Navarretia torreyella 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 : 61-66

Navarretia torreyella L.A.Johnson & D.Gowen sp. nov. Figs 4 , 7

Type.

United States of America. California: El Dorado County, Peavine Ridge Road, along left fork (11N55) about 4.7 miles from junction with Icehouse Road , 38.7961°N, 120.4770°W, 1480 m, 26 June 2013, L.A.Johnson, R.L.Johnson, & A. Yankee 13-230 (holotype BRY! [BRY-619469]; isotypes JEPS! RSA! and to be distributed).

Diagnosis.

A species similar to Navarretia divaricata , but distinguished by having three equal stigmatic lobes and three fully developed fruit valves, and generally larger corollas with a deep maroon distal tube and throat abruptly transitioning to nearly white or less commonly pink lobes, the lobes drying lighter than the much darker throat and tube.

Description.

Taprooted annual herbs to 7(-10) cm tall and 14(-20) cm wide, sometimes larger, generally wider than tall. Primary stem erect, terminating in an inflorescence head 1-2(-3) 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, sparsely minutely glandular pubescent to glabrescent, less often villous, the trichomes generally less than 0.5 mm long; distal-most branches filiform, generally no more than 0.3 mm in diameter. 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 at tachment, 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(-12) 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 the rachis, with the distal 1/2 of each lobe more or less glabrous or with a few minute, stipitate glands. Flowers actinomorphic, calyces mostly 4.5-7.8 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 sparely, obscurely, and minutely glandular distally; intercostal membrane v-shaped at sinus. Corolla generally ± equal to the calyx costae at anthesis but exceeding the calyx as fruit matures, narrowly funnelform, glabrous, 4.0-6.0(-6.8) mm long, lobes 0.7-1.5 mm long × 0.5-1.0 mm wide, proximal tube white, distal tube and throat maroon, abruptly transitioning to white or whitish to uncommonly pink lobes; tube base expanding and investing the fruit apex. Stamen filaments unequal, 0.25-1.2 mm long, inserted unequally to subequally 0.2-1.2 mm below corolla sinuses, included in throat to exserted less than half the length of the corolla lobes; pollen white (uncommonly light blue), apertures pantoporate, acolpate; sexine seimitectate, reticulate, heterobrochate. Ovary three-chambered, stigmatic lobes three, included in to slightly exerted from the corolla throat; capsule ~ 2.5 mm long, dehiscing circumcisally around the base with valves splitting upward. Seeds generally 2-5 per locule, medium brown, ovoid-angular, mucilaginous when wet. Nuclear gene loci showing diploid PCR amplification patterns.

Habitat, distribution, and phenology.

Navarretia torreyella occurs on basalt flats, pyroclastic rubble, and clay soils from 1000-2100 meters elevation, in Butte, El Dorado, Nevada, Placer, Plumas, and Tehama Counties, California. Depending on latitude and elevation, it blooms from (May) June-July (September), beginning its flowering a little earlier than N. divaricata , and about the same as N. crystallina , when these taxa co-occur.

Conservation status.

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

Etymology.

In honor of John Torrey for the plant he recognized, before others, as distinct at the species level.

