Navarretia paradoxiclara L.A.Johnson & D.Gowen, 2013

Johnson, Leigh A., Gowen, David & Jensen, Angela B., 2013, Cryptic speciation: distinguishing serpentine affiliated sister species Navarretia paradoxiclara and N. paradoxinota from N. intertexta (Polemoniaceae), Phytotaxa 91 (2), pp. 27-38 : 32-33

Navarretia paradoxiclara L.A.Johnson & D.Gowen , sp. nov. ( Fig. 3A, E, I, K, M–P ).

A species similar to Navarretia intertexta in gross morphology, but distinguished by having larger flowers with stamens shorter relative to the corolla lobes and predominantly occurring on serpentine derived soils. Similar and genetically closer to Navarretia paradoxinota , but distinguished by having notably larger corollas that are light blue (fading to white) rather than pure white.

TYPE: — U.S.A. California: Calaveras County, In moist drainage (serpentine nearby) leading to New Hogan Reservoir, within the Oak Knoll Campground, ca. 2.5 miles SE of Hwy 12 on S. Petersburg Rd. , 38.18049° N 120.79178° W, 208 m, 28 May 2009, Johnson, Gowen, & Mort 09-032 (holotype BRY!; isotypes JEPS! MO!, NY!, RSA!).

Taprooted annuals, erect with primary axis exceeding in length secondary axes, infrequently much (± 6) branched from the base with subequal secondary axes; primary axis to 30 cm tall; secondary axes 0–10(–20) in number, 0.2–5(–15) cm long, all axes terminating in an inflorescence; tertiary axes rare, and only from very well developed secondary axes. Stem and branches mostly tan to reddish brown, pubescent with white, retrorse eglandular hairs generally less than 1 mm long, sparsely pubescent at lowermost nodes, ± evenly pubescent distally. Cotyledons two, linear, entire. Lowermost nodes, ± 4 in number, bearing opposite leaves; leaves flexuous, entire to pinnate with filiform rachis and lobes, ± glabrous with blunt tips. Higher nodes alternate, leaves rigid, once to twice pinnate, tips acerose. Proximal nodes often congested, forming a rosette, leaves to 5 cm long, withering with age; internodes above the base commonly 2–3 cm long, associated leaves (subtending secondary axes, if present), diminishing upward from 4 cm to 2 cm in length, typically with 1–2 pairs of short linear lobes proximally, 2–3 pair of unequally branched lateral lobes, and the rachis then extending to a well separated, unequally three-pronged apex (the central prong or rachis terminus longer than the two lateral prongs), glabrous to sparsely pubescent with uniseriate hairs along the abaxial rachis and margins. Inflorescence 1–10(–20) in number, head-like, terminating all axes, ca. 8–26 mm wide exclusive of bract tips, up to 40 mm inclusive of bract tips, bearing up to 30 flowers in 1–3 flowered cymose units. Bracts strongly 3-dimensional from lobes diverging from rachis in multiple planes; outer bracts pinnate with a nongreen, flattened and linear proximal rachis ± 2 mm wide by 5 mm long, the lateral margin bearing 3–4 pairs of branched needle-like lobes; the proximal rachis transitions into a green, needle-like apex equal in texture/ appearance to the lateral lobes, three pronged distally with the central prong (rachis terminus) longer than the two lateral prongs; the abaxial rachis base and margins pubescent. Inner bract proximal rachis non-green, obdeltoid, concave-clasping with entire lateral margins such that all lobes are ± distal, flanking either side of the 3-pronged rachis apex; rachis base long pubescent abaxially and marginally. Flowers. Calyx (6.5–)8–11(– 12.5) mm, accrescent, of 5 costae connected into a tube 4–5 mm long by an intercostal hyaline membrane, this membrane v-shaped distally; the costae broadest at the distal tube, non-green proximally and tapering to a narrow base, narrower than the intercostal membrane, green distally, the lobes unequal with two typically longer than the other three, sometimes one or both of the longer lobes 3-pronged; calyx tube glabrous internally, pubescent and minutely glandular externally with uniseriate chain-like hairs increasing in length from the base upwards, longest near mouth of calyx tube, the lobes pubescent just above this region both adaxially and abaxially; minute glands short stalked, ± 50–60 µm in diameter. Corolla funnelform, light bluelavender or white (fading to white), exserted from calyx, 9.5–13.5 mm long, lobes rounded, 2.0– 2.9 mm long, 1.25–2.5 mm wide, with a single vein entering the base and branching pedately soon after. Stamen filaments 3.0– 4.7 mm long and inserted 1.2–1.85 mm below sinuses; the anthers 1–1.45 mm long, exserted to tips of corolla lobes or below. Pollen deep yellow; apertures pantoporate, acolpate; sexine semitectate, reticulate, heterobrochate. Ovary oblong, two-chambered; style 8.0– 11.5 mm long, stigma lobes two, 0.38–0.71 mm, Capsule 2.3–3(–3.8) mm long, proximal ¾ membranous, distal ¼ corneous, short apiculate, indehiscent at maturity. Seeds (4–)8(–10), medium brown (hue 5YR, value 3, chroma 4 to hue 2.5YR, value 3 chroma 4), ovoid-angular, 1–2.1 mm long × 0.8–1.1 mm wide, testa reticulate, mucilaginous when wet.

Habitat, Distribution, and Phenology:— Navarretia paradoxiclara occurs in serpentine influenced soils that are at least seasonally moist or best characterized as drainages, alone or among grasses. This taxon is presently known from at least 10 sites in Tuolumne and Calaveras counties, California and blooms primarily from mid May to late June (early July).

