Navarretia modocensis L.A.Johnson & D.Gowen, 2017

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 : 53-56

Navarretia modocensis L.A.Johnson & D.Gowen sp. nov. Figs 4 , 5

Type.

United States of America. California: Shasta County, west side of Hwy 89, ca 10 miles north of junction with Hwy 299. Meadow area adjacent to railroad tracks, 41.0450°N, 121.6326°W, 2992 ft, 17 June 2004, L.A. Johnson 04-130 (holotype BRY! [BRY-614952]; isotypes JEPS! RSA! and to be distributed).

Diagnosis.

A species similar to Navarretia vividior , but distinguished by being diploid, rather than allotetraploid, generally being more conspicuously villous in the proximal inflorescence, possessing slightly larger flowers and more exserted corollas that tend toward pinkish-lavender lobes with darker throat above a yellowish tube rather than bluish to bluish-lavender lobes with darker throat above a yellowish or whitish tube, and having white (rarely blue) rather than blue pollen.

Description.

Taprooted annual herbs to 12(-15) cm tall and 20(-25) cm wide, sometimes larger, often wider than tall. Primary stem erect, terminating in an inflorescence head 1-2(-4) cm above the cotyledons; generally greatly exceeded by secondary stems, with tertiary, and quaternary stems present on larger plants; 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 reddish-brown, commonly glandular-villous, the trichomes generally greater than 0.5 mm (often ± 1mm) long, sometimes glabrescent; distal-most branches generally 0.3-0.5 mm in diameter. Cotyledons two, linear, entire, united at base. Leaves somewhat glandular-pubescent 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(-5) paired or unpaired linear lateral lobes 1-10 mm long attached along the proximal 3-5(-15) mm of the leaf, with an elongated, linear terminal segment. Inflorescences head-like, largest generally ≥ 12 mm diameter (exclusive of bract tips; ≥ 18 mm with bract tips), mostly 12-20-flowered, sometimes more, glandular-pubescent, generally with conspicuous, minutely-glandular villous trichomes proximally. Inflorescence bracts <15(-20) 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 minutely-glandular villous abaxially, often more densely villous adaxially and proximally along the lobes just above the bract base, glands becoming more prominent and their stipe diminishing in length toward the bract tips. Flowers actinomorphic, calyces mostly 5-7.5(-12) mm long, tube ~ (1.5-)2-2.5(-3) 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, glandular-puberulent on the intercostal membrane with the costae at least somewhat glandular-villous, the gland stipes longest along the costae at the junction with the intercostal membrane, diminishing in length toward the chlorophyllous costae tips (may be glabrescent with age); intercostal membrane v-shaped at sinus. Corolla generally equal to exceeding the calyx costae at anthesis and exceeding the calyx further as fruit matures, narrowly funnelform, glabrous, 6.0-8.2 mm long, lobes (1.0-)1.2-1.5(-1.95) mm long × 0.8-1.4 mm wide, tube white proximally, yellow distally, throat lavender-purple or purplish streaked, lobes light to dark pinkish-lavender; tube base expanding and investing the fruit apex. Stamen filaments unequal, 0.3-1.3 mm long, inserted unequally 0.3-1.6 mm below corolla sinuses, anthers ± included in throat to exserted less than half the length of the corolla lobes; pollen white (uncommonly blue), apertures pantoporate, acolpate; sexine seimitectate, reticulate, heterobrochate. Ovary three-chambered, stigmatic lobes three, included in to slightly exserted from corolla throat; capsule ~ 2.6-3.8 mm long, dehiscing circumcisally around the base with valves splitting upward. Seeds generally 4-9 per locule, medium brown, ovoid-angular, mucilaginous when wet. Nuclear gene loci showing diploid PCR amplification patterns.

Habitat, distribution, and phenology.

Navarretia modocensis occurs in volcanic influenced soils in forest openings and sagebrush slopes from (390-)800-1700 meters predominately in the Modoc Plateau of northeastern California and adjacent southern Oregon, but extending south to the western flank of the northern Sierra Nevada/southern Cascade Range in Butte County, California, and with a long-distance disjunct occurrence in San Benito County, California. Flowering occurs primarily in (April-)June-July.

Conservation status.

Navarretia modocensis has many occurrences throughout its range and is often abundant when encountered. It is a species of Least Concern following IUCN (2012) Red List version 3.1 criteria.

Etymology.

From the Latin - ensis, origin or place, combined with Modoc, in reference to the Modoc Plateau on which this taxon predominantly (but not exclusively) occurs.

