Ipomopsis congesta (Hook.) V.E.Grant subsp. matthewii 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 : 21-24

Ipomopsis congesta (Hook.) V.E.Grant subsp. matthewii J.M.Porter , subsp. nov. ( Fig. 4 )

Similis Ipomopsis congesta (Hook.) V.E.Grant subsp. congesta , sed folia integra vel pinnatifida differt; similis subsp. crebrifolia (Nutt.) A.G.Day, sed folia basibus 12–22 mm differt; et similis subsp. frutescens (Rydb.) A.G.Day, sed caules 6–25 cm differt.

Type:— J.M. Porter , K.D.Heil, & A.Clifford 12673; U.S.A. New Mexico. San Juan County, Navajo Indian Reservation, “The Hogback,” overlooking Eagle Nest Arroyo , rimrock ( Point Lookout Member of Mesa Verde Sandstone Formation ), 8 May 2001. (holotype: RSA; isotypes: BRY, GH, RM, SJNM) .

Polycarpic perennials, bearing a branching, subterranean, woody caudex, sometimes taprooted when young, plants sometimes subshrubby at the base. Stems 1–20, decumbent, ascending or erect, simple or with up to 5 or more branches, 4–30 cm tall, with both glandular and eglandular trichomes, usually densely floccose. Leaves mostly cauline on each stem at flowering, 8–30 mm long, lower leaves linear and entire 0.8– 1.2 mm wide, upper leaves pinnatifid, with (2–)3–7 narrow segments, the terminal lobe, 0.7–1 mm wide, trichomes glandular and eglandular, more or less white villous or floccose, usually restricted to the adaxial veins. Inflorescence solitary or paniculately arranged heads, flowers in dense, capitate, cymose heads, 14– 25(–27) mm broad, flowers sessile or subsessile, inconspicuously bracteate, bracts 2.5–6(–10) mm long, simple, outermost trifid to 5-lobed, with 1–2 flowers subtended by a bract. Calyx usually 3.5–5.5 mm long, free portion of the lobes 1.0– 2.5 mm long, mucronate or shortly aristate, the margins hyaline, densely villous, the fused portion (tube) sparsely glandular puberulent, the green, herbaceous costae alternating with a narrow hyaline region. Corolla salverform, cream-white, 4.5–7.5 mm long; the tube 3.5–5(–6) mm long, only slightly, if at all surpassing the calyx, external corolla glabrous, rarely with a few glandular trichomes, internal tube with glandular trichomes (or papillae) along the more proximal staminal vasculature; lobes spreading, (1.5–) 2–3 mm long, elliptic-ovate to elliptic-oblong, rounded to acute at the apex. Filaments diverging from the corolla tube at the sinuses of the lobes, slightly curved, 0.8–2(–3) mm long, the anthers 0.5–1.5 mm long, exserted beyond the orifice of the corolla, pollen cream-white. Style reaching the anthers and slightly exserted, proximal portion sparsely glandular, the trichomes sometimes to 1 mm long. Ovary ovoid, bearing a few glandular trichomes at the apex, locules 3, 1 ovule per locule. Fruit a loculicidal capsule, 2.5–3.5 mm long, dehiscing from the apex, valves usually remaining fused at the base, with 1 seed per locule, often with some locules empty. Seeds 2–2.5 mm long, 0.6–1 mm wide, ellipsoid-fusiform, wingless, light brown to pale chestnut, funicular scar linear, elongate, surface irregular, becoming mucilaginous when wetted, embryo green, slightly curved, often visible within the seed.

Distribution:—Scattered across the Four Corners region, Ipomopsis congesta subsp. matthewii occurs in arid habitats, primarily associated with pinyon-juniper woodland. However, it may be found in sagebrush flats (often with scattered pinyon-juniper) or in desert scrub vegetation, where it may be found in badlands. Rarely, it has also been found in ponderosa pine-Gamble oak woodlands. Most frequently it is found in sandy pockets of sandstone outcrops, ledges, and ridge tops. It can also occur on sandy soils or clay soils, particularly if covered by alluvial or pediment gravels. Occurrences range in elev. from 1675–2440 m (5500–8000 ft).