Representative specimens examined

(paratypes). UNITED STATES OF AMERICA. California: Butte County, about 8 miles east of Feather Falls, 1.5 miles east of Camp 18, yellow pine forest, 1000 meters, 4 June 1982, Ahart 3533 (CAS, CHSC); About 2 road miles west of Camp Eighteen along La Porte Rd (94), then 0.2 miles north along Frey Ranch Rd , 39.6291°N, 121.1970°W, 1266 m, 10 June 2015, Johnson & Ahart 15-022 (BRY, JEPS); About 1.2 miles east of Camp Eighteen along Lumpkin Ridge Rd, 39.6277°N, 121.1473°W, 1290 m, 10 June 2015, Johnson & Ahart 15-028 (BRY, JEPS, RSA); 1.6 miles north of La Porte Road (and 1.8 miles south of south end of Sly Creek Reservoir Dam), T20N R8E S19, 23 June 1980, Schlising, Ikeda, & Banchero 3765 (CHSC); Lumpkin Ridge, T 21N R7E S36, 20 May 1981, Schlising & Banchero 4059 (CHSC); Near Bull Hill Road, about 3 miles south of Butte Meadows , 1600 m, 16 August 1983, Ahart 4303 (CHSC); Jackass Flat, Mooreville Ridge, about 1 air mile northwest of the Lost Creek Reservoir Dam, 3800 ft, 20 June 1993, Ahart 6999 (CHSC); sides of a poor road, east of cow corral, about 200 yards north of the county road, about one mile west of the intersection of county road and Lumpkin Road, about 7 miles northeast of Feather Falls , 39.6294°N, 121.1983°W, 4204 ft, 2 July 2014, Ahart 19551 (CHSC); Southwest end of Lumpkin Ridge, about 6.75 air miles northeast of town of Feather Falls and 1.3 miles east of the Camp 18 site on topo maps, about 300 ft southeast of USFS Rd 22N27, 2 miles east of its junction with the county road and USFS Rd 22N94 to Fall River , T 21N R7E S36 SW1/4 of SE1/4, 4250 ft, 6 June 2005, Castro 1483 (CHSC); El Dorado County , off road to Ice House Reservoir, 6.4 miles from jct. with Hwy 50, 1596 m, 38.79754°N, 120.40287°W, 30 May 2013, Johnson 13-218 (BRY, JEPS); Along Park Creek Road, 2.8 miles from junction with Mormon Emigrant Trail, 15 May 2004, Johnson 04-076 (BRY); Sly Park Vicinity, Park Creek Road off of county road E16, 38.7493°N, 120.4970°W, 1207 m, 8 July 2014, Johnson 14-143 (BRY); Nevada County, Nevada City, 20-22 June 1912, Eastwood s.n. (CAS); Indian Springs Campground, near Cisco, 26 June 1965, Day 65-146a (DAV); 1.5 miles west of Cisco Grove , on a western slope above South Fork of Yuba River , 5800 ft, 21 July 1953, Crampton 1516 (AHUC; mixed collection with Navarretia divaricata ); Scotts Flat Reservoir on Deer Creek about 5 miles east of Nevada City, 3100 ft, 25 August 1965, True & J. T. Howell, 2548A (CAS); Near Grass Valley in red clay in the yellow pine and oak belt, 25 May 1919, Heller 13197 (CAS, UC); 2 miles west of Grass Valley , gravelly meadow edges, 2350 ft, 5 June 1939, Rose 39208 (CAS); hills northwest of Grass Valley , 2700 ft, 14 June 1967, Rose 67134 (BRY); Spur road off north side of Hwy 20, ca. 9-10 miles from I-80 at Yuba Pass, 1565 m, 39.3171°N, 120.7514°W, 3 June 2014, Johnson et al. 14-017 (BRY); West of meadow under power lines, about 150 yards west of paved road, southwest of Lake Spaulding , 39.3174°N, 120.6400°W, 5160 ft, 23 July 2005, Ahart 12185 (CHSC); Small meadow near a small wash between the curve and below the bridge on old highway 80, east of Donner Pass and Donner Summit, 39.6521°N, 120.3169°W, 6767 ft, 31 July 2005, Ahart 12227 (CHSC); North of Hwy 20 about 8.2 miles west of I-80 ramp at intersection with road 20-16, 39.3171°N, 120.7239°W, 1617 m, 11 July 2016, Johnson 16-015 (BRY); Placer County , Cisco, 25 June 1910, Hall 8712 (UC); Rainbow, north of Cisco , 5700 ft, 23 June 1946, Rose 46201 (UC); Strawberry flat, the Henderson ranch, near Indian Creek , T 15N R10E S23 SW1/4 of SW1/4, 3280 ft, 24 May 1969, Kawahara 870 (CAS); South facing slope of pyroclastic flow overlooking the North Fork of the American River , just off Sawtooth Ridge Road ca. 1.1 mile west of Dawson Spring , 39.2179°N, 120.6227°W, 1670 m, 3 June 2014, Johnson et al. 14-028 (BRY, JEPS); Along Forest Hill Road, ca. 0.2 miles northeast of NF-66 ( Humbug Canyon Rd. ), 39.1605°N, 120.6618°W, 1578 m, 3 June 2014, Johnson et al. 14-048 (BRY, JEPS); Off of Sugar Pine Road (NF10) enroute to Sugar Pine Reservoir, ca. 3.6 miles from Forest Hills Road, 1181 m, 39.1239°N, 120.7588°W, 9 June 2015, Johnson 15-013 (BRY); 50 yards northeast of the small natural lake, on the east side of the paved road to Lake Valley Reservoir , about 1/ 4 mile east of Yuba Gap and Highway 80, 39.3158°N, 120.6039°W, 5840 ft, 22 July 2002, Ahart 9896 (CHSC); Plumas County , south of Cascade and Lava Top, access via FS road 21N22YA, 39.6827°N, 121.1665°W, 1402 m, Johnson & Ahart 15-023 (BRY, JEPS, RSA); Along Lumpkin Ridge-La Porte Rd, ca. 6.9 miles northeast of junction with Golden Trout Crossing, 39.6982°N, 121.0783°W, 1541 m, 10 June 2015, Johnson & Ahart 15-030 (BRY, JEPS, RSA); About 1/ 4 mile north of Lumpkin Ridge Road , on Lumpkin Ridge, about 4 air miles northeast of Camp 18, 39.6856°N, 121.1061°W, 5013 ft, 3 July 2006, Ahart 12905 (CHSC, JEPS); North side of Onion Valley , east of Quincy La Porte Road , about 1 mile northwest of Pilot Peak , T 22N R10E S5 SE1/4, 6000 ft, 13 September 1995, Ahart 7640 (CHSC); About 100 yards north of the paved basalt road, about 3.75 air miles northwest of Tamarack Flat , about 5.75 air miles northwest of Little Grass Valley Reservoir , 39.7615°N, 121.0980°W, 5507 ft, 9 July 2006, Ahart 12929 (CHSC); West edge of top of Goat Mountain , 0.7 km southeast of the southeast side of Little Grass Valley Reservoir , 39.7181°N, 120.9664°W, 1721 m, 6 July 2011, Janeway 10497 (CHSC); Tehama County , across fence on southwest side of Hwy 36 about 5.1 miles west of jct with Hwy 32, 40.3278°N, 121.4735°W, 1459 m, 26 June 2017, Johnson & Johnson 17-042 (BRY) .