Representative specimens examined

(paratypes). UNITED STATES OF AMERICA. California: Butte County, North east of Forest Ranch , 2.3 miles east and southeast of Hwy 32 along Garland Road, T 24N R3E S33, 2960 ft, 13 June 1980, Schlising & Azevedo 3699 (CHSC) and 20 May 2017, Johnson 17-027 (BRY); Top of North Table Mt , ca 200 meters north of junction of Cherokee Rd and east edge of Mt, T 20N R4E S17 NE1/4, 18 May 1979, Jokerst , Devine , & Greenstein 0462 (CHSC); Ponderosa way on Musty Buck Ridge between Cohasset and Big Chico Creek, T 24N R2E S25 SE NW 1/4, 2500 ft, 16 May 1989, Oswald 3773 (CHSC); About 0.5 mile east of Kunkle Creek , about 2.5 miles south-east of Paradise , 1500 ft, 29 May 1979, Ahart 1906 (CHSC); Lassen County , Off Hwy 139, up slope and across fence at a pull out area on west side of road, near mile marker 49.5, 40.9833°N, 120.7950°W, 1615 m, 9 July 2014, Johnson 14-176 (BRY); Murrer Meadow Rd, ca 3 miles east of Eagle Lake Biological Station , 5100 ft, 24 June 1967, Stern 2418 (CHSC); 100 yards south of Hwy 44, 34 miles northwest of Susanville , 26 June 1974, Stern 5863 (CHSC); Ash Valley RNA/ACEC, T37N R11E S5, 5100 ft, 29 June 1993, Schoolcraft 2303 (CAS; mixed collection with N. divaricata ); Near Bar Springs road, just off Hwy 299 near top of grade, ca. 7.4 miles east of county line, 41.0669°N, 121.2315°W, 1383 m, 14 July 2016, Johnson 16-075 (BRY); Modoc County , Vicinity of mile maker 9.37 off of Hwy 299, near Roney Flat and Johnson Creek Roads, 41.30602°N, 120.89584°W, 1428 m, 25 June 2011, Johnson & Smith 11-064 (BRY); Fandango Pass vicinity, helicopter pad 5.1 miles from Hwy 395 on Fandango Pass Road (Rd 9) at south end of Buck Creek Fire station and intersection with Jack’s Rd, 41.8639°N, 120.2849°W, 1586 m, 9 July 2014, Johnson 14-170 (BRY); About 0.3 miles up 41N25 (Shake Canyon) off of Hwy 139/299, 41.3825°N, 120.9437°W, 1415 m, 9 July 2014, Johnson 14-173 (BRY); 0.6 miles along Johnson Creek road from jct with Hwy 299/139 about 9.5 miles north of county line, 41.3120°N, 120.8874°W, 1479 m, 14 July 2016, Johnson 16-083 (BRY); 0.7 miles east of Hwy 139 along road FS 46, 41.5090°N, 120.9750°W, 1519 m, 14 July 2016, Johnson 16-086 (BRY); Little Hot Spring Valley , 15 June 1894, Baker s.n. (UC 23519); San Benito County , Condon Peak vicinity, 36.3205°N, 120.63817°W, 4683 ft, 5 June 2016, O’Dell s.n. (BRY; JEPS); Shasta County , Fuller Flat. north side of Hwy 299, just west of mile marker 64.71, about 14 miles west of junction with Hwy 89, 40.86532°N, 121.8238°W, 3792 ft, 17 June 2004, Johnson 04-124 (BRY, JEPS, RSA) and 14 July 2016, Johnson 16-072 (BRY); Meadow 0.3 miles past Rock Creek , south side of road 37, 41.0245°N, 121.7164°W, 3180 ft, 17 June 2004, Johnson 04-137 (BRY); Rock Creek vicinity; north of Summit Lake Road (Forest Route 38N10) ca. 3.5 miles from junction with Clark Creek Road, east of Lake Britton & Hwy 89, 41.0218°N, 121.7217°W, 973 m, 17 June 2005, Johnson & Zhang 05-157 (BRY); 0.4 miles south of Hwy 299 along Cassel road, 40.9561°N, 121.5772°W, 926 m, 14 July 2016, Johnson 16-073; Ahjumawi Lava Springs State Park , 3 miles north of McArthur , along Spatter Cone Trail, 3450 ft, 24 May 2004, Fischer & Marr 0213 (CHSC); Fall River Springs , 3500 ft, June 1903, Hall & Babcock 4213 (UC); Montgomery Creek , 27 June 1912, Eastwood 631 (CAS); Willow Creek Ranch about 6.8 miles south of the intersection of California State Rd 89 and Squaw Valley Rd in McCloud , 2900 ft, 29 May 1995, Almeda & Eisenhardt 7417 (CAS); Tehama County , High Cascade Range. Along Road 27N08 to Deer Creek and Onion Butte, ca. 4.6 miles from jct with Hwy 32 at sharp hairpin bend (ca. 0.9 miles from Deer Creek ), 40.1624°N, 121.5999°W, 994 m, 11 June 2015, Johnson 15-047 (BRY); North side of Hwy 36 in ecotone gradation between mixed conifer forest and chaparral at mile marker 75.49, about 12.8 miles west of jct with Hwy 89 to Mt. Lassen , 40.34734°N, 121.7299°W, 1138 m, 12 June 2015, Johnson 15-057 (BRY); Along Ponderosa Way at crossing of Soap Creek ;, 40.3933°N, 121.8033°W, 2450 ft, 30 May 1997, Taylor 16013 (BRY, JEPS); North side of Ishi Wilderness at Rancheria Creek Trailhead, 40.2052°N, 121.6141°W, 3210 ft, 31 May 2003, Hillaire & Elliott 327 (CHSC); 33 meters above Road 27N08, 2.3 km east of major hairpin turn, 40.1586°N, 121.6019°W, 1000 m, 30 May 2004, Janeway & Castro 8123 (CHSC); North facing slope along Powerline Road just southwest of north fork of Little Antelope Creek , 40.1833°N, 121.9186°W, 1890 ft, 25 April 2009, Castro & Janeway 1778 (CHSC) ; Oregon: Jackson County, Near trail, 19 June 1931, Howell 6766 (CAS); 7 miles southwest of Prospect , 2100 ft, 28 June 1939, Hitchcock and Martin 4992 (IDS; OSC); On Copco Rd. , about 1/ 2 mile south of Ranch , just north of California line, T41S R4E S10 SW1/4, 18 June 1990, Rolle 237 (OSC); Klamath County , 11 miles east of Blye , 30 June 1937, Peck 19655 (WILLU); Hwy 140, about 4.7 miles west of Lake County line, to south of highway near old dirt road, 12 July 1995, Johnson 95-053 (BRY); Lake County , 24 miles northwest of Lakeview , 25 June 1927, Peck 15226 (WILLU); jct of FS-3870 and highway 140, 42.2632°N, 120.7197°W, 1529 m, 29 June 2017, Johnson & Johnson 17-159 (BRY) .

