Navarretia divaricata Greene, Pittonia 1(8): 136. 1887 emend. L.A.Johnson & D.Gowen

Johnson, Leigh A. & Gowen, David, 2017, Ex uno, multis: taxonomic revision in Navarretia divaricata (Polemoniaceae) and the recognition of four additional cryptic or near-cryptic species, PhytoKeys 91, pp. 39-83 : 43-47

Navarretia divaricata Greene, Pittonia 1(8): 136. 1887 emend. L.A.Johnson & D.Gowen Figs 1 , 2

Gilia divaricata Torr. ex A.Gray, Proc. Amer. Acad. Arts 8: 270. 1870, non Gilia divaricata Nutt. 1848.

Navarretia divaricata Type: United States of America. California: Mariposa County, Yosemite trail, 1866, Bolander 4908 (lectotype, designated here: GH scan! [GH-00274983]; isolectotypes: UC scan! [UC-23512], YALE scan! [YU-065402], US scan! [US-322192], KEW, middle of three specimens, flanked on either side by Rattan s.n, scan! [K-000769074]). Replaced synonym.

Gilia atrata M.E.Jones, Contr. W. Botany 12: 55. 1908.

Navarretia divaricata Type: United States of America. Idaho: Washington County, Salmon Meadows, 4000 ft, 22 July 1899, Jones s.n. (holotype: POM, top three specimens on sheet, scan! [POM-75127]).

Navarretia prolifera Greene var. breviflora M.Peck, Proc. Biol. Soc. Washington 50(24): 94. 1937.

Navarretia divaricata Type: United States of America. Oregon: Douglas County, Diamond Lake, 3 July 1936, Peck 19234 (holotype: WILLU! [WILLU-18416]; isotype: OSC scan! [OSC-43592]).

Type.

Based on Gilia divaricata Torr. ex A.Gray, non Gilia divaricata Nutt. 1848.

Emended description.

Taprooted annual herbs to 15 cm tall and 20 cm wide, sometimes larger, often wider than tall. Primary stem erect, terminating in an inflorescence head 1-5 cm above the cotyledons; generally greatly exceeded by secondary stems, with tertiary, and quaternary stems present on larger plants (these higher order stems may be reduced in length and the inflorescence heads ± congested); higher order branches arise from axils of proximal inflorescence bracts, axils of leaves subtending the primary head, or less commonly, leaves within 1 cm of an inflorescence head; branches ascending to spreading and ± leafless, except for leaves subtending higher order branches or within 1 cm of a head; stem and branches tan to reddish-brown, glabrous or sparsely minutely glandular pubescent to glabrescent, less often villous, the trichomes generally less than 0.5 mm long. Cotyledons two, linear, entire, united at base. Leaves somewhat finely stipitate-glandular proximally, less so distally; leaves at the lowermost nodes opposite, linear-filiform, and widened at the point of stem attachment, the proximal nodes often congested with overlapping leaf bases. More distal leaves alternate, entire, or more commonly with 1-3 paired or unpaired linear lateral lobes 1-5 mm long attached along the proximal 3(-15) mm of the leaf, with an elongated, linear terminal segment. Inflorescences head-like, generally ≤ 10 mm diameter (exclusive of bract tips; ~15 mm with bract tips), mostly less than 15 flowered, villous proximally, obscurely glandular. Inflorescence bracts <10(-13) mm long, ± palmatifid to subpalmatifid; outermost 1-2 bracts with a short achlorophyllous base and 2-3 pairs of lateral lobes flanking an elongate terminal lobe, the distal pair of lateral lobes sometimes shorter and reflexed somewhat out of plane relative to the other lobes; bract bases become larger and clasping centripetally with lateral lobes reduced to a single pair departing from near the apex of the bract base flanking the central terminal lobe, all bract lobes chlorophyllous, entire, long tapering acute. Bracts sparsely villous abaxially, more densely villous adaxially and proximally along the lobes just above their point of attachment, with the distal 1/2 of each lobe more or less glabrous or with a few minute stipitate glands. Flowers actinomorphic, calyces mostly 4.5-7.5 mm long, tube ~ 1.5-2 mm; costae entire, long tapering acute, strongly unequal to subequal with typically two costae longer than the other three; costae narrowing proximally, the shorter ones narrower at base than the intercostal membrane and the longer ones subequal with the membrane; calyx tube achlorophyllous, minutely glandular-puberulent on the intercostal membrane with the costae at least somewhat villous, the trichomes longest along the costae at the junction with the intercostal membrane, the free portion of the costae glabrous to very sparsely and minutely glandular distally; intercostal membrane v-shaped at sinus. Corolla generally shorter than longest calyx costae at anthesis but exceeding the calyx as fruit matures, narrowly funnelform, glabrous, 3.5-5.0 mm long, lobes 0.6-1.0 mm long × 0.4-0.9 mm wide, proximal tube white, distal tube and throat yellow, sometimes red-streaked, transitioning to white lobes suffused with pink or lavender at tips (drying pink); tube base expanding and investing the fruit apex. Stamen filaments unequal, 0.1-0.6 mm long, inserted unequally to subequally 0.2-1.0 mm below corolla sinuses, included in throat to slightly exserted; pollen white, apertures pantoporate, acolpate; sexine seimitectate, reticulate, heterobrochate. Ovary two-chambered, unequally three-valved with two values bearing a septum and the third valve smaller and lacking a septum entirely or nearly so, stigma obscurely three-lobed, minute, unequally divided with two stigmatic lobes nearly entirely fused, generally included in the corolla throat; capsules mostly 2-2.5 mm long, dehiscing circumcisally around the base with valves splitting upward loculicidally (completely) and septicidally (partially). Seeds generally 5-9(-12) per fruit, medium brown, ovoid-angular, mucilaginous when wet. Nuclear gene loci showing diploid PCR amplification patterns.

