Navarretia vividior (Jeps. & V.L.Bailey) L.A.Johnson & D.Gowen, comb. et, 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 : 48-53

Navarretia vividior (Jeps. & V.L.Bailey) L.A.Johnson & D.Gowen, comb. et stat. nov. Figs 3 , 4

Navarretia divaricata var. vividior Jeps. & V.L.Bailey, Fl. Calif. 3(2): 156. 1943. Navarretia divaricata subsp. vividior (Jeps. & V.L.Bailey) H.Mason, Ill. Fl. Pacific States 3: 449. 1951.

Navarretia vividior Type: United States of America. California: Lake County, Mt. Hanna, 15 July 1897, Jepson 15045 (Holotype: JEPS! [JEPS-2659]). Basionym.

Type.

Based on Navarretia divaricata var. vividior Jeps. & V.L.Bailey.

Emended 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 10-25-flowered, glandular-pubescent with some 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(-4) 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 somewhat 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 4-7.5(-8.5) mm long, tube ~ 1.5-2.5 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 or shorter than longest calyx costae at anthesis but exceeding the calyx as fruit matures, narrowly funnelform, glabrous, 5.0-7.2 mm long, lobes 0.8-1.1(-1.4) mm long × 0.6-0.9(-1.2) mm wide, tube white proximally, distal tube white or yellow, throat bluish, sometimes streaked with magenta, lobes medium bluish-lavender; tube base expanding and investing the fruit apex. Stamen filaments unequal, 0.3-1.2 mm long, inserted unequal ly 0.3-1.3 mm below corolla sinuses, anthers ± included in throat to exserted less than half the length of the corolla lobes; pollen blue (white rarely?), apertures pantoporate, acolpate; sexine seimitectate, reticulate, heterobrochate. Ovary three-chambered, stigmatic lobes three, included in to slightly exserted from corolla throat; capsule ~ 2.3-3.3 mm long, dehiscing circumcisally around the base with valves splitting upward. Seeds generally 5-7(8) per locule, medium brown, ovoid-angular, mucilaginous when wet. Nuclear gene loci showing allotetraploid PCR amplification patterns.

Habitat, distribution, and phenology.

Navarretia vividior occurs in soils influenced by volcanic activity with favorable water status such as the edges of ephemeral pools and transient rivulets, to open flats or gentle slopes in forested areas from 300-1600 meters elevation. It occurs predominately in the North Coast Range from Lake and Sonoma Counties, California in the south to Humboldt, Trinity, and the western edge of Shasta County in the north (Fig. 4 ). A few populations cross the central valley and occur on the western flank of the northern Sierra Nevada/southern Cascade Range in Butte County, California, where they flower in late May-early June, rather than late June-July as is typical for species in the North Coast Range.

Conservation status.

As defined here, Navarretia vividior is distributed more narrowly than previously considered. Some historical populations have not been revisited for ± 100 years, though other known occurrences have been revisited multiple times over the past 20 years with no apparent change in local numbers (beyond what may be expected during drought years). Following IUCN (2012) Red List version 3.1 criteria, this species is most accurately characterized as data deficient, though it likely borders between being Vulnerable to a species of Least Concern.

Etymology.

This specific epithet is derived from the Latin vividus, lively or vigorous, with the comparative ending - ior, more so, in reference to the larger, more robust habit and flowering heads this taxon possesses in comparison to N. divaricata . Alternative epithets of var. Navarretia divaricata vividia and var. Navarretia divaricata fertilior were considered by Jepson and Bailey for this entity, as indicated by annotations on the type sheet.

Representative specimens examined.