Notes.

Hall 8712 (UC) describes this collection as faintly malodorous though we have not detected a scent ourselves in this taxon. The Shelton s.n. syntype of Gilia divaricata Torr. ex Gray belongs here.

When growing with N. divaricata , a subtle difference in habit and coloration is discernable, with N. torreyella somewhat more spreading, its primary inflorescence head closer to the ground, and the plants overall more anthocyanic (purple tinged); however, as these features can vary in both taxa, they should not be relied upon for determination. Instead, corolla coloration of both fresh and dried flowers as outlined in the diagnosis distinguishes N. torreyella from all of the species treated herein. This taxon also has the thinnest, most filiform branches of the species considered here.

Key to the taxa formerly treated as Navarretia divaricata

1	Corollas 3.5-5 mm, lobes white or the tips tinged pink to lavender when fresh, drying pink (generally much darker than throat and tube), tube and lower throat yellowish when fresh, similar when dried (sometimes streaked with red); stigmas minute with 2 of 3 lobes fused almost to tips, fruit with 1 (of 3) valves half as wide and lacking a septum	N. divaricata
-	Corollas 4-8(+) mm, commonly with blue, lavender, pink, or whitish lobes and similar or darker maroon throat when fresh, drying with lobes and throat blue to purplish, or whitish lobes with reddish-streaked or dark maroon throat; stigmas equally 3-lobed; fruit equally 3-valved, each bearing a septum	2
2	Corollas bicolored when fresh with white or less commonly pink lobes abruptly transitioning to a dark maroon throat, drying in similar manner; distal half (or more) of inflorescence bract lobes and calyx costae glabrous or nearly so	N. torreyella
-	Corollas variously concolored to bi- or tri-colored when fresh with transition between lobes and throat gradual, bluish, bluish- or pinkish-lavender, or less commonly white, drying in like manner or with darker and/or reddish-streaked throat; distal half of inflorescence bract lobes and calyx costae generally glandular	3
3	Largest inflorescence heads exclusive of bract lobes ≤ 10 mm diameter (≤ 15 mm with bract lobes), conspicuously glandular proximally and distally (some villous trichomes present proximally); branches filiform ( ± 0.3 mm diameter), trichomes mostly <0.5 mm, sometimes wanting; corollas 4-6 mm, blue (generally with blue pollen) or whitish (with white pollen)	N. aeroides
-	Largest inflorescence heads exclusive of bract lobes mostly ≥ 12 mm diameter (≥ 18 mm with bract lobes), ± villous proximally, glandular distally; branches more robust ( ± 0.5 mm), trichomes commonly> 1 mm; corollas 5-8(+) mm, bluish-lavender or pinkish-lavender, pollen blue or white	4
4	Corolla generally 5-7 mm, bluish or bluish-lavender when fresh distally with whitish to yellowish tube, pollen blue; inflorescence heads often more glandular than villous; plants mainly west of the Central Valley in the North Coast and Klamath Ranges (uncommon in Butte County)	N. vividior
-	Corolla generally 6-8(+) mm, pinkish-lavender when fresh distally with yellowish tube, pollen white (uncommonly blue); inflorescence heads often more villous than glandular; plants mainly east of the Central Valley in the Modoc Plateau and Cascade Ranges, disjunct in San Benito County	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 7. Navarretia torreyella. A Pressed specimen showing plant habit, scale bar = 1 cm (Johnson et al, 13 - 230) B Flowering head in the field, scale bar = 2 mm (Johnson 16 - 008) C-E Equivalent magnification, scale bar = 2 mm. C Pressed flowering head showing coloration of dried flowers (Johnson 16 - 008) D, E Fresh flowers showing coloration (Johnson 13 - 218) F Fresh flowering head with maturing fruit, showing stretched corolla base that typically clings to fruit through maturity common in all of the species detailed herein, scale bar = 2 mm (Johnson 16 - 008). All photographs by L. A. Johnson and vouchers deposited at BRY.