

FIGURE 2. A. Habit and architecture of Aliciella cliffordii, scale= 3.0 cm, J.M.Porter, K.D.Heil, & A.Clifford 13528 (Photo J.M.Porter). B. Cymose inflorescence and flowers, scale= 5.0 mm, J.M.Porter, K.D.Heil, & A.Clifford 13528 (Photo J.M.Porter). C. Bombilius sp. visiting flowers of Aliciella cliffordii, scale= 5.0 mm, J.M.Porter, K.D.Heil, & A.Clifford 13528 (Photo J.M.Porter).

FIGURE 3. A. Habit and architecture of Aliciella haydenii, scale= 3.0 cm, J.M.Porter & K.D.Heil 12455, (Photo J.M.Porter). B. Cymose inflorescence and flowers, scale= 3.0 mm, J.M.Porter & K.D.Heil 12455, (Photo J.M.Porter).