Phenology:—Flowering begins in late April and generally continues through late June or early July. Exceptionally, flowering may continue into September. Fruiting begins in mid-May and lasts through August or rarely into September.

Etymology:—The specific epithet, matthewii, honors Matthew Heil, energetic field botanist of the Four Corners region, whose life was tragically cut short.

Additional specimens examined (paratypes):—U.S.A., ARIZONA. APACHE COUNTY: Navajo Nation, Alcove Canyon, 1/ 4 mi W of confluence with White Rock Wash, W Slopes of Carrizo Mountains , 27 April 1995, B.Hevron 2269 ( ASC) ; Navajo Reservation, Blackhorse Creek on the south side of the Carrizo Mountains , 30 May 2004, G.Rink 3354 ( ASC, NAVA) ; Navajo Nation, top of Mexican Cry Mesa, 14.1 mi from Cove , 27 Sept. 2001, K.Heil & A.Clifford 18383 ( SJNM, RSA) ; Navajo Nation, Lukachukai Mts. at Cove , 30 May 1998, S.L.O'Kane, Jr. 4370 ( NAVA, SJNM) ; Navajo Reservation, W foothills Lukachukai Mtns , ca. 9.5 mi E Round Rock Trading Post, T36N, R28E, S10, 6500 ft, 5 Jun 2001, A.Clifford & D. Atwood 01-610 ( SJNM) . COLORADO. ARCHULETA COUNTY: Arboles , 3 June 1899, C.F.Baker 534 ( POM) ; Sambrito Wetlands Area, hillside above Sambrito Cr. , 37° 0’ 23” N, 107° 28’ 43” W, 6125 ft, 8 Jul 2003, K.Heil, et al. 22120 ( SJNM) . LA PLATA COUNTY: 2 mi E of Ignacio , 6400 ft, 21 Jun 1964, J.N. Mann s.n. ( SJNM) ; Near La Boca Ranch , ca. 0.75 mi SE of railroad bridge, 650 ft, 18 June 1999, K.Heil 13242 ( SJNM) . MONTEZUMA COUNTY: Ute Mountain Indian Reservation, mesa top, T32N, R15W, S19, 23 June 1985, J.M. Porter 1301 ( SJNM) ; 7 mi. N. of Cortez. Limestone soil, 27 May 1982, R.Beal s.n. ( ASC) . MONTROSE COUNTY: 7 mi NE Nucla , T47N, R14W, 2 Sept. 1947, W.A. Weber 3578 ( RSA) . NEW MEXICO. SAN JUAN COUNTY: Navajo Nation, The Hogback, T 30N, R16W, S11, 4 May 1985, J.M. Porter 1044 ( SJNM) ; Navajo Nation , ca. 12 miles WSW of Sanostee, 11 June 2001, K.Heil 16676 ( NAVA, SJNM) ; ca. 2-3 mi N of Bloomfield , T29N, R10W, S3, 5650 ft, 11 May 1992, K.Heil & D. Hyder 7074 ( RSA, SJNM) ; 1 mi S of La Boca , Colorado, W side of Los Pinos River , T32N, R17W, S17, 16 May 1984, J.M. Porter 80-175 ( SJNM) ; Ute Mountain, Ute Indian Reservation , Barker Dome, T 32N, R14W, S10 SE qtr., 2000 m, 28 Apr 1989, K. Heil & J.M. Porter 4981 ( SJNM) . UTAH. SAN JUAN COUNTY: La Sal Mountains , 3 June 1914, M.E.Jones s.n. ( POM) ; U.S. Highway 666, 20.6 km E of Monticello , T34S, R26E, S7, 2070 m, 15 June 1993, N.H.Holmgren & P.K. Holmgren 11876 ( RSA, NY) ; along U.S. 666, at mile marker 14, ca. 3 mi E Eastland turnoff, 37° 49’ 56” N, 109° 05’ 47” W, 6793 ft, 30 May 2002, K.Heil & W. Mietty 19044 ( SJNM) ; 10.4 mi E of U.S. 191, on CR332 ( Hickman Flat Rd. ) ca. 3.5 mi N Pinte Knoll, 37° 56’ 57” N, 109° 09’ 17” W, 6775 ft, 22 May 2003, K.Heil & J.M. Porter 21813 ( SJNM) ; 11.6 mi E of U.S. 163, near Monticello, East Canyon , 37° 56’ 44” N, 109° 09’ 26” W, 6400 ft, 2 Jul 2002, K.Heil et al. 22018 ( SJNM) .