Etymology:— This specific epithet is derived from the Latin paradoxus —contrary to expectation, and clarus —bright, famous. Clarus is to honor Dr. Robert (Germanic for fame + bright) "Bob" Patterson, a faculty at San Francisco State University with a long interest in student training and in the systematics of Polemoniaceae . His selfless help has contributed meaningfully to the first author's research through the years. Paradoxus refers to the unexpected genetic similarity of this taxon to N. paradoxinota , and unexpected degree of molecular divergence from N. intertexta .

Additional specimens examined (paratypes):— U.S.A. California: Calaveras County, dry serpentine soil along a stream bank at the tip of New Hogan Reservoir in the N shore camp area, 2.5 mi S. E. of Hwy 12 near Valley Springs , 31 May 1972, McNeal 999 ( BRY, CPH, UT); Burson , 500 ft, 30 May 1923, Jepson 9948 ( JEPS); Salt Springs Valley , 1000 ft, Tracey 5650 ( JEPS); Upper Littlejohns Creek Drainage about 2.7 airmiles SW of Copperopolis , 1200 ft, 18 June 1984, Stone & Clifton 630 (& 633, JEPS) ; 5.8 miles west Altaville (just west of Elkhorn Station , along Calif. Highway 4) , 1420 ft, 6 June 1958, Bacigalupi, Constance & Alava 6375 ( JEPS); Dry serpentine soil in small gullies beside the Hwy 49 bridge over the N Fork of the Calaveras River, 16 May 1972, McNeal 887a ( BRY, CPH, UT); Serpentine drainage leading to New Hogan Reservoir , 38.17692°N, 120.78546°W, 206 m, 28 May 2009, Johnson, Gowen, & Mort 09-031 ( BRY, JEPS, RSA); Off Hwy 49, in a small channel in steep slope leading to N. Fork of the Calaveras river near bridge, serpentine soils , 38.21907°N, 120.69914°W, 227 m, 28 May 2009, Johnson, Gowen, & Mort 09-033 ( BRY, JEPS, RSA) . Tuolumne County, 3.6 miles west of Yosemite Junction, Highway 120, 24 May 1953, Cantelow s.n. ( CAS); Six-Big Gulch, upper portion, along Red Hills road, Red Hills , 1050 ft, 22 May 1984, Taylor 8516C ( JEPS); Chinese Camp , 1300 ft, 30 May 1915, Jepson 6328 ( JEPS); Dry rocky serpentine soil by a small stream crossing Sims Rd, 0.6 miles south of Hwy 120 between Yosemite Jct. and Chinese Camp, T1S R14E S5, 11 May 1972, McNeal 849 ( CPH, IDS); Just north of jct of State Hwy 59 and Hwy 108/120, ca. 3.9 miles southwest of Yosemite Jct. , seep area with serpentine soils , 37.84379°N, 120.5067°W, 324 m, 28 May 2009, Johnson, Gowen, & Mort 09-035 ( BRY, JEPS, RSA); Northwest side of Red Hills Road en route to Chinese Camp , serpentine , 37.84887°N, 120.45863°W, 313 m, 28 May 2009, Johnson, Gowen, & Mort 09- 039 ( BRY, JEPS); North side of Red Hills Road ca. 0.7 miles southwest of jct. of Hwys 120 and 49, serpentine influenced soils , 37.86542°N, 120.43976°W, 381 m, 28 May 2009, Johnson, Gowen, & Mort 09-040 ( BRY, JEPS, RSA) .

Notes:—We first became aware of this species when reviewing herbarium sheets from BRY in 2008 and made our first collections in 2009. Though suspecting this material was not typical N. intertexta , we were surprised to see the close genetic relationship between this material and the smaller flowered, allopatrically distributed material under study at that time that we here designate N. paradoxinota . The ecological propensity for both species to occur on serpentine derived soils, however, is congruent with the genetic relationships. The larger flowers of N. paradoxiclara compared to N. intertexta and N. paradoxinota are striking, and make this species showier both in the field and on herbarium sheets. This species was included in Johnson et al. (2012 b) as " Navarretia sp. nov. 2-CA."

Figures

FIGURE 3. Features of Navarretia paradoxiclara (all Johnson, Gowen & Mort 09-032) and N. paradoxinota (all Johnson, Gowen & Mort 09-021), with some comparison to N. intertexta and N. propinqua. All vouchers deposited at BRY unless otherwise indicated. A– D. flowers, top and side views, scale bar = 1 cm. A. N. paradoxiclara. B. N. paradoxinota. C. N. intertexta Gowen 1133, 1134-B. D. N. propinqua Johnson & Johnson 11-076. E–H. Corolla dissections, scale bar = 1 cm. E. N. paradoxiclara. F. N. paradoxinota. G. N. intertexta (left = Gowen 1133; right = Ahart 3453 [CAS]). H. N. propinqua Johnson & Johnson 09-067. I–J. Plant habit (note, either species can have a single leader (I) or be variously branched (J), scale bar = 1 cm. I. N. paradoxiclara. J. N. paradoxinota. K–L. Inflorescence, scale bars = 2 cm. K. N. paradoxiclara. L. N. paradoxinota. M–N. Outer inflorescence bract, N. paradoxiclara, scale bar = 1 cm. M. Adaxial view. N. Lateral view. O–P. Inner inflorescence bract, N. paradoxiclara, scale bar = 1 cm. O. Adaxial view. P. Lateral view. Q. Pollen grain, N. paradoxinota, scale bar = 10 µm. R. mature capsule, N. paradoxinota, scale bar = 1 mm (distal end to the left). S. Partially hydrated seed with thin halo of mucilaginous spiracles, N. paradoxinota, scale bar = 1 mm.