Notes.

Navarretia modocensis are mephic when fresh. Three of the nine paratypes listed for N. divaricata var. vividior belong here: Baker s.n. [UC-23519], Hall & Babcock 4213 [UC-127856], and Austin 827 [UC-133706]. A collection from Yosemite National Park, Mason 12480a [UC-908393], likely belongs here also, representing a second disjunct population for this species. Pollen is usually white in N. modocensis , but blue pollen has been observed.

Navarretia modocensis is the species most likely to be visually confused with N. vividior . Though N. modocensis can have larger inflorescence heads, calyces, and flowers than N. vividior , the range of measurements in these features overlap. We are fairly confident (on fresh flowers) that N. modocensis has a yellow corolla tube on fresh flowers, and that populations with blue pollen are uncommon, but we are less certain that N. vividior always has yellow on its tube, or that populations with white pollen do not exist (given white results from the absence of pigment; we have collected species in several genera, including Navarretia that are characterized by blue pollen but occasionally have populations with white pollen). The more robust features of N. modocensis contrast more sharply with N. aeroides , which has smaller flowers, smaller inflorescence heads, thinner branches, and is visually much more glandular and less villous in its inflorescence heads. The smaller dimensions and very different corolla coloration patterns also readily distinguish N. divaricata and N. torreyella from N. modocensis .

Figures

Figure 4. Distribution of Navarretia vividior (circles), N. modocensis (squares), N. aeroides (triangles), and N. torreyella (4 - pointed stars) occurrences across their native range in California and southern Oregon, United States. Instances of observed syntopy are indicated with the hexagon and pentagon, as defined in the legend.

Figure 5. Navarretia modocensis. A Pressed specimen showing plant habit, scale bar = 1 cm (Johnson 04 - 130) B Flowering head in the field, scale bar = 2 mm (Johnson 16 - 072) C, D Pressed flowering heads showing range of typical coloration, equivalent magnification, scale bar = 2 mm (Johnson 04 - 124, 04 - 130, respectively) E-G Fresh flowers showing typical coloration and white pollen, equivalent magnification, scale bar = 2 mm. (E, F = Johnson 14 - 170; G = Johnson 16 - 0 72). All photographs by L. A. Johnson and vouchers deposited at BRY.