Discussion:— Ipomopsis congesta is a widespread species, occurring from the Pacific Northwest south to California, and east to South Dakota, Nebraska, Colorado, and New Mexico. Across this range, I. congesta displays considerable variability ( Constance & Rollins 1936, Cronquist 1984, Day 1980). Currently eight races are recognized ( Cronquist 1984, Day 1980, Welsh et al. 2003), either at the varietal or subspecific rank. These races differ in habit (from tall, wand-like and basally woody, to decumbent or cespitose herbs), leaf morphology (from linear or pinnatifid, to palmatifid leaves), vesture (glabrous to hairy or glandular), flower color (white to blue-purple), filament length (much shorter than the anther to twice the anther length), anther/ pollen color (white to blue, or yellow), ovule number, as well as ecological features. These races can be quite distinctive. As a result, some races are occasionally treated at the species rank, e.g., I. frutescens ( Rydberg 1913: 471) V.E. Grant (1956: 361) , I. crebrifolia ( Nuttall 1848: 11) Dorn in Clark & Dorn (1979: 41).

In the Four Corners region of Arizona, Colorado, New Mexico, and Utah a distinctive form of Ipomopsis congesta has been collected since the 1800’s. Various authors have considered it to represent I. c. subsp. congesta ( Weber and Wittmann 1996) , I. c. subsp. crebrifolia (Nutt.) A.G. Day (1980:112) ( Cronquist 1984), or I. c. subsp. frutescens (Rydb.) A.G. Day (1980:112) ( Weber and Wittmann 1996). In fact, the race found in the Four Corners region differs qualitatively (see below) from all three of the subspecies to which it has been assigned. We consider this taxon distinct and recognize it at the subspecific rank.

Ipomopsis congesta subsp. matthewii represents the southeastern Colorado Plateau race of the widespread I. congesta . As a whole, this species has received less attention in contrast to the I. spicata complex ( Wilken and Hartman 1991) or the I. aggregata complex (e.g., Grant and Wilken 1986; 1987; 1988a; 198b). The historic confusion of I. c. subsp. matthewii with other subspecies is understandable because it possesses traits reminiscent of I. c. subspp. congesta , crebrifolia , and frutescens ; however, it remains amply distinct morphologically and ecologically. Most similar to I. c. subsp. congesta , I. c. subsp. matthewii differs in generally having a more suffrutescent base, cream-white flowers and pollen, and displaying heteroblastic leaf variation (early leaf production of simple, entire, linear leaves, later production of pinnatifid leaves with long, linear segments). Ipomopsis congesta subsp. matthewii has also been confused with I. c. subsp. crebrifolia , a subspecies restricted to western Wyoming and adjacent Montana that is a woody, tufted subshrub with exclusively linear, short leaves. As previously noted, I. c. subsp. matthewii is often somewhat woody at the base and possesses at least some entire, linear leaves like I. c. subsp. frutescens . Unlike I. c. subsp. frutescens with its tall, wand-like stems, nearly all entire leaves, and two ovules per locule, I. c. subsp. matthewii has decumbent to erect, branching stems, basally entire but distally pinnatifid leaves, and one ovule per locule.

In spite of the new taxonomic recognition of Ipomopsis congesta subsp. matthewii , this subspecies is neither rare nor in need of specific conservation effort. It occurs over a wide area, from Montrose and Archuleta Counties, Colorado, to San Juan County, Utah, south to Navajo and Apache Counties , Arizona, and San Juan County , New Mexico. Even so, this is not a particularly common race, occurring in small populations, scattered across the region .