Habitat, distribution, and phenology.

Navarretia divaricata occurs on a variety of soils from (800)1000-2600 meters in foothill and mountain habitats. It is widely distributed from Santa Barbara and Tulare Counties, California in the south to just within the borders of British Columbia, Canada, in the north, and east to Nevada and Idaho (Fig. 2 ). Flowering time is (May)June-July(September).

Conservation status.

Navarretia divaricata has many occurrences throughout its broad range, and is typically abundant when encountered. Occurrences near the periphery of its range (e.g., in British Columbia) may be limited in numbers, but the species is otherwise well established. It is a species of Least Concern following IUCN (2012) Red List version 3.1 criteria.

Etymology.

From the Latin divaricatus, to spread or diverge at a wide angle, which aptly describes the characteristic repeated pattern of nearly leafless stems diverging from under flowering heads to give rise to additional, well-separated flowering heads in this species and its morphological allies.

Representative specimens examined.

Canada. British Columbia: Anarchist Mt., 1.6 km north of Hwy 3 rest stop, 6.5 km due east of Osoyoos Lake , 49.0275°N, 119.35°W, 1350 m, 10 July 2000, Lomer 3838 (UBC Scan); Anarchist Mt. , 7 km due east of Osoyoos Lake , 1.5 km due southwest of summit, 200 m northeast of tower, 49.02658°N, 119.34786°W, 1380 m, 9 July 2013, Lomer 8435 (UBC scan) .