UNITED STATES OF AMERICA. California: Butte County, North of Magalia at Old Coutelenc Site , along Coutelenc Rd , 1.7 miles south of junction with Skyway at Lovelock , 17 June 1980, Schlising & Banchero 3739 (CHSC); Upper Bidwell Park , Chico , on the north rim overlooking Sycamore Creek Canyon about 1.8 miles east of the power lines, 1280 ft, 30 May 1983, Oswald 578 (CHSC); South of Cohasset off of Cohasset Highway, T 23N R2W S8, 4 June 1979, Schlising 3435 (CHSC); Colusa County , Along Pacific Ridge Road (16N06) 0.5 miles east of road to Spanish Ridge and Hough Springs , 39.2236°N, 122.625°W, 3450 ft, 3 July 1998, Janeway & Isle 5711 (CHSC); Humboldt County , Waif on river bar, valley of Van Duzen River opposite Buck Mtn, 1000 ft, 27 June-30 July 1908, Tracy 2721 (UC); Little Van Duzen River , 23 June 1937, Eastwood & Howell 4813 (CAS); Lake County , Mt. St. Helena , 6 June 1915, Eastwood 4706; Bartlett Springs, 9 June 1938, Jepson 18938 (JEPS); Boggs Lake , at base on Mount Hanna , 29 June 1945, Mason 12629 (DS); Margin of Boggs Lake , 3000 ft, 29 June 1945, Baker 11088 (CAS); 3.5 miles by road east of Seigler Springs , margin of Snow's Lake , 2150 ft, 26 June 1956, Crampton 3646 (AHUC); About 1/ 4 mile down Harrington Flat Road from Bottle Rock Rd, where access road follows under powerlines, 2850 ft, 7 June 1981, Knight & Knight 4148 (CAS); Bogg's Lake Vicinity , between Harrington Flat Road and the Bogg's Lake parking area, to the north side of road along the power line corridor, 38.8904°N, 122.7837°W, 862 m, 13 May 2004, Johnson 04-032 (BRY), and 12 July 2016, Johnson 16-029 (BRY); Bartlett Mountain Ranger Station , 24 Sept 1949, Crampton 494B (AHUC, DAV); About 6.3 miles from Hwy 20 along Bartlett Spring Road, at junction with Pinnacle Rock Road, 39.1345°N, 122.7769°W, 3915 ft, 10 May 2005, Johnson 05-021 (BRY) and 12 July 2016, Johnson 16-032 (BRY); Mendocino County , Chamise Mt. north of Bell Springs , 3500 ft, 9 July 1934, Tracy 13326 (UC); Jeep trail above Bell Springs Rd 11 miles south of jct with Harris Rd, 39.9667°N, 123.6113°W, 1092 m, 28 June 2017, Johnson & Johnson 17-119 (BRY); Eden Valley Ranch , vernal pool area, north of Hearst , 1249.6 m, 11 July 1978, Wheeler 506 (CAS); Napa County , Gravelly slope in chaparral, edge of the Crater Country, 13 July 1930, Howell 5347 (CAS); Robert Louis Stevenson State Park ; along trail to Table Rock on East side of Hwy from parking area off Hwy 29, 38.6513°N, 122.5867°W, 802 m, 30 May 2009, Johnson & Gowen 09-049 (BRY); Shasta County , Harrison Gulch Rd, 2.4 miles north of Hwy 36 west of Platina , T 29N R10W S3, 3100 ft, 1 July 1996, Oswald, Ahart & Ondricek-Fallsheer 7906 (CHSC) and 13 July 2016, Johnson 16-062 (BRY); 0.3 miles north of Hwy 36 along NF-29, 40.3699°N, 122.9380°W, 893 m, 27 June 2017, Johnson & Johnson 17-052 (BRY); Sonoma County , Mayacmas Mtns, Caldwell Pines, ca. 3.5 miles west of Glenbrook and Bottle Rock Rd, 38.841°N, 122.815°W, 2350-3000 ft, 27 June 1975, Neilson & Sholars 3294b (DAV); Tehama County , West of Red Bluff and south of Platina on Forest Route 45 (Tedoc Mtn. Rd), 6.3 miles south of Hwy 36, 3800 ft, 12 July 1995, Oswald & Ahart 7022 (CHSC) and 13 July 2016, Johnson 16-058 (BRY); Trinity County , Along Wildwood Rd north of Hwy 36 between mile post 7 and 8, where drainage crosses and goes under road, 24 June 2005, Gowen 452 (BRY, JEPS) and 13 July 2016, Johnson 16-063b (BRY); 6.2 miles north of Hwy 36 along Hastings Tie Road, 40.5092°N, 123.5041°W, 1597 m, 27 June 2017, Johnson & Johnson 17-082 (BRY) .

Notes.

Navarretia vividior are mephic when fresh. Crampton 494B [AHUC-38375] (collected by McMillan at the Bartlett Mountain Ranger Station) is a voucher for a tetraploid chromosome count from pollen mother cells. Our low copy nuclear gene data indicate this species is an allotetraploid, with N. modocensis the paternal diploid parent and N. aeroides the maternal diploid parent. Four of the eight paratypes listed for N. divaricata var. Navarretia divaricata vividior belong here: Chestnut & Drew s.n. [UC-52826], Tracy 2721 [UC-133661], Tracy 13326 [JEPS-70300], and Jepson 18938 [JEPS-70299]; a fifth paratype, Condit s.n. [UC-455660], is difficult to place with confidence; it was collected from 'garden soil’ and thus its original source is uncertain, but its smaller corolla features suggest N. vividior . The Torrey 302 syntype of Gilia divaricata Torr. ex Gray also belongs here.

Navarretia vividior is readily distinguished from N. divaricata with its larger flowering head, larger flowers, equally divided 3-lobed stigma, typically blue pollen, and bluish-lavender corolla lobes and throat (Fig. 3 ). Stems are typically larger in diameter and invested with long, gland tipped hairs. The evenly 3-lobed stigmas, larger flowers, and some variation in flower color with colored lobes, throat, or both apply to three additional species that have heretofore generally fallen under the umbrella of N. divaricata subsp. vividior . These species are described and differentiated below.

Figures

Figure 3. Navarretia vividior. A Pressed specimen showing plant habit, scale bar = 1 cm (Johnson 16 - 066) B Flowering head in the field, scale bar = 2 mm (Johnson 16 - 063 b) C-F Equivalent magnification, scale bar = 2 mm C, D Pressed flowering head showing typical coloration of dried flowers and blue pollen (Johnson 16 - 029, 16 - 063 b, respectively) E, F Fresh flowers showing typical coloration (Johnson 16 - 063 b). All photographs by L. A. Johnson and vouchers deposited at BRY.

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.