Infraspecific taxa of Ipomopsis congesta can be identified using the following key:

1 Leaves, or most of them, palmatifid to subpalmatifid, usually hairy (this complex of subspecies displays a near continuum of variation and could easily be considered a single subspecies; however, I maintain taxonomic recognition of these entities to emphasize the geographic restriction of subsp. nevadensis , and ecological differences of subsps. montana and palmifrons ) .............................................................................................................................................. 2

- Leaves entire, trifid, or pinnatifid, but not palmatifid .................................................................................................. 4

2 Filaments shorter than the anthers, anthers situated at the corolla tube orifice, flowers blue-lavender to white; apparently restricted to the Toiyabe Mountains, Nevada ........... I. congesta subsp. nevadensis (Tidestr.) Kartesz & Gandhi

- Filaments longer than the anthers, anthers situated beyond the corolla tube orifice, flowers white to lavender ......... 3

3 Most leaves crowded on short, vegetative shoots, forming a conspicuous basal mat; dwarf alpine or subalpine plants, seldom as much as 10 cm tall ...................................... I. congesta subsp. montana (A.Nels. & P.B.Kenn.) V.E.Grant

- Leaves mostly scattered along the stems, sometimes clustered as in the proceeding, plants to 20 or 30 cm tall, intermountain valleys to montane habitats ................................................ I. congesta subsp. palmifrons (Brand) A.G.Day

4 Leaves glabrous, rarely sparsely tomentose on abaxial surface, mostly crowded on short, vegetative shoots, forming a basal mat; stems usually simple above the branched base; middle and upper elevations of mountains.................... 5

- Leaves sparsely to densely hairy, rarely glabrous; sterile vegetative shoots rarely developed, stems often branched above; foothills, valleys, and plains.............................................................................................................................. 6

5 Leaves mostly entire; plants, up to about 12 cm tall ........................... I. congesta subsp. crebrifolia (Nutt.) A.G.Day

- Leaves mostly trifid, often some pinnatifid or entire, plants to 20 cm tall ..................................................................... ............................................................................................................. I. congesta subsp. viridis (Cronquist) A.G.Day

6 Cauline leaves mostly entire; plants 5–40 cm tall; herbaceous or woody above the caudex ....................................... 7

- Cauline leaves trifid to pinnatifid, entire only on proximal stems; plants rarely over 20(–30) cm tall, herbaceous above the caudex .......................................................................................................................................................... 8

7 Stems 10–40 cm tall, from a woody caudex and base, wand-like; Washington and Kane Counties, Utah, Coconino Co., Arizona ........................................................................................ I. congesta subsp. frutescens (Rydb.) A.G.Day

- Stems usually less than 15 cm, from a diffuse underground caudex, short and herbaceous; Green River Shale Formation of Duchesne and Uintah Counties, Utah ........................................................... I. congesta var. goodrichii Welsh

8 Basal leaves entire, 3–6 cm long, proximal leaves pinnatifid; W South Dakota, E Wyoming, E of the Continental Divide...................................................................... I. congesta subsp. pseudotypica (Constance & Rollins) A.G.Day .

- Basal leaves pinnatifid or entire, less than 3 cm long, widespread but W of the Continental Divide.......................... 9

9 Basal leaves pinnatifid; corolla white, anthers bright yellow; widespread but in northern portion of species range ... ........................................................................................................................................... I. congesta subsp. congesta

- Basal leaves entire, upper trifid to pinnatifid; corolla cream, anthers cream to white ............................................... 10

10 Stems woody well above the woody caudex; southern Colorado Plateau, widespread on sandy soils and pockets of slickrock sandstone ........................................................................................................... I. congesta subsp. matthewii

- Stems herbaceous above the woody caudex; a narrow endemic on hills of Arapien Shale Formation, Sevier County, Utah ......................................................................................................................... I. congesta var. ochroleuca Welsh

Figures

FIGURE 4. A. Habit and architecture of Ipomopsis congesta subsp. matthewii, scale= 2.0 cm, J.M.Porter, K.D.Heil, & A.Clifford 12673, (Photo J.M.Porter). B. Capitate inflorescence and flowers, scale= 2.0 mm, J.M.Porter, K.D.Heil, & A.Clifford 12673, (Photo J.M.Porter).