United States of America. All seventeen specimens cited by Jepson (1943) under Navarretia divaricata var. divaricata (JEPS); California: Alpine County, North side of Hwy 88 along an old dirt road, 38.7240°N, 119.9542°W, 2262 m, 10 July 2014, Johnson 14-179 (BRY); Amador County, down slope from Tragedy Springs Monument, 38.6389°N, 120.1465°W, 2406 m, 8 July 2014, Johnson 14-159 (BRY); Butte County , ca. 0.7 miles up Humboldt Rd to Butte Meadows from Hwy 32, 40.0407°N, 121.6056°W, 1204 m, 11 June 2015, Johnson 15-045 (BRY); Calaveras County , at entrance to old lumber road off of Hwy 4 ca. 1.6 miles northeast of Camp Connell Maintenance station , 38.3356°N, 120.2278°W, 1 July 1993, Johnson 93-112 (BRY); North side of Avery-Sheep Ranch Road ca. 2 miles from Hwy 4, 38.1969°N, 120.4003°W, 1098 m, 4 June 2014, Johnson et al. 14-068 (BRY); El Dorado County , ca. 0.1 mile west of NF-8N55 on Omo Ranch Road, 38.5542°N, 120.5401°W, 1172 m, 29 May 2013, Johnson 13-189 (BRY); off road to Ice House Reservoir, 6.4 miles from jct. with Hwy 50, 38.7975°N, 120.4028°W, 1596 m, 30 May 2013, Johnson 13-219 (BRY); Sly Park Vicinity, Park Creek Road 3.7 miles from county road E16, 38.7493°N, 120.4970°W, 1207 m, 8 July 2014, Johnson 14-141 (BRY); Lassen County , west of county road A21along dirt road leading to large moist swell, 40.3601°N, 121.0039°W, 1589 m, 2 August 2006, Johnson 06-129 (BRY); Nevada County, 39.32°N, 120.75°W, 1565 m, 3 June 2014, Johnson et al. 14-018 (BRY); Placer County , Off Sawtooth Ridge Rd ca. 1.1 mile west of Dawson Spring , 39.21787°N, 120.6226°W, 1670 m, 3 June 2014, Johnson et al. 14-029 (BRY); Plumus County , south of Cascade and Lava Top, access up FS road 21N22YA, 39.6827°N, 121.1665°W, 1402 m, 10 June 2015, Johnson & Ahart 15-024 (BRY); Tehama County , two miles south of Hwy 36/89 on Hwy 32, 40.2661°N, 121.4509°W, 4590 ft, Johnson 04-140 (BRY); Shasta County , off Hwy 89 ca. 1.3 miles north of jct with Hwy A 19, 41.1612°N, 121.6609°W, 1257 m, 25 June 2011, Johnson & Smith 11-058 (BRY); Trinity County , 10.3 miles north along Hastings Tie Road (FS-4N12), 40.5626°N, 123.5236°W, 1577 m, 27 June 2017, Johnson & Johnson 17-094 (BRY); Tulare County , Rabbit Meadows vicinity off Forest Service road 14S11 near 14S13A, T14S R29E S18, 2300 m, Johnson 94-065 (BRY); Tuolumne County , Twaine Harte vicinity, north side of highway 108, just west of hairpin in Lava Drive, 38.0266°N, 120.2369°W, 1159 m, 5 June 2014, Johnson et al. 14-077 (BRY); east of the Punch Bowl, north side of 4N12 ca. 5.4 miles east-northeast from junction with highway 108, 38.2386°N, 119.9496°W, 2178 m, 5 June 2014, Johnson et al. 14-108 (BRY); Idaho: Blaine County , two miles above Alturas Lake , above Alpine Creek , 7400 ft, 23 June 1941, Cronquist 2661 (IDS); Boise County , at the 62 mile marker of Highway 21, ca. 8 miles south of Lowman , 44.02°N, 115.62°W, 1859 m, 14 July 2003, Porter & Machen 13772 (BRY); Idaho County, Trail to Lake Serene , 45.1961°N 116.1907°W, 2169 m, 12 July 2013, Smith et al. 11162 (BRY, SRP scan); Valley County , FR 626, west of Sagehen Reservoir , Sagehen Basin, 44.3196°N, 116.1547°W, 1845 m, 17 July 2011, Smith 9922 (BRY, SRP scan); Brundage Mt. , 7000 ft, 23 July 1940, Davis 2930 (IDS); Nevada: Douglas County , Carson Range, Genoa Peak Road, 2560 meters, 13 August 1974, Williams 74-D-83 (RENO scan); Nevada, Washoe County , Carson Range, Little Valley , near bridge, 1981 meters, 22 June 1974, Tiehm s.n. (RENO scan); Oregon: Crook County , Little Hay Creek vicinity, along Forest Service Road 150 0.1 miles north of jct with forest service road 2610, T13S R19E S22, Johnson 97-133 (BRY); Harney County , Steens Mountain Loop Byway , 1 km below jct with Kiger Gorge turnoff, 42.69°N, 118.59°W, 8251 ft, 21 July 2007, Johanson 07-76 (WTU scan); Jackson County , Just north of Hwy 66 along Moon Prairie Rd, 42.1263°N, 122.3321°W, 1161 m, 29 June 2017, Johnson & Johnson 17-147 (BRY); Josephine County , Upper Biglow Lake , 10 July 1949, Baker & Ruhle 385 (IDS scan, WTU scan); Washington: Asotin County , Anatone Butte vicinity ca. 7.2 miles west of highway 129 on West Mountain Road , 1400 m, 2 July 1994, Johnson & Johnson 94-049 (BRY); ca. 0.5 mile west of Anatone Butte , 10 miles southwest of Anatone , T 7N R45E, S2, 4700 ft, 12 June 1949, Cronquist 5847 (ID scan, WTU scan WS); Chelan County , Rainbow Creek drainage, ca. 4.5 miles north of Stehekin River , 3000 ft, 19 June 1971, Naas & Naas 851 (WWB scan); Columbia County , Godman Spring campground, 18.5 miles southwest of Dayton , 46.10°N, 117.7861°W, 1753 m, 12 July 2004, Legler 2014 (WTU scan); Klickitat County , dry hills, Falcon Valley , 18 July 1908, Suksdorf 168 (WTU × 3 scan) .

Notes.

By elevating Navarretia divaricata subsp. vividior to species status, as done below, N. divaricata subsp. divaricata henceforth is designated simply N. divaricata . Fresh N. divaricata have no noticeable odor. The reported presence of this species in Montana is based on a single, misidentified specimen collected in an agricultural field; that specimen differs from N. divaricata in branching structure, having pinnately divided leaves, and possessing different bract and calyx morphology (our determination is N. squarrosa (Eschsch.) Hook. & Arn. [MONT-68910 scan!]. Though the protologue provides no mention of N. divaricata , Peck’s rationale for recognizing N. prolifera var. breviflora as distinct from N. divaricata can be inferred from the key and species descriptions in the first edition of his Manual of the Higher Plants of Oregon ( Peck 1941). Examination of the type reveals it is simply a narrow, more erect N. divaricata that fits well within the range of variation observed in this species. Peck, perhaps influenced by Mason (1951), came to realize this and removed N. prolifera var. breviflora from the second edition of his manual ( Peck 1961).

Navarretia divaricata differs from the remaining divaricately branched taxa of Navarretia (i.e., those treated below plus N. crystallina L.A. Johnson & D.Gowen, N. miwukensis D.Gowen & L.A.Johnson, and N. prolifera , in having two of its three stigmatic lobes nearly entirely fused, and in having unequal fruit valves with two of the three valves normal sized and bearing a septum, while the third valve is smaller and lacks a septum entirely or nearly so ( Cronquist et al. 1984; personal observation). These other species have three lobed stigmas and three equal valves, each bearing a septum. Navarretia divaricata 's flowers, ± 4 mm long, are among the smallest in Navarretia and typically dry with pink lobes with a white or yellowish throat and tube, though the throat and tube may at times be streaked with red. On herbarium specimens, the contrast between darker, pink lobes and light, whitish or yellowish tube is preserved on many flowers even on specimens 150 years old (Fig. 1 ). This coloration pattern enables this species to be determined without dissecting flowers to observe the stigma or fruit valves.

Figures

Figure 1. Navarretia divaricata. A Pressed specimen showing plant habit, scale bar = 1 cm (Johnson 14 - 143) B Flowering head in the field, scale bar = 2 mm (Johnson 15 - 045) C-E Equivalent magnification, scale bar = 2 mm C Pressed flowering head showing typical coloration of dried flowers (Johnson 16 - 024) D Dried fruit, showing dehiscence from base upward typical in all of the species detailed herein (Johnson 16 - 024) E Fresh flowering head dissected with corolla removed to show style and two-lobed stigma (with third lobe nearly entirely fused to one of the two apparent lobes; Johnson 16 - 024) F, G Fresh flowers showing typical coloration (Johnson 15 - 045), equivalent magnification, scale bar = 2 mm. All photographs by L. A. Johnson and vouchers deposited at BRY.

Figure 2. Distribution of Navarretia divaricata (black squares) occurrences across its native range in